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Evolution - The First Four Billion Years
Evolution - The First Four Billion Years
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with essayson the social and political sienifi-
cance of evolutionary biology today.
A second, encyclopedic section travels the
spectrum of topics in evolution with concise,
informative, and accessibleentries on individ-
uals from Aristotle and Linnaeus to Louis
Leakey and Jean Lamarck; from T. H. Huxley
and Edward O. Wilson to Joseph Felsenstein
and Motoo Kimura; and on subjects fronr al-
truism and an-rphibiansto evolutionary psy-
chology and Piltdown Man to the Scopestrial
and social Darwinism. Readers will find the
latest word on the history and philosophy of
evolution, the nuancesof the scienceitself, and
the intricate interplay anong evolutionary
study,religion, philosophy, and sociery.
Appearing at the beginning of the Darwin
Year of 2oo9-the zooth anniversary of the
birth of Charles Darwin and the rjoth an-
niversary of the publication of the Origirt o-f
Species-this volume is a fitting tribute to the
s c i e n c eD a r w i n s e t i n n l o t i o n .
M TcHAEL RUSE i s L u c y l eT . W e r k n r e i s t e r
Professor of Philosophy and Director of the Pro-
granr in the History and Philosophy of Science,
Florida State (Jniversiry.He is the founder and edi-
tor ofthejournal Biologyand Philosophyand has ap-
peared on "Quirks and Quarks" and the Discovery
Channel. His books include Tlrc Euolution-Crcdtiort
Struggleand Danyin and Desigtrftoth from Harvard).
J oS EPH T R A V T S i s D e a n o f t h e C o l l e g eo f
Arts and Sciencesand Robert O. Lawton Distin-
guished Professor of Biological Science at Florida
State Universiry.
www.hup.harvard. edu
-Edward O.Wilson
rsBN378-0-b7rr-03175-3
90000
llillfflilllilllllllllll
ilililil]tl
VoLUTloN
The First Four Billion Yers
EDlD BY
MlHAL RUsE
JosPH TRAVls
W|H A FoRWoRD BY
DWARD o. WlLsoN
H E E L K N A PP R s s o F
A R V A R DU N l V R s | YP R s s
A t ' 4 B R I D G
A. ssAHUss
L o N D o N , N G L A N D
2009
ontnts
olulr volutin
FrnisoJ.A
volution of th Gnr
Brin hrles
Human volution
Hnr . H
votutionyBilogy
ihl F' nto
Contnts
Adaptatin 105
Josph ruis nd Duid N. Rzik
olulavolution 12
Frnisol. l
0 Forword
ishi
48 durd. Wilson
t nd Pospt 70
401
Two nturis aftr its author's irh and 150 yars aftr its puli:rtion,
935 hrls Darwin's the rigin of Speis an fairly rnkd as th most
important book vr writtn. Not th most widly rad, to sur. opis of
949 th origi hav not bn plad in hotl rooms aross Amria; its vritis
ar not prahdfrom pulpits alr Sunday; and no politial ladr taks an
951
oath of offi with a hnd on its ovr. It is th mastri that first ad.
drssdth living world and (with he Descnt of following) human-
ity pl within it, without rfrnto any rligion or idology and upon
massivsintifi vidn providd aross sussiv dds. Its argumnts
hav grown ontinuously in stm as th st foundation for hran slf-
undrstndingand th pilosophial guid for human rion.
So solidly hav th filds of iology uilt upn th Darwinian onption
of volution that it maks sns tday to rogniz it as on of th two laws
(univsalprinipls if you wish) that govrn ou undrstanding of lif. Th
firs law is that all th lmnts and prosss tht dfin living organisms ar
ultimatly odint to th laws of physis and hmistry. This formulation
has bn th diving for of molular and llular iology during th past
half ntury' Th sod law, th foundation :rnd produt of volutionary
iology as wll s muh of organismi and nvironmntal iology, is that all
lmnts and prosss dfining living organisms hav n gnratd y
volutionthrugh natural sltiln.
Th two laws hav provn to frrlly omplmntary, a prim rquirmnr
for th rognition of sintifi laws gnrally. Furthr' to an inrasing d-
grthy ar ing omind to ahiv samlssanalyss of partiular i-
logial phnomna. Th first law addsss how a phnonrnon ours and
th sond addrssswhy it ours. Mitohondria and othr organlls, for
xampl, ar built and work thus and so; and thy originatd in on way and
not som othr. ah dsription omplts th xplanation for th othr.
odrn biology as a whol has littl maning without th joining of both ap-
proahs guidd y th two lws.
vn without th imptus of th origin ssquintnnial-fortuitously th
samas th Drwin irth intnnil-2009 is an pproprit tim to prsnt
u
ultx Foreord
volutionary iology in its nompassing modrn form. I dout that any singl
writr, or any nsml of sveral writrs, ould summariz this urgoning
disiplin in a omprhnsiv and authoritativ form. Th multipliity of au-
thors in uolutio: The First For Billio Yers has gon far to*ard that goal.
Its array of prts provid stat of th art for ah sujt in turn. Th sin
is prsntd hr' ut also som of th main pratial onsquns and in.
vitably, with fundamntal rlvan, th onsquns of iologial volu- Introdu
tionary thought for philosophy and rligion. Th grat qustions-..Who ar
w?'' ...Whrdid w om from?'' and ...Whyar w hgrg}''-gan nswrd
only, if vr, in th light of sintifially asd volutionary thought.
NIihlR
h disovry of volr
.W.strn
thought, ranl
mong sintifi dis<
From sing natur aI
forvr hanging, w
drstanding volution
In sin th word
idas. First, thr is tl
ganisms, living and dt
natural pross of h
othr, diffrnt ons. l
of th anitlt Grks.
ganisms volvd rally
sidrd a asi elm
pulishd is on tbe
organisms on th plar
that opratd rlntls
th natral world. An
tology to moleular i
vidn that volutiol
Sond, uolution is
glo. This an rfr t
volution of orhids,
suh as th volution <
reptil iaw. volution:
vidn that rangs fi
to similaritis and diff
tin of som faturs.
stood, whil th volu
organlls of lls, is l
h third ida is th
that volution has not <
ldrn form. I dout that any singl
ould summaizthis urgoning
tiv form. Th multipliity of au-
,rs has gon far toward that goal.
lr ah sujtin turn. Th sin
in pratial onsqunsand in-
:onsqunsof iologial volu-
Introdution
. Th grat qustions-cGWho ar
Y arewe h?''_an answrd ihl Rse d JosphTruis
d volutionarythought.
ronstrutd ut to orrr idas aout th fors that driv volutionar bhavior. Thr ar f
hang.In the rigi Darwin propsed what today is almost univrsallyon- whih rval that ds1
sidrd th major for in organi hang, natural sltion.But natural s- ovrd. Not that th.
ltion is not th only importnt volutionary for; natural sltion oprats rathr' thr is muh
on th variation in shaps, sizs, and iorrs of organisms atd y muta- spis or faturs' pa
tions in thir gns' and th rt and mirgnitud of th hangs that ar la lvls' and to rfinr
drivn y natural sltinare govndy hw thos gnsontrol dvlop- stonishing divrsity
ment. An enormous amount of osrvation and eprimnt has doumntd ook show ot only h
th ation of naturl sltionand som of th gnti rsPonssto sltion. li and how w might
But modrn disovrisin gntisand dvlopmnt have raisd many nw This ook is organi
qustions and guidd us toward sorsurprising answrs out how all ths on rh varhing tt
fors omin to driv volution. thought to th ontro
Outsid sin th word uolution rfers to a natural pross onsidrd origin of lif to th p
the inspiration for host of soiologial, litrary, politial' philosophial, m a j r f a t u r so f v o
and rligious ids. From writs xploring th animal natur of humans to ras lik molular .
psyhlogistssarhirrgfor th origirrof humn havior in volutionary his. Th sond pat onti
tory to thologians grappling with th impliations of volution for ou un- topis. Ths inlud t
drstanding of th Divin, th influn of volution outsid sin f p o p l a f m i l i a r ,t
ds that f any othr sintifi disovry vr mad. And as modrn on- urs in th disovy
trovrsis in th shools illustrat, partiularl in th Unitd Stats' it pro. oks that rount tht
voks rtiolls lmost s stong as issussurrounding th eginningand nd a disiplin'
of hnran lif. This organizat\on a
In this ook w xplor all ths fts of volution. Our uthors prsnt or hr intstsand ;
th vidn for volution as fat from th fossil rord to gnomis, illus- th sin an fous <
trat th histry f groups from atri to irds, and dsri our urrnt intrstd in th intlI
idas about how hs historis and tlris vidn am to b. Our uthors sin an ntr throl
also xplor tlr influn of voltrtion on plrilosopy, rligion, soiology, spifi topis or hist
psyhology, and many othr aras and addrss th urrnt ontovrsis with ssays on rlatd
aout th tahingof volution in shools.Th ook ovrs th historial dis- plling introdution to
overis, suh as homology, that opnd th path for disovring volution; point th way towad
th phnomena' suh s industrial mlanism, that playd maior rols in in- biliogaphisw , hih
spiring primntal studis of volution; th appliations of volutionary Although w hop t
prinipls to aras smingly far fild, suh as omputr sin;and th volum to inform,
ontributions f th major figurswho shaped th history <lfvlutionary si- sholars. W find vol
n and th disiplin as it xists today. intrsting, and ind
Prhas most important' this book prsnts voltionary sin as a mod- mrg with th sam
rn, dynami disiplin. All too oftn disussions of volution outsid th si- t i o n , i t i s i m p o t a n tt .
n itslf giv th imprssion tht th subjt am fssilizd in th hallng of antiiot
nintnthntuy' This is partiularly so whn th suitaility of volution as tious disass,it shor
a sujt for sondary-shoolstudntsis ing argud.Th irony is tht th a l t r a t i n so f h a i r a t
disovriesof modrn sin,from molular iology to otl1putationalin- planations of who '
novation' hav providd onrplling vidn that Darwin ould only dram m a n y o f o u r l o s l yh
of sing. From thos disovrishav also om appliations that Darwin h last of ths r
might nvr hav imagind, suh as Darwinian mdiin. Prinipls of volu- Provn so ontrovrs
tion ar ing applid to a wid vrity of sintifi prolems, from undr- popl in th Unitd S
stnding snsn to stratgis for onsrvation to plaining hunran of lif on arth. lt is u
ltroductio i
th lrs tht div volutionary havior. Thr ar fw topis in sin with so many xiting nw fats'
hat today is almost univrsallyon- whih rval that dspit all w hav larnd, thr is muh mor to dis-
:, natural sltion'But natural s- ovrd. Not that thr is futhr ned to sustantiat volution as fat;
ar for;ntulsltionopats rathr' thr is muh mor to don to tra th volution of groups of
ms of organisms rated y muta- spisor faturs, partiularly omplx faturs t th llular and molu-
ragnitud of th lrngs that ar lar lvls, and to rfin our thory of volution to aont mor fully for th
y how thosgnsontrol dvlop- astonishing divrsity of lif as w ontinu to disovr it. Th ssays in this
ln and primnthas doumntd book show not onl how far w hav om but whr th sintifi horizons
f the gntirsponssto sltion. li and how w might mov toward thos horizons.
vlopmnt hav raiso many nw This ook is organizd into two parts. Th first part ontains long ssays
lrising answrsabotrthow all ths on th ovrarhing thms in volution. From th history of volutionary
thought to th <rntrovrsisovr duation, from th fossil rord and th
:rs to naturl Pross onsidrd origin of lif to th pross of adaptation, th long ssays offr primrs on th
l, litrary, politial, philosophial, majo faturs of volution. Many of thm off los looks at partiular
lg th animal natur of humans to aras lik molular volution' gnomi volution' and Darwinian mdiin.
mn bhavior in volutionary his- Th sond part ontains a larg numbr of shortr ssays on mor spifi
pliationsof volution for our un- topis.Ths inlud ssayson malor groups of organisms with whih most
f volution outsid sin far popl ar familiar' topis that a important fats of volution, major fig-
ry vr mad. And as modrn on- us in th disovry and shaping of volutionary sin' and th ritial
r l a r l y i n t h U n i t d S t a t s 'i t r o - ooks that rount th history of disovry, dvlopmnt, and maturation of
;urroundingth ginning and nd a disiplin.
This organization allows th adr to xplor volution arding to his
of volution.our authors prsnt or hr intrsts and bakground. h radr who wishes to b immrsd in
r fossil rord to gnomis, illus- th sin an fous on th mjor sintifi thms, whil th adr who is
to irds, and dsriOur urrnt intrstdin th intlltual history and influn of th sujt away from
:vidnam to . our authors sinan ntr through a sparat st of ssays. A radr intrstd in vry
n philosophy, rligion, soiology, spifi topis o historial figurs an find th appropriat ntris, along
addrss th urrn ontrovrsis with ssays on rlatd topis. Ou goal was to provid an iting and om-
Th ook ovsth historial dis- plling introdution to volution along with a basi rfernwork that ould
h path for disovring volution; point th way toward a dpr study of individual issus. Th ssays inlud
sm' that playd major rols in in- ibliographis' whih srv as guids to furthr and dpr rading.
th appliations of volutionary Although w hop that our fllow sholars njoy ths ssays' W want this
rh as omputr sin; nd th volum to infom, duat' and it radrs who ar not profssional
ld th history of volutionarysi- sholars.\w find voltion in all of th word's manings to a provoativ,
intersting,and indd aw-inspiring topi' and we want our radrs to
nts volutionarysieneas a mod- mrgwith th sam fling. But yond th itemnt of studying volu-
ssionsof volution outsid th si- tion, it is important to undrstand and appriat it. voltion plains th
subjt am fossilizd in th hallng of antibioti.rsistant pathogns and th sourg of novl inf-
lhn th suitailityof volution as tious disass, it shows us wht W an xpt limt hang and our own
ringargud.Th irny is tht th altrationsof haitt to do to th natural world, and it may offr profound
ular iology to omputational in- xplanations of who w ar and why we hav as w do. It also hallngs
that Darwin ould only dram many of our losly hld lifs about man's, and woman's, pla in natur.
c om appliations that Darwin Th last of ths issus is prhaps th fondation of why volution has
Lianmdiin.Prinipls of volu- provn so ontrovrsial. Survy aftr survy shows that fwr than 50% of
.sintifiprolms,
from und- poplin th Unitd Statsapt that volution is rsponsil for th divrsity
]srvation to plaining human of lif on arth. It is unlar how muh of that opposition is asd on a poor
Introdution
Th arly Days
Th arly Days
inlud nils formd in a rudimntay Way in thir dsign for hu- humn thought and
mnkind, right at th start. This was thir raso, thn, and ths th pogrss.
profssd aims that guidd thm in making skin, hair and nails grow at Th British physii;
th xtrmitis of our |ims. (Timus, 76d_e, in oopr 1997, 1277) harls) Was a para
mankind:
In an inrdily influntial disssion, Aristotl in D prtibs nimlium
Imp:
(1,984\identifid th fators at work hre as ..final auss.'' Ths ar auss
Of la
that our not just to produ or do somthing (th fingr parts dry and mak With
ails) ut fo the sak of som kind of purpos (th nails prott th fingr And r
nds). Thy show som kind of forethought or intntion. For this rason, fi.
Arost
nal auss annot rdued to lind, unguidd law, as is dmandd in vo- An r
lution. Th world, partiularly th world of orgnisms' must in som sns
hav n dsignd rathr than just produd undr its own stam tly natual (.D
prosss.(Sdly2008 is th dfinitivstudy.)
This is all rh nd pr.
Th Jws, and following thm th arly Christians, had religious rasons
gn' whih auss
for th rjtion of volution. It gos against th ration storis of th arly
haptrs of Gnsis, whih portray a world ratd miulously y God and Hw lo
thn popld y him through divin fiat ovr a short tim span. But do not From .
think that rligion as suh was thn and always an asolut ar to volution- ow fr.
ism. Th hurh fathrs (th major hristian thologians of th arly n- h pilt
turis) workd toward an undrstanding f th ilial txt tht wuld allow
intprtation, partiularly in th fa of advans of sin. Sint Augustin
was agr not to lt anint ration aounts stand in th way of modrn Similar idas wr t
thought. H himslf, living that God stands outsid tim, spulatd in a Pbilosophi zoologiq
kind of protovolutionary fashion that th Divin had fomd sds of lif produd th first ful
that thn sprang into full ing whn thy wr plad hr on arth. How- to our own spis fr
vr' on should not ad too muh into any of this. Lik th Grks, th n produd from l
Jws and Christians wr simply not looking in th dirtion of volution and ity and othr natural
would hav thought final auss an unnswrabl oition to signifiant d- Although th mta1
vlopmntalism. As is wll known, ths kinds of auss am a founda' ris of volutionary i<
tion of on of th major proofs of God's istn, th Argumnt from vidn. Aristotl ha
Dsign, whih movs frm dsign her on arth to th xistn of th divin sha ommon pattrl
artifir. and th volutionists
.!hy,
thn, did volution start its rise in the ightnthentury? Th an. tris (Figur 1). Likr
swr is simpl. It was at this time that popl startd to hallng th 8o unnotid (Figur
Christin pitur of world histry-a providntial pitu of a world r. sition. Th fossil r
atd y God, whr humans ar mad in his imag ut hav falln and ar would invok first as
abl to ahiv salvation only through his undsrvd gra. Som gan ful. As a systmati p
to argu that prhaps humans hld thir fats in thir own hnds and were magr inded.
ould progrssivly improv thir own lots. It Was this ida of pOgrss- fat that no on hal l
th lif that th world and its dnizns ar on a trajtory upward nd ltion. ost assm
that this upward ris is mad possil y (and only y) th unaidd fforts fors, ut littl mor
of th world's human inhabitants-that gav ris to th ida of organi that haratristisa
volution (Rus 1996). nthusiasts for progrss tndd thi thinking ately to th futur g
into natur and dvlopd th id of volution-progrssiv hang up- that th inhritan
ward from th simpl to th ompl. They thn rad this ida ak into Lmrkism*ut bt
Th Histor fuolutior Thougbt 3
f wa in thir dsign for hu- human thought and soial prati as onfirmation of thir lifs about
thir rason, thn' and ths th progrss.
king skin, hair and nails grow at Th British physiian and man of sin rasmus Darwin (grandfathr of
76d-r, in Coopr 1997,|277) harls) was a paradigm who hymnd in vrs lif's upward ris to hu-
mankind:
'ristotl i D prtibs imlium
Imprious man, who ruls th stil rowd,
as ..final auses.''Ths ar auss
of languag'rason' arrd rfltionproud,
:hing (th firrgrparts dry and mak .With
brow rt who sorns this arthy sod,
urpos (th nails prott th finger
And stlshimslf th imag of his God;
;ht or intntion. For this rason, fi-
Aros from rudimnts of form and Sns,
guidd law, as is dmandd in vo-
An mryon point' or mirosopi ns!
of organisms)must in som sns
d undr its own stam by natural (. Darwin 180' 1: anto I,lins 09-14)
udv.)
This is all th nd produt of th progrssivdvlopmnt of human intlli-
y hristians,had rliglous rasons
gn,whih aussarrd is rfltdin humans' sintifiahivmnts:
nst th ration storis of th arly
d atdmiraulously y God and How lovs nd tasts,and svmpathisommn
vr short tim spn. But do ot From vansntnotis of sns;
lwys an absolut ar to volution- H<lw from yilding touh and rolling ys
stian thologians of th arly n- Th pils immns of humn sinris!
'f th ilial txt that would allow
(anto III,lins 4346\
dvans of sin.Saint Augustin
ounts stand in th way of modrn Similar idas wr to found lswh,most notaly in Fran. In his
tands outsid tim, spulatd in a Philosophie zoologiqu (1809), th taonomist Jan.Baptist d Lamark
r Divin had formd sds of lif produd th first full-blown volutionay thory-a pitur of upward ris
F w plad hr on ath. How. to ou own spisfrom th most primitiv forms of lif, whih in turn had
, any of this. Lik th Grks, th nprodud from mud and slim through th ations of hat and ltri-
ng in th dirtion of volution and ity and othr natural fors.
wralojtion to signifiant d- Aithough th mtaphysial ida of progss was the main fator bhind th
kinds of auss am a founda- ris of volutionary idas, it is not tru that thr Was no prtinnt mpirial
j's istn, th Argumnt vidn.Aristotl had notd that organisms of vry diffrnt spissm to
from
arth to th istn of th divin sharommon pattrns or struturs-wht today ar known as iromologis-
and th volutionists wr rady to intrprt ths as signs of ommon ans-
n th ightnthntury? T an- tris (Figur 1). Likwis, th susssof animal nd plnt brdrsdid not
: popl statd to hallng th go unnotid(Figur2). But gnrallyth vidntook vry sondary po-
ovidntial pitur of a world r- sition. Th fossil rord, somthing that today many (if not most) popl
his img ut hav falin and are would invok first as th proof of dvlopnrntalorigins, was lss than hlp-
is undsrvdgra. Som gan ful. As a systmati prof of progssiv hang, th glanings from th roks
ir fats in thir own hands and Wmagrindd. In any s, ounting against th mpirial sid was th
)ts.It was this ida of progrss- fat tht no on had any grat undrstandingf wht might hav ausd vo-
i ar on a trajetory upward and lution. ost assumd som kind of vagu' upwardly thrusting for or
(and only y) th unaidd fforts fors, but littl mor. Gnrally, vryon was ommittd to th folk blif
gv ris to th ida of organi that haratristisquird irr on gnration ould transmittd imrndi-
progrss xtndd thir thinking atly to th futur gnrations-Lamark was so nthusd y this pross
.olution-pogrssiv hng up- that th inhritrrnof aquird haratristishas sin om known as
1y thn rad this ida ak into Lamarkism-ut yond this was siln.
4 The istor of uolutior Thought
A DARWlnl
tl8D li llr
l, '!? or
Th idology of progrss was wht ountd, and it was for this rason that
most popl around 1800 would hav rgardd volutionism lss as a ral
sin and mo as a ptnd,smwhat lik animal magntism (ms- divin industialist a
mrism) and th rading of haratr from skull shap (phrnology). vn of th world througt
!udgd by th standards of that tim, volution was what may fairly alld Critis of voluti
a psudosin. Oviously, hristian opponnts of volution dislikd in- Gorgs uvir, m;
tnsly the antiprovidntial undrpinnings of th dotrin. But volution was mummifid bodis o
not assoiatd with total nonlif, athism, or vn what latr in th nin. from gypt. Althou1
tnth ntury Thomas Hnry Huxly was to all agnostiism. ost volu- tmporary forms an,
tionists wr dists who livd in God as unmovd movr' a ing who had with th positiv at
st th world in motion and now lt it unfurl without nd lf miraulous in- that rally ountd.
trvntion. For th dist' inded, volution was proof of God,s powr and in- hatrd of a dotrin
tntion rathr than disproof. vrything was plannd forhnd and wnt than thy ould d z
into fft through th lws of natur. In rasmus Darwin's words: ..\)hat lution, whn oppon.
magnifint ida of th infinit powr of GRAT ARITT! T us provd tru and saft
oF AUss! PARNT oF PARNTS!Ns NTIU!' (1801' 2: 247|. Thr is a as for vry othr o
link hr, spially in ngland, with th Industrial Rvolution. Popl wr final aus. Th Fr
harnssing th fors of natur-watr, ol' and othrs-to produ goods not ignord; indr
through mahins rathr than y hand. Th god of th dist was th ultimat lif. Th ky to un
The llistor of uolutior Thought 5
A Dlnwl*tA lDA.
t.lfitD lr 1n. .rlt ilor.
l' i!' o l
r a singlearhtyp,and yet th
l w a sr h B r i i s h n a t o m i s r Figure2. Priz pig. An industrial rvolution means that pople lav th
s,''althoughit ws not until after ountrysidand go t th fatorisin th itis. n, far fwer peopl left
at thy eameunivrsallyaptd on the land to produ th nation's food. Thr has to an agriultural
revolution,and at th nd of th ightenthntury popl disovrdtht a
major key to suesswas th redingof ttr-qualitylivstok-fattr pigs'
hairirshep,and iggrattl.(From Punh, |865.)
simply sujt to th physial laws of natur ut was organizd. Th parts in human natur, t
wr dirtd to th nd of th funtioning whol, and ah individual fa. Now all of this is i
tur playd its rol in th ovrall, purpos-dirtd shm of things: of organi ration
th hour hand is t
Natural history nvrthlss has a rational prinipl that is lusiv to
w do of that latt
it and whih it mploys with grat advantag on many oasions; it is
rtrogrssion of th.
the oditios of eistee or' popular|,
fil uss. As nothing may missil, but a forl
ist whih dos not inlud th onditions whih mad its xlstn
rs 1846, 40040
possibl, th diffrnt parts of ah ratur must oordinatd in suh
a Way as to mak possil th whol organism, not only in itslf ut in
On th oth sid
its rlationship to thos whih surround it, and th analysis of ths on-
aout th signifian
ditions often lads to gnral laws s wll foundd as thos of alula-
modifid ut thinkin1
tion or xprimnt. (uvir 1817' 1: 6\
ral man of Sottish
uvir's point simply was that th organism is far too intgratd-organizd grs: ..It would augl
and ompl-to allow signifiant hang in any dirtion. It is rtainly too wr inftd with th
intgratd to allow hang from on spis to anothr. organisms at mid. taintd with Matria
point would litrally nithr fish nor fowl and hn would simply un- rfltions that still fi
a]to xist or surviv. volution was in som snsa thortial impossiility, th slaknss of sho
as wll as mpirially unfoundd. diating th unsoun<
Rligion was involvd too. Progrss gos against th hristin dotrin of startd into puli fa.
Providn. Nvrthlss, although Cuvir thought that thr was vidn n, and of sapping
of Noh's flood, nithr h nor othr serious sintists wantd to mak th trstingly' this kind
as by simpl rfrn to Gnsis. Indd, y th ginning of th nin- viws. Although, fro
tnth ntury' all wr starting to ralize that th rth's history must far flt th nd to makr
oldr than th traditional 6,000 yars that on an work out from th g- aov is from th fifl
nalogisgivn in th Bil. It is not that th Bil is fals, ut rathr that it mor rdit to th Cr
nds intrprtation. Som solved th prolm y thinking of th si days of of ours, nithr
ration as si long priods of tim; othrs solvd it y supposing that thr ral viden, and fin
wr long, unmntiond gaps twn th ilial days. God's ration thre- all to miral may
for was a long, drawn-ut pross' ut it was not volutionay. good sintifi soluti<
Th ontrovrsywas at an impass,and not muh had hangd y th mid-
dl of th nintnth entury. on the on sid wr th volutionists' om-
mittd to progrss and ardnt in thir lif that organi dvlopmnt was Charls Darwin
th perft omplmnt to this idology, with nthusiasm outstripping mpir-
iai knowldg. onfirming this pattrn' ]n 1844 th Sottish pulishr harls Robrt Darl
Robrt hamrs wrot (anonymously)a highly popular work on volution, train in th family
Vstiges of tb Ntrl Histor of rtio, in whih h argud that vry- droppd out, ord r
thing was in a stat f upward oming and that wht happnd in th so- rady Darwin had st
ial world mirrord what happnd in th iologial world: std in th living w
anatomist and an -v
Th qustion whthr th human ra will vr advane far yond its work (th young h'
prsnt position in intllt and morals, is on whih has ngagdmuh tion was an ida to r
attntion. Judging from th past' w annot rasnaly dout that grat nvrthlss, that thl
advans ar yt to mad; ut if th prinipl of dvlopmnt ad- whol of hristianit;
mittd' ths ar rtain, whatvr may th spa of tim rquird for was a fator in his r
thir ralization. A progrssion rsmling dvlopmnt may trad th stalishd Chur'
b Histor of uolutior Thought
Lturut was organizd.Th parts in human natur' oth in th individual and in larg grouPs of mn . . .
ng whol, and ah individual fa- Now all of this is in onformity with what w hav sn of th progrss
'-dirtdshmof things: of orgai ration. It sms ut th minut hand of a wath, of whih
th ho hand is th transition from spis to spiS. nowing what
lnal prinipl that is xlusiv to
w do of that lattr transition' th possiilit of a didd and gnral
/antagon may osions;it is
rtrogrssionof th highst spis towards a manr typ is sa ad-
|, fil uss.As nothing may
missil, ut a forward movmnt sms anything ut unlikly. (Cham-
litions whih mad its istn
s 1846.400402|
tur must b oordinatd in suh
ganism'not only in itslf ut in
on th thr sid wr th opponnts' ommittd to Providn, vol
Lit, and th analysisof thson-
aout th signifian of final aus' and apting that Gnsis must
vll foundd as thos of alula-
modifid but thinking that this ould radily don. David Brwster' gn-
ral man of Sottish sin and iographr of Nwton, ould s th dan.
;m is fr too intgrtd-organizd grs: ..It would augr ill for th rising gnration, if th mothrs of ngland
, in any dirtion. It is rtainly wr inftd with th rrors of Phnology: it would augr Wors wr thy
to
;isto anothr. organisms at mid- taintdwith atrialism.'' Th prolm, Brwstr gloomily onludd with
lwl and hn would simply un- rfltionsthat still find muh support in many irls today, stmmd from
m snsa thotialimpossiility, th slaknss of shools and univrsitis: ..Prophti of infidl times, and in-
diating th unsoundnss of our gnral duation, .Th Vstigs . . .' has
s aginst th Christian dotrin of statdinto puli favour with a fair han of poisoning the fountains of si-
lr thought that thr was vidn n)and of sapping th foundations of rligion', (BrWstr1844' 503). (In-
ious sintists wantd to mak th trstingly, this kind of attak sms to hav onfirmd hamers in his
d, y th ginning of th nin- viws. Although, from th start, progrss was th litmotif of his ook' h
tht th rth history must far flt th nd to mak his point vr mor xpliit. Th passag just quotd
at on an work out from th g- av is from th fifth dition of Vstigs and rplas a passag that givs
th Bil is fals' ut rthr that it mor rdit to th rator. [Figur 3])
lm y thinking of th si days of of ours, nithr sid was ally satisfatory. Idology is no substitut for
.s solvd it y ral vidn, and final aus an b ignord ut dos not go away. Rfring
supposing that thr
ilial days. God's ration thr- all to mial may soially and psyhologially omforting, ut it is not a
Was not volutionary. good sintifi solution. Th tim had om for a signifiant stP forward.
not muh hd hangd y th mid-
: s i d w r t h v o l u t i o n i s t s' o m -
lif that ogani dvlopmnt was Charls Darwin
i t h n t h u s i s mo u t s t r i p p i r r g m p i r -
,, in \844 th Sottish pulishr hls Robrt Darwin (1809_1882) was snt to dinburgh Univrsity to
highly popular work on volution, train in th family tradition of mdiin (Figur 4). Aftr two yars h
lz, in whih h argud that vy- droppd out, ord with th lturs and voltd y th oprations. Yt al-
and that what happnd in th so- ady Darwin had startd to mi with sintists, spially naturalists intr-
iologial worid: stdin th living world. On of his aquaintanswas Rort Grant, an
anatomist and an avowd volutionist. So, quit apart from his grandfathr's
will vr advan fa eyond its wok (th young harls rad rasmus's major tratise,Zooomia), volu-
is on whih has ngagd muh tion was an ida to whih Darwin was introdud at an arly ag. It sms'
Lnotrasonalydoubt that grat nevrthlss,that th youthful Darwin aptd in a fairly litral form th
prinipl of dvlopmnt ad- whl of hristianity, inluding th arly haptrs of Gnsis, and that this
th spa of tim rquird for was a fator in his rdirtd hoi of arr: to an ordaind ministr in
ing dvlopmnt may trad th stablishdhurh of ngland. To ahiv this nd on ndd a dgr
8 The Histor of uolutionar Tbought
from an nglish univrsity, and so, in 1828, Darwin was pakd off to
hrist's Collg, amridg.
In 1831 Darwin (who ontinud mixing with sintists)got his ig rak.
Aftr h graduatd, his arr as a lrgyman Was put on hold through th of-
fr of a lngthy voyag on HS Begle,just about to start on a survying trip
around South Ameria (Figur 5). A major influnt this point was (viar- Figur 4. harls Darl
iously) th Sottish gologist Charls Lyll, who at th eginning of th d- 1 8 3 9 j u s t f o r h e m l
ad egan pulishing his massiv Priipls of Golr>g(thr wr thr liknss(no surpris's
volums; Darwin took th first with him and had th othrs snt out). AI- portraitist in ngland
though h was no volutionist, Lyll insistd that th physial world must from a very rih, midd
.Wd
xplaind in trms of natural auss of a kind now still working (Figur 6). wif' was Josia
mass prodution.Alth
This had a great fft on Darwin, whos first systmti work was in gol-
wok for a living. Th
ogy, nd prpad him to think aout th ogani world likwis lss in bi-
plains w n th |
lial trms and mor in trms of natural auses. In 1835, laving South frindly fashion. Dar
Amria, the Begl saild into th Paifi oan and visitd th Galipagos Wdgwood family.
Arhiplgo' a 8rop of islands on th quator, far from land. Thanks to th
governor' who pointd out that th giant tortoiss indignous to th arhi-
plago wr diffrent from island to island, Darwin am to s that this hld plaind was throu[
for th Galipagos fauna gnrally-th irds in partiular, th finhs and radin arly 1837
th mokingirds, wr puliar to thir spifi homs (Figur 7). Th dif- li thinkrs, what w
frns had to b signifiant, and th only way th signifian ould .Without
volution.
The Histor of uolutior Thought 9
-/B
---B
_---F
iii,i:.:.'.:' P
-i
AN ATTE1I
Figur 5. }{S Begle.The Bgle was aptind y Rort Fitzroy, thn only
2 earsold. Bing a aptainon warship would a very lonly Post aus
on would st asid from th rw. Fitzroy thforlookd for somon
who ould b his ompnionon th trip, somonnot in the navy ut II
gntlmanabl to pay his own bills. Darwin fit th jo prftly,rapidly
volving from aptin's frind to full-tim ship's naturalist.Th rw usd to all
him ..Philos,''short for Natural Philosophr_the trm thn usd for sintists.
.sithout
allss' h ould nvr th Nwton of iology. Darwin wantd to
find a biologial for that would explain th volution of lif and was uni-
vrsal in th world and th ky hind all organi motions' and to this nd h
workd fvrishlyfor th nxt 18 months, spulatingnd ding an tnsiv
litratur rviw.
Darwin soon ralizd that th proal ky to hang ly in somthing
paralll to th way in whih animal and plant rdrsfft hang, namly,
th sltin of th dsil and th rjrion of all othrs. Brrt how was oD
to gt a natural quivalnt to th rdrs' artifiial sltion? At th nd of JOII]
Sptmr 1838 Darwin rad n ss on th Prinipl of Popltioz (sixth
dition, 1826| th Rvrnd Thomas Rort Malthus. In his work
althus argud that food supplis would always b outstrippd y potntial Figur 6. Priniples o
population growth. Hn' unlss thr was ..prudntial rstraint,'' thr his mthodolgialin
wr bound to ongoing struggles for xistn as popl omptd for th
availl food and supplis,inluding living spa (Figur 9). Darwin turnd
althus's rasoning on its had. H pointd out that thr ould no
prudntial rstraint in th animl and plant wrld' and hn,bauspopu-
lation growth rtainly Was not rstritd to humankind, thr would an
ongoing' organi-wid struggl for xistn. or than this, suss and
failur in th struggl would (on avrag) a funtion of th diffrnt
Tbe Histor of uolutior Thought 11
PRINCIPLBS
G o Lo GY,
ElIiG
Abingdon
\
indIoo
owel
A|bemr|o e hr south meri tofuis
fudo dbingdonii shldo bul
fiom Abidon l5lnd
fudo vi.in
fom Almr| |sland
e*udo ephiwiu
from Dunn lnd
on
BindIo
Iow l
t^
\,J he south merin toise
studobulfu
s
GALAPAGOS
ISLANDS
l..s-y
J
| > Indefstigable
\*-/ / J /
/'2
.rigton <-'/,/ h.ath.am
harIes
/> ood
rawn in July 1837' is from a privat Figur 9. Th althusian xpltlsion. This arton appard in the omi
lisprolem.Lamark had a tr. lmk in 1851. On of tlr rasons tht l)rwin was al to onvin popl
)nan find thm in th writings of of th truth of volrrtion Was' apart frtlrn lris frindly styl, th fat that h rlid
g to show th ordr in whih God on idas tlrt wr widlv rltd in Vitorian Britirin.
I+ Th Histor of uolutionr Thought
8, ut by thc t,Iiment
purposful natur that pitoizs final irus. \Y. Wvrl : Ba
Darwin sat on his volutionary idas for 15 yars' during whih tim h
's7' ..l. l t no lt of s
turnd to a massiv study of arnals. ar not quit sur why this dlay nclud, thrcfor'
ruritY. or ill.aPli mdl"tioD, thiDk or:
vll sfudi.'d io t
ourrd, although y this tim Darwin hd falln sik with a mystriousail. ..*o.u'' .h @ lar o i
Bo; .4dvlre
Was not rlishing th hug dattlrat his idas wr ound to aus. Also a
major fator must hav n his rlutan to upst powful sin-
stalishmntfigurs (inluding mridg mntors) who had nouragd
th young Darwin in his work. On of th things h did during th paus was
to ntwrk with youngr sinrists'who ould rally around him whn h did
go publi. Finally, howvr, Dzrrwin was pushd into ation whn, in th
.Walla, Do' Bfolq, K|'
middl of 1858, h rivd short ssay y Alfrd Russl a oll- o.ros' l!l' 1859.
tor in th Malay pninsula' that had virtually th sam prmiss and onlu-
sion that h had disovrdsom 20 yars arlir.
oN oRlGlN oF sPls
Darwin quikly wrot up his idas, and ()n th rigin of Spies b es Figur 10. Th titl pag
of Ntrl Seltion, or th Preserutio of Fulred Res i th Struggle hky in quoting th p
for Lif apprd in th fall of 1859 (Figur 10). Th work ginswith a dis- th topi of th law-gor
ussion of artifiial sltion rrd animal and plant rdrs'grat susss. Drwin's mntorswhr
This is partly huristi,t prpar th rdr by following th path tht Dar. ontinud in this r<llir
win himslf had takn to disovry, and partly justifiatory,suggstingthat known opponnt of v
Darwin snt a omplir
what w an do, natur an surly do ttr.Following is a disussion of th
y anning the ook fr
variation tht lways ours in th wild, somthing that Darwin oviously
h ws astr (had).
ndd as th building loks of hang. If vry organism Wr xatly lik
its parnts, thn thr ould no volution' At this point Dawirr was rady
to introdu th strugglfor istennd' following on this, th mhanisnr
of natural sltion: Not that vn mor
for rprodution. lt is
A struggl for istn invitaly follows fr<lmth high rat t whih ll sxal dsirsof a phi
organi bings tnd to inrils. vry ing, whih during its natural way' givn naturally
liftim produs svral ggs or sds,must suffr dstrution during
som piod of its lif, and during som sason or oasionl yar' oth- Canit...thou1
rwise, on th prinipl of geomtrial inras, its nurrrl-lrswould hav undoutdl1' .
quikly om so inordinatly grat that no ountry ould support th ah bing in th g
produt. Hn, as mor individuals ar produd than an possily sur- ur in th ours o
viv, thr mllst in vry as a struggl ftlr istn,ithr on indi- doubt (rmmbri
vidual with an()tof tlr sm spis,or lvith th individuirls of distint sily surviv) that
spis,or with th physil onditions of lif. (. Dawin 1859' 63) ovr othrs' would
Tbe istor of uolutior Thought 15
for 15 yrs, during whih tim h PltEsIivAloN o!' FAVOURD RAcs rN T TIiUGGL0
! ar not quit sur why this dlay ..'l nlud' thefe{orc, lt no msn out
thik
of o wk
o mritain'
oncit
ttt
of
t
F'o'r LJ .
liety, o ill-spplied modati0n,
had falln sik with a mystrious ail- lrrn u sr fu f o s t ell fudi d in the mk of God's
worJ, o in thr k f Gorl's ltork; diviniiy or pllilsolit; ut
st of his lif' and so undoutdly h rth0r let meD eDdvoul nn ldles grss r ploficine in both.''
| d'turnt o! Iain(]'
Lisidas wr ound to as.Also a
B^
Ry OHARLDS DR1r!TN' 1{'A.,
-ltan to upst powrful sin- lL o trE
lJoURN^l
Y^L' GEoIDGI^L' LlNNt^n, .' sor8;
.ticfu@n
tlrir kind? On tlr othr hnd W Iyfl sur that any r,rizrtionin th ltion partiularly pr
last dg injurious worrld rigidly dstrlyd.This prsrvatiorrof th sam spis:sl
favouralvariations and th rjtionof ilrjurious variations,I all Nt. istis rthr thn sp
u r l S l t i t l n(. 8 0 _ 8 1 ) s sltion (inludin
aving thus prs
In latr ditilrrsDawin intodtrd th ltrrtivtrm gtruiul of th fittest, dud th famous m
wlrih was prhps a littl unfotulrat,for it has ld to ndlss laims that s- sam lass hav soml
ltion is a tutollgy, rduing simply to th lairn tht thos who surviv ar il largly spaks th
thos who surviv. But althouglr this is oviously tru, sltionmans mor spis xtant today.
than tlrat. It laims, ior ttr lr flr wors' th21ton vrag tlros who surviv volutionary paths of
a diffrnt frr-nthos whtl do not, and thilt sussin tlr struggl is a fun- vry first shard orig
tiorr of thos diffrns.This may ()r may not tru, ut it is not a tautology. and ths, if vigorou
Darwin inludd a sondary sltiv mhanism. H argud that th ranh, so y gnrz
stugglis not always for food irnd spa ut an dirtly fo mats. This whih fills with its d
mhanisrn, stll seleti>tl, Darwirr dividd into twtl Prts. Thr was s- rs th surfa with i
ltion through lal omat (flrf;rrals). Tlr antlrs of th dr would win 1859, 129-130)
th produt of this. hn thr was slti<lnthrugh fmal hoi (of th Now, with som -
most dsiral m:rls).Th tail fathrs lr gnral plumg of th paok prsnt th sond
-l
or sirrrilarird wol] pr<ldr"rt lf this (Figtrr 1). Noti hrwsual s- Darwin took th ra<
instint, palontology
and showd that ph
through natural sl
support th mhan
fl sur that any variation in th ltion partiulalyputs th mphasis on omptition btwn mmrs of
r"dstroyd.This prsrvation of th samspis:sltionis dirtd toward individual-nfitingharatr-
of injuriousvaritions'I all Nat- istisrathr than spis-nfiting haratristis.Darwin tndd always to
ssltion (inluding natural sltion)ating in this way.
Having thus psntdth min Inhanisms of hang, Darwin intro.
trnativ tJm suruiul lf the fittest, dudth famous mtaphor of a tr: ..Th affinitis of all th ings of th
,r it has ld to ndlsslairns that s-
samlasshav somtimsn rprsntdy a grat tr. I livthis sim-
l i n lt h t t h r l s w h o s t t r v i v il largly spaks th truth.'' Th lavs and twigs at th top reprsnt th
bviously tru, sltion mns mor spisxtant today. Thn, as w go down th branhs' w hav th grat
' that on avragthos who surviv volutionary pths of ystrday. All th way down w go until w rh th
tht strssin th strtrgglis fun- .,As
vry fist shard origins of lif. uds giv ris y growth to frs uds,
not tru' ut it is not a tautology' and ths,if vigorous, ranh out and ovtop on ll sids many a fblr
mhanism. argud that th ranh, so y gnration I liv it has n with th grat Tr of Lif,
bllt an dlrtly for mts. This whih fills with its dad and rokrrranhsth rust of th th, and ov-
idd into two parts. Th was s_ rs th surfa with its vr branhing and autiful ramifiations'' (. Dar-
;). Th antlrsof th dr would win 1859, 129_130) (Figu 12).
:tion through fr-nal hoi (of th Now, with sol minor problms brr.rshdaway, Drrvin was rady to
or gnral plurr-rag of th paok prsnt th sond part f his thory. For a good two-thirds of th rigi,
; (Figur11). Noti how sual s- Darwin took th radr through th vaious ranhs of iologial sin-
instint,palontology,iogography,lassifition,m<lrphology,mrvol.,gy-
and showd tht phnornna in ths ranhs ar plained y volution
thrughnatural sltion,and, onvrsly,thsvarious branhs point to and
support th mhanisnr of volrrtion through sltion (Figur 13). hr
h animalrdingld Drr,vrnto
:tion.Naturl sltion
rlpsurvival,lik thikroars,
tion fr ttriutsthat giv pla-
mor foiousfightingqulitis.
[a.(Bustl.t()rigin ws writtn Figur 12. Th tof lif s given in t rigi. Noti that th mphasis in
or illustrating.Most of Darwin's this pituris lss on showing lif rahing up and mor on th way in whih it
iouslyillrrstratd.) spreds'
18 The Histor of uolutilrl Thought
would all th nr
p l a i n ds i t n l s e
n o l o n g r t h r , r .
rought out all of D
whr now th Wrr
not find it. vn if w
dotrtftll that w wou
prssd and mtamor1
o s t o t r r a so f
-l -l _l -l l
trir-rtion(iogograph
is that on finds th va
L__ltt ll tt II tl
Why, fo instan,do
L__lL__lII L__l
that ar xhiitd y tl
lt
lnstint Paleontology cographial lssifiation orphology mryology
simply that th found
th mainlnd, nd on.
sify undr th nw s
Distribution
Figur 7). Systmatis
Figur 1.3.Th struturof Darwin's rgumnt.l)arwin did not hit y hanon volution. As Linnau
this kind of argumnttion,whr a unifyir-rg hyplthesisplainsin many ally in nstdsts,thu
'Whwll,
diffrntarasand in turn is justifidor suportdl.thsras. who opology lrrsthos
alld it a ..onsilin
of indutilns,''mad muh of it whn disr.rssing
diffrnt spis.ovi.
sussfulthtlrisof physis,spiall1' Nwttlnianmlrnisand th wav
thory of light.Whwll idntifidth irussat th ntrof onsilirr ryology likwis was .
s
..truauss,''wht Nwttln hd alld Urd(l|ts. Showinga onsilinwas mbryos of sor diff-
thrforn ssntilstfatgyfor worrld-bNwtlt-lof iology. whras th adults r
fat tht in th wonr
simila (thywould nor
th rwo adults would l
Wr som prolmati isstts.Soial hlvior, prtitrllyth:rt shown by Hr' as lways thou
th soial insts (th hymnoptra-th ants' th s, and th wasps), analogy with th worl
would asily plial if sltion favlrdth group. But frrm an indi- ..Fanirsslt
hand:
vidualisti prsptir,,why slror"rldol1 gt stril lr,orkrs lvlro smirrgly ar nrly 8rown up:
spnd thir whol livs dvotdto th lvll-ingof othrs irl th nst or hiv? struturs lrv n a
How an natural sltion,with its mphasis on th importan of rprodu- posssssthm'' (. D'
tion, rodu organisms that an<rtrprodu? }.inirlly,Darwin' wo al- So w ar ld to tt
ways favord individulisti pproahs ovr grLlp ppr(lhsaus h hristianity lrd volr
thought th lattr too opn to hting and ploitation, didd tht in suh mmrs of th \/d
ass th group mmrs ar so ll,ll intgatdthirt it is prmissilto trat ld that h Was W
th ll,hol hiv as kind of suprorganismin lr,hith individual instsar unslf-onsiousrfr
parts of th whol. Just as sltiltltr work oll th v' for ampl, whih sm to fully satis
xists for th nfitof th whol rganism, so th workr ists for th n- dntly ratd.''This r
fit of th whol hiv. tr with what w kno
Palontology also raisd diifiultis. It lrad its good points-for instan, th prodution and
that s W go down tlr rorl,inrasingly w fin1orgnisrs that srnto shuld hv n du
hav faturs midwy twn fatr-rrsof xtrrt <lrganismsthat ar widly and dath of th indi
diffrnt.owvr, against th positiv, tr wr gps in th fossil rord, Wht God dos not d
and' vn wors' th fossil rord gan atlptl-vt t start of whr w nss in supposing oth
h istor of ulltilrThogbt 19
-l
doutfulthat . would find lif; th prssur from abov had surly om-
prssdand mtirmrphosdth fossil rmains lrlow (Figur 14).
ost oth ars of iology fll int<-l pla witlr mor s.Gographi dis-
l
triution(iogogrplry) rvas triumph ttsD:rrwin plaind iust why it
is tht on fin]sth variolts pattrns of arrirland plant lif aourrd th glo.
L_l
__lL__l
dos on hav th strng sorts of distriutions and pattrns
!hy, for instt-l,
that ar hiitd y th Gal5pagos Arhiplago and othr island groups? It is
simply that th ftrundrsof ths isolatd islirnd dnizns am y han from
th mainland, alld on thy wr stalishl,thy startd to volv and divr-
]lassifition Morphology mryoIogy sify undr th nw sltiv prssurs to wlrih thy wr now sujt (s
Figur 7). Systrirtislikwis hiits th kind of pattrns on pts from
lnt. Dwin did not hit y han on volution. As Lirrrrattsdmonstratd, organisnrs an b lassifid hirarhi-
lg hyplthsisl:rins in many
allyin nstdsts,tlrusslrowinga pttnthat vlspast history (Figr 15).
portd by ths :rrzrs.Whwll, who
Morpholog1,hirs thos similaritis,lld lromologis,btwn orgnisms of
n l t l h r l f i t r rh n d i s u s s i n g
:wtonin mhnis and th wav diffrntspis.oviously thsspk to slrardanstry(sFigur 1). m-
s at th tr of a onsilin as ryologylikr,l,isWas a partiular point of ridfor Drwin. Why is it tlrt th
us. Showing a onsilin was mryosof sotldiffrnt spis,suh as man and th dog, ar vry similar,
Nwton of iology. whrasth adults ar vry diffnt? Darwin argud tht this follows frorn th
fat tht in th wom th sltiv fors on th two mryos would b vry
similar (thywlr"rldnot thrfor torn apart)' whras th sltivfors on
th rwo adults wold vry diffrnt(thy wor"rld torn apart) (Figur 16).
,havior,partir-rlrlvthat shown y
r, as always tlrrouglror-rt Darwin turnd to th
his disussions lf vllr-trion,
t ants, th s, and th wsps)' anabg,vr,r,ithtlr world of rdrsin tlrdr to larify and support tlr point t
vord th group. But from an indi- hand: ..Fiu-rirs sltthir horss, dogs, rld pigons, for rding' wlrn thy
gt stril ,orks who smingly ar narly gro\vn up: thy ar indiffrrrt rvhthr th dsird qualitis nd
l-ingof othrsirr th nst or hiv? struturshav bn aquird arlir or lat in lif, if th full-grown animal
asis on th importanof rprodu- posssss thm', (. Darwin 1859,446).
lrodu? Finally, Drwin, who al- So w ar ld to th onluding passgsof the rigi. Darwin, whos
ovr group approahs aus h hristianityhad r,olvdinto a form of disrn (Lyll, a Unitarin lik many
Ldploitation,diddthat in suh mmrsof th Wlgwood family' Was mjor influn hr), nvr on'
gratdthat it is prnrissilto trat ald that h w,:rsr,vlrkingin a God_kl mod, and thr ar frqunt
r in whih th individalinstsar unslf-onsiot'ts frnsto th rttl' ..Authors of th lrighstnrinn
lork on th v, ft>rampl, whih smto fulh. satisfidwith th virv tlrlrt ah spishas n indpn-
m' so th u.orkr ists fr th n- ,.To my mind it
dntlyratl'''This was not Darwin's ptlsition. aords t-
tr with wht w know of th laws irnprssdon mattr y th rator' that
had its good points-for instan, thprodutionalrd tintion of th past and prsntinhaitantsof th world
gly w find orgarrismsthat sm to should hav n du to sondary auss' lik thos dtrmining th birth
)f tant organisrs that ar widly and dath of th irrdividual'' (. )rwin 18's9' 488). God aks th world.
.hrwr gaps irr th fossil rord, What God dos Irot 1ois gt involvd in th world, and sinhas no busi-
r aruptly at th star of what w n s si n s u p p t l s i n gr l r h r r r ' i s .
20 h Hist<.lr lf uolutior Tbought
Tray.
o Shetl-arl.
N Glacial Drift. i fl y Wapon.
o Brick Ert. E8
z
Nowih 1
Red }Crg.
Comllin )
o
Flun. 3
s lase.
"" .'"-"-.. . . . ;I E
F Gypr.
A,
UJ
e
t- !9o{9n
rltl,
}t"",.
fgu]ta. Canivr
l
astricht' Cyloid. l _ osaun.
Uppe Chalk. Ci"ooid. l !rg,g' olyptyodo'
Iwe Chalk'. s Brs, Bone.
Uppe leensand Prrlian Coodilia.
Iver Grensand
o
N 'lYeld
la1..
N IatingSan.l Igunodn.
'l Puck Bd. _l\Iasupial' - Cheloni y Bone.
5 Kimmeidgian. P]iosauu'
ofodian.
ellovian.
"E- * .f:{
ltoest ble.
Bath-Stone. EE
o
z StonofieldSlt' o
asupial.
o Grat oolit.
o
Li. Ithoptegia.
"{-*-";:":"w
lrl
U' BonoBed.-Ar vrAL|
Figue 15. Th Linnaa
w i t h t h m m r so f | o
o m P h n s i v s) r sa t l
s e t ( k n o w n a s t a a ,s i n 8
shard past' althoughas
without rgardto volu
interprtationgavit n
hypothsisof shrdd
.
l1 DEscEN |
!
#;dse
)F APPAANOE or ANIA, ,rFE
l'
rffi
>1.0o0,oo0speies (nim|s)
.";;i;.^-"6.
a--;.;"; ;.
di
"*Wehf?
&
I
6
..-;;H'r't"+$:s
d
5,,l60 seies (songbirds)
trl
ld
,d.l. osauu.l
id. r l8lrs' PoIptyodn.
I
' y Bone. l
lin Coodilia. l
I
odo.
pial,- Cheloniy Bon
u8.
,hyopteygi. "-t#
i, by Foot-pfints.
totygia.
Figure15. Th Linnaanhirarhy.organisms n all plad in a hirarhy
itodontia. with th mmrsof lowr-lvlsts ing groupd into vr fWr(ut mor
omprhnsiv) stst highrlvls.All organismslongto on and only on
st(knownas taa, singulartaon) at ah lvl (tgory).This refltsa
'ia, by Toot.pints. sharedpast, although as With th fossil rord' muh of this was workd out
lL|A ganoh. Ineta without regardto volution. Hn, giving th hirarhy an volutionary
interprtationgav it nw maning and at th sam tim onfirmd th
-si hypothsisof shard dsnt.
Ei
.o
I ganOld. ."f
I plao.ganoid.+-
ig.{ plaoid.-
$1'$:'*:w
l] DEscN oF MAN
What aout humans? From th first, Darwin was always ston.old rtain
Fuoids. zoopl\I,os.
that humans ar part and parl of th volutionary pross. His nountr
on te Bgl oagewith th nativs at th ottom of South Amria, th
60. This is takn from Rihard Tirra dl Fugians' had onvind him of this. In the origi, howvr, tati-
wn aman implalnemy of ally Darwin didd not to mak muh of th human qustion. H did not
matrialisti; ut, as young mn
wnt to onal his thinking; h simply did not want it to swamp his gnral
n proalyama
pointsaout volution and its pross.Hn, in on of th grat undrstat.
ing th ourseof history)'ut on
<thesas implying volution. As mnts of all tim, h said mrly' ..Light will thrown on th origin of man
re fossil rord thinking that it might and his history'' (. Darwin 1859' 488). Darwin fully intndd to gt ak to
entially ratdorganisms. th topi and dal with it at lngth. This h did in 1871, offring a major
two-volum work,The Desent of n.
22 Th Histor of uoltilr T'htlugbt
naturlisti prsPtiv
tl ssay on how humal
with spiritualism and .
natural vnt. Thr ml
off of this onlusion,
tristis that ould not
human hairlssnssan
ally livd with nativs,
powr unusd by pril
prodd y sltion.
th Thor of Nturl '
Suh a onlusion w
good ar8umnts. Natul
tiv human faturs,
that is haratristi o
aury and dsirabilit;
.Wlla
wr not suh
rtainly not out of th(
sion of sxual sltio
.Why
ar mn ig and s
stors who wr got tl
domsti? Baus th
st jo with th famil;
anstors found sua
womn (using Darwin'
th standard of ut1
x p t d l y ,a l l o f t h i s i
timnts abot th lat
}.lg.l. Uppl ffgrlrultr.'rJ}*3f;Ji:kr'
didd minn ov
l,owefi$tre thlt of s dog,
mntal powr in man
326-27).
o. For.broln, et.l lrtisphers,
t.
;f. Fist visal arb'
g. solrdvislarh.
No argumnt is n
r,. ]ld.brin. .,rln,8 (llt'lign'is. I{. vetbsl olurnns and multl his thinking wr star
. Ilind.lrrai. erlllunt, medulla in pe:or d0vlor(.ut.
obloug8t. may hav n a rvol
d. e. it.**'ltli]l.
'il }tnri
ties.
. I. I,. o ti yx. tri. It is littl wondt
By univrsal alaim,
Figur 16. Similarityof th rnryoof humirnand dog. Again, W sth
that nglish Valhalla,
strngthof the rigi in r.r.raking
sof fts lik rlrs'wll known to all
iologists.(rom e Dstrl|-.|
Aft Darwin
Aftr Darwin
TtI
. . G tS t i
I . i g u r l 9 . h
uninhaitlusth
<rfvryn'saW sWag
Vitorians th nd for tr
nginring.(Fr om P ul
T ..SILNT }IIGWAY''.MAN
vrything oys this law. Humans ar mor omplx or htrognousthan th individual and th d.
just-quotd ssay on prl
othr animals, uopans ar mor omplx or htrognous than savags'
and th nglish languag is mo omplx o htrognousthan th tongus trm for both. In th
..dsntwith modi
of othr popls (Figur 20). was
..volvd.''Darwin alwa
Inidntlly, it was rrt Spnr who popularizd th word euolutio,
Until th middl of th nintnth ntury' th trm was gnrally rsrvd him a woolly and loos t
for th dvlopmnt of th individual (ontogny). For evolution in th mod- in the rigiz, a work tl
rn sns (phylogny), most popl usd words lik
..trnsformation'' tndd as a maior ontr
(Rihards 1'992). Sing no ssntil diffrn twn th dvlopmnt of us a trm tht appad
to philosophy, litratur
tim of Tbe Dscnt of 1
day had om univrs
and as omfortaly as a
A good rligion' su
of volution' th artiul;
winism. Th asi patt
nss of volution-sir
ai
;2
> mond to man. This is
vry last, not impd i
with th volutionary p
should promot it and <
y volution. Start wit}
nv ours-nd it is :
R
soioonomi soial p<
i6
ptition, and th wak
f3
supportd amlioration
th fat that undr th r
its unhalthy' imbil
ing,thoughwll-mani
th purifying pross br
a th multipliation of tt
Sqtl failing provision, and ,
*
providnt by hightni
z'tt2
*i*
si
( S p n r| 8 5 1 , 2 3 _ 3 2
r: Christians diffr, soI
manings of thir rlig
from anything ls, ma:
world than in th iolo1
. wr lirtarian; othrs
<:
volution; and still oth'
.a !.
appaling to a kind f
i.. low mmbrs of a sp
hism. Som wr fmi
Som wr paifists (S
Figur 20. Th tr of lif as drawn y rnst Hakl. Noti just how th
All usd volution lss
ida of progrssinflundhis pitur of lif's history. (From HekeL1897,
2:188.) living world-as a full-
0 Th Histlr of ulltilrThoght
l"{
\
Th Synthti Thory N-\.J
'\\
Things prsistdthis way into th arly dlsof th twtith ntu nd
in som rsptsam vn wors whn ontinntal thirrkrs,notaly th
mbryologist Hans Drish in Grmany and th philosophr ni Bgson
in Fran, gan pLrshinga kind of no-Aistorlianthory of lif tht szrw
r
spil, nonphysial final fors guiding tlr path of volution. Dris's nt-
lhisnd Brgson,sls uitu' th foundatins of ..vitalisr,''wr iust
{
not th lmnts of forward-looking modrn sin.Rlif finally am
with th dvlopmntof Mndlin gntisnl its mlding rvith Darwiniurn
sltiorr,although vtrhr th pross took til-r-r nd lvirs not stri]ishtfor-
ward (rovine |97I). Grgor ndl, Morvian mtlk who livd irr th
Austro-Hungian mpir, disovrdth ssntialprinipls of hrdity in
th 1860s. Unlik Darwin, h sa'ur, th trnsmissionof hartristisas ..pzlr-
tiulat,'' living tht faturslik olor nd siz n p.rssdon without
th thrat f ing lndd wy (Figur 2l ). Howvr, it was not Llntil th
bginning of th twntith nttrrythat popl am to ppitth impor.
tan of Mndl's work and saw tht it providd ky lmntin th story of
volution. You miglrt think tht this was a trril missd opportunity, and
that, hd Dwin rad Mndl's ky papr, h would hllv rlizd tht tow
h had th answrs to all of tlr issus of hrdity tht his thory of ntural
sltion dmandd. Hen, th thory would hav mlvd forward mor
quikly' muh rlir. This is proaly not sO. vn whn th work of
Mndl was disovrd, it to<lk tim to ssinriltit. Marrt. -loplat first
thought it an altrnativto Drrvin rathr thn a omplllnt (slo.)'So
F i g u r 2 l . T h p a p l a n
vn if Darwin lrad rad Mndl-nd th monk's work rvas pulishd in a ndl to work out th
journal wll-nough known that, if h sarhd,Darwin would hav found many diffrentvariti
it-thr is no rason to think tlrt thr wold hav n il ..urka'' mo- so forth. ndl ould
mnt. ndl inidntallydid rd th origi' Ir-rrrstinglv, h trvrthought passdfrom on gnf
of his own work as prtinnt to Darwin,s prtlln-rs. Judgirrgfrom th anno. rsultsw a littl too l
tations that ndl mad in t margins of th Grmn transltion of th he was only rordingr
ovrzalouslytrying to
rigi' h was fr mor intrstdin th thllogialimpliations of volr-r-
sevntnth-ntury pl
tion than in th troules of hrdity that Drwirr fad.Th is rally no strr-
pris hr sit.t,:rftrall' N,Indlwas first nd formost mn of God :rnd
o n l y s o n d a r i I ya p l a n t s i n t i s t .
Thr wr fairly straightforward rasons r,vhyat first, whn Mndl was was rvald, and it
rdisovrdat the ginning of th twntih ntury, no ()l1thought tlrirt Finally, around 1930,
h was spaking to Drwinin issus'Natr'rrll,rly ntiistsfr.rsd on taly R' A. Fishr (19
ig vriations and so tndd to favor a kind of saltat thory of ovrall Wright (197' t9z)
hang, that is volution y lag jumps. Slowly' howvr' thanks prtiu- slvs and ar transmi
larly to work in th sond dd of th ntury y Thlmas Hunt organ works nly in th grou
and his studntst olumia Unir,rsityin Nrv York, th ntllr of th gn work to rh mplti
Tbe Histor rlf uolutionr bought 1
sns why at fist,whn Mndl was was rvald,and it ould sn as th omplmnt tO natural sltion.
)ntith ntury' n() on thought that Finally,aound 1930, a numr of highly giftd mathmatial biologists,no-
aturally,arly gntiistsfousd on t a l yR . A . F i s h r ( 1 9 3 0 )a n d J . B . S . H l d a n ( \ 9 2 \ i n n g l a n d a n d S e w a l l
kind of saltatory thory of ovrll !right (|91, I92) in Amri, showd how ndlian gns sort thm-
s. Slowly' howvr, thnks partiu- slvsand ar transmittd irr groups (arrssntial finding, givn that sltion
ntury y Thomas Hunt Morgan works only in th group situation),and thn it ws possilto bring Darwin's
in Nw Yrk, th natur of th gn work to th ompltion that it ndd.Along with mutation (th oming of
2 h Histor of uolutionr Thought
IoN
l*{itNl C :EYl}l,ll 1g
nti fingrprinting for d
dtaild in dward o.
Snthsis (1975), hav po
.W
on lf th iggstprrlms
Figlr 24. Th ovr of th first issu lf ulltir.t.
of thsarly profssionalvolutionistswas of findingpopl,working to thir
standards,who wr nt studyingfruit flisor similar organisrns.Th journal
was launhdwith gntfrom th Amrin PilosophialSoity(flunddy
BenjaminFranklin) in Pildlphia.Rvlingly,t only prsonOn th grxnts
ommittwho opposdth fundingWs th mryologistdwin onklin. Figure25. Puntuatdqu
mryologyWas thn th qunof th illogialsirrs and onklin was not g r a d u a l i s m .. l d d g i r
onvindtht volutionor-rld th asisof ral profssionirlsin.Prhaps l t r n t i v p i r u ( B ) ' i t g l
bausf thsdisiplinarytnsions,volutionistsrsplnddv ignorrng sltationist,whih rathr s
mbryologyand it was not until th 1980s,y wih tim m<llular iology omparativtimsals.F
had ompltlyangdth landsap,that it gin -t a mirjlrlmntin hangan tk pl duri
volutinarystudis. arlygts rorddand a
'' -*:
"i." h Histor of urllutilttrThought 7
: r ;:-'- i ' ;
"".
:
signifian of diit at th m<llularlvl) irnd nw thniqus(tlrus of g-
UTION
-t
,L oF oANI YoLUlN
nti fingrprintingfor dtrrnininghrdity). Soirlhavior nd instint'
dtailedin dwrd o. !7ilson's magnifint survy Soiobiolog: Th Nc,
Sthsis(1975I,hav powrful nw modls of plnation, lik kin sl-
,".l!tl:.: .,.1,,,t''t,',,,,: tion nd riproalaltruism, <rnfirmingDrwin's htrr-rh that th right wy
to plin th intiaisof soial intrirtions,inluding grotrp hvior, is
Ie +r N r.? from an individualistiprsptiv.Baking th thortial woks ar majo
mpirialstudis_for instan,that of Tim lutton-Brok and his assolats
(1982)on th rd dr of Sltl.lnd-tht slrlwtht volution has indd bn
,.,:;:,l:i powerfulfator in th vrllution of soial bhvior. Somwht ontrov-
.it,l
ril saetion i wirl Pqul* siallv, th fild (oftn hr known as uolutior pshlllgy) lso in vry
.ust , . -.' . . ' . . . . . . . . . ' . . . . . Darwinian fashion loks at humankind, trying to xplain suh things s in-
:,
**"eL .
h {attn8 fntiidand sual havior nd mating pattrns in trms of na rr-rralsl_
ai{*P. 'srcirl ,,,.tt,'.,.'t$'
.. '.':'.,:1, tion' Palontologyhas rr revolutionizd. Nlt tlnly ar tl.leontinud
7pod*y. R,,4sr ,i;
mior fossilfinds-rntly the disovry in th ndian Arti of a link -
.t
twn fish and amphiins (Dashlr, Shuin, and Jnkins 2006), not to
:,t,',u!{t;
mntionth finding in Ind<lnsillof littl hr'rmanlikfiglre (<ltoflorsi-
, ,,:
,.:|i
*8: . :,:lil 51-n. ,a t u r a l l yk n o w n a s t l r . . h o i t ' ' ) ( I } o w n t l . 2 0 0 4 ) - u t m o l u l a
. .,.:l:i*
63 thniqushav nald popl to xplor th past ours of volution with
I i' 'll;* vrgratrprision.At th mor Onptuallvl, nw thoris hav n
. I r.,r'.r1ti
proposd(for instan,th th<lryof puntuatd quiIiria of Nils ldrdg
''"1.
, ,,.''-:|':. and th lat StphnJay Gould 11972|\ (Figur 2.5),and idas drawn fron-l
othrarasof volutionary studis hav bn applid to undrstand th pat-
s9 trnsof th past (tr instan,th way in rvhih th lirtJohn J. Spkoski Jr.
.. . ,tli
[1976lusd ologialthoris aout islanl iogograplryto throw light on
103
past pattrnsof animal divrsity, as wll as on th Grat At-nrianInt-
111 hang,an vnt that ourrd ablut 10 million yrs ago whn Sluth nd
.- :" ir
ryffi
,/r.q-..:--,I}
ffi
Anteaters
Armdi||os
Capybaras
Glyptodonts
Figur 27. Pttrns of foss
Monkeys
y o u a s s u m r h t t h o n t i
Opossums how muh volutioniststo
Porupines w hv ome from th ol
Phorusrhaids
Bears Sloths
Camels ouans
Systmatis has bn tran
Cats oxodonts
th oming of omputrs
Dogs
absorbd, quantifid, al
EIephnts likwis is opn to nw u!
Horses th synthti thorists t
Pecaries oame euolutior det,el
Rbbits ptual bakwatr to th
Raoons volutionary studis (ar
Skunks instn,aout th ways
apirs mans shar th sam un
Wese!s th ways in whih org
among th hottst aras (
Figur 26. lr Grat Amrin Intrhng. By trating North Arrrri and Thr ar' of ours'
Sluth Arrlri s if thy wr t''v islands ing olonizd, on an rvork out th importngg-1hg ygr
th ptl ts li lr.lg anl th numrs tlf invadirrg gr.ouPs ln wluld volutionists' partiularl;
Ii rium.
pt at qr"ri s o m t i m sq u s t i o n s o f .
Th Histor of ullutior Tbought 39
Antrcti
Mesosaurus Glossopteris
--ffi
Permin Reptile PInt
*.*
ry#
, . -,.,s *
Anteaters
fr /w
Armadillos
\. Cpbaras
Glyptodonts
Figur27. Pattrnsof fossil organisms.Thsdistriutionsmak snsonly if
Monkes
youassumthat th ontinntswr joind. Phnomnalik thsshow jusr
Opossums howmuhvolutioniststoday still work in th Darwinian mod nd yt how far
Porupines we havom from te rigi itslf. (Unitd StatsGologial Survy.)
Phorusrhaids
Sloths
Touans
Systmatis has n trnsformd y nw ladisti thniqus,muh aidd y
oxodonts th oming of omputrs that hv allowd massiv amounts of data to
asord,quantifid,alulatd, and undrstood (Hull 1988). ophology
likwisis opn to nW undrstandings.Aov all' mryology' a sujtthat
th synthetithorists tndd to ignor, has now, spiaily undr th nw
nameuolutilrdeulopmt (evo-dvo), n onvrtd from a nonon.
ptualakwatrto th most xiting ar of rsarhtoday in th fild of
volutionarystudis(arroll 2005). Fantsti nw findings hav mrgd,for
instan,aout th ways in whih organisms s divrs as fruit flis and hu-
mans shar th sam undrlying gneti mhanisms for dvlopmnt, and
th ways in whih organisms ar uilt and vriations ar produd ar
amongth hottst aras of rsarh (Figurs 28 and 29).
ng.By trating North Amria and
Thrar, of ours, ontrovrsisand diffrns.No on wants to deny
rds ing olonizd, on an wlrk out th importan-th vry grt importn-of natural sltion.But som
numrs of invding groups on would partiularly in aras lik palontology and mryology, whr
volutionists,
somtims qustionsof adaptiv signifianar (as was arlir th as for
40 Tb Histor of uolutilnr Thrlught
HAD- x. oo-
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ABDON
Figu32. Th elephant's
nos am aout aus,
that many storisaout a<
)w-tintast,found in th ogs of
(Gilrtet a|. |996,368). No dout postrity will tll us whthr sntimnts
for its antlrs.Thy ar so larg it is
' nof adaptivadvantag.A popu' lik thsar wis and prsintor mrly th unjustifidffluvia of nthusi-
roughsxualsltion,th lks with astsfor a nw disiplin tht is trying to stalish itslf as important. ithr
)ttingth fmls.It is suggstdthat way' th vigor and itmntof ontmporay volutionry thory ar on-
d g r o w i n g n d p r h a p s n i m m a t u r fimd.
r later in life h and his offspring would Dos this thn man that th nonsintifi'mor idologial sid to volu-
this showshow arfulon should b tionarythinking has now vanishd? Dos volution n longr funtion for
r. In on sns'th antlrs r
manypoplas a kind lf sular rligion? Not at all. on th ln hand, hug
I and prhapsnot for muh of th
numrsof artils and olks (not to mntion radio and tlvision pro-
grams)show or justify idologial impliations srid to or drawn from
.Wilson
volution.dward o. is paradigm, for h has pourd out a stram
of ooks dsigndto show that volution is progrssiv nd that from this
;s within morphognti filds provid .Wilson'
flowsnot just mral hortations-for th prsrvation of th rin
n-just as skltaland organ homolo-
forsts-utn altrnativviw of ration to that of trditiona1rligion:
volution is ttr than vr. Th rol
v r ,i s s nt O p l a y l s s a n i m P o r t n t But mak no mistak aout th power of sintifimatrialism. It prs-
l morphologisgnratdy dvlop- ntsth human mind with an altrnativ mythology that until now has al-
o hang if it is not to om as irrl- ways,point for point in zons of onflit, dfatd traditionl rligion. Its
:hanisis to ontmporary physis'' narrativform is th pi: th volution of th univrs from th ig ang
11 h istrlrrlf ulutitl1'Tbought
lf fiftn illion \Irsago thrlugh th origin of th lrrrntsand ls- 10,000 yas old and that i
tial odis to th gir.rnings of lif on artlr. Th v<llutionar1'
pi is rgy in just six days' follor
mythology in th sns that th laws it addus h and now ar - vrsal flood' sav only for
livd ut an nvr dfinitivIvprovd to form a aus-and-fft (Rus 2005; Numrs 20(
()lltinuul fronr physis to th slial sirrs,frol this world to all story hr' ut th onstan
othr wrlds in th visil univrs, and akward through tim to th nthusiastsfor this kind of
ginning of th univrs. vry part of istn is onsidrd to thory ut as somthing m.
ldir-rt ttl plrysil larvs quiring no trnI ontrol. Th sirrtist's tionl lif systms that
dr,ltiontl parsimony in plarrtion xluds th divin spirit and hld morl norms. In th'
oth.rtran()Llsagnts' Most importantly, w hav om to th ruial ..It rl
Hnry . orris: is
stag in th istory of iollgywhn rligilnitslf is sujttl th .pla- punishmnt for murdr n
nrtionsof tlr natural sins.As I hav trid to show' slioiologyan mnt for othr sriousril
aount for tlr vry lrigin of mythology y th prinipl of natural s- v o l u t i o n r yd t r m i n i s mt
ltiln ating on th gl-ltiallyvolving mtrial strutur of tl-rhu- is 1989, 148). Apprntl
rrrnbrin. ..ath
Rus,'' wll_known
If this intrprtationis orrt, th final ]isivdg njoyd y si- th morl rot. lvlor rn
ntifi naturalism will lm from its apait to plin traditior-ralrli- has morphd into th morr
gion, its hif ornptition,as a wholly mtril pn<rnrnon. Tholgy gnt Dsign Thory, whih
is nlt likly to surviv irs n indpndntirrtlltualdisiplin. (Wilson q u i r s p i a l r a t i v i n t r
1 9 7 9 ,r 9 2 ) is mor than just a onfli
pting idologis. Th fou
Alrothr sirrtistmuh givn to rhis kind of spulatio-altlrough h dis- Brkl law profssor Ph
likd trditionirl notiot-tslf progrss and wantd to sustituthis own idas l d t r m s :. . T h C h i s t i n
aout th rndonrnsslf lrang-was StprrJay Gould, partiulaly in his an asy targt atrs it ]
popLrlrssays' ..his Vir,v<lfLif,'' pulishd n-rtlnthlt, in |'trtrlistlr.l,. with worldly ids lik hul
Vry arly did Gould fil to draw slmkind ot moral mssagfrlrrrhis writ- whih r atually profoun
ings, 'uvhthraout th raism ndmi in our slityor th nd for onsr. ingly, this humanisti mt:
vtiorr.Not that Gould gv tradititln;rlrligiln muh mo sop than did gay mariag, aortion, ar
Wilsln. Any idas aout lur ing th fvord hiidrr-rof Gld, tlrt w hu-
rnans might th rason for th ati()n'ar simply huris. ..Sin di-
nosaurs Wr n()t nroving toward markdly largr rains, and sin suh a Conlusion
prospt may ii utsiltl-rapailitisof rptilia dsign . . ' w lnust as-
sLlmthat onsiousnssr,vouldnot lravvolvd On our prlantif a osmi - h history of volutiona
tastrtl;lh had rr<ltlirimd th dinlsaurs as r,itirns.In an ntirly litral until 1859, it was littl m.
sns' W oW otlr istn's larg nd rsrningmanl-tills,tl orrr luk idology of progrss, th r
stars'' (Gould l989, 1ti). Hr Go[d antiip1ltdth so-alldnrv athists' It was opposd to traditiol
lik his grat British ountrpat in th rlm tlf popular sin writing' litral truth of th Bil l
Riha]Dawkins. In the T|la God Dlusil(206\Dawkirls i1r]us tlrt hris- that hang oms only t
tiarrity and otlrr rligilrrsar th r:ljlrsours tlf hur-rrrkind's
ills, and tlrirt on th rigin of Spis ,
w nd :r sular philosophy as a sustitut-a sulaphilosophy informd truth of volution on an
y Dirrwiniirn thinkir.rg,whih itslf mks app.lsto traditional rliiorlsnot tion was ignord or down
mrly tltiosut asolr-rtly fls.Phils<lph Danil Dnlrtt'sDt'itz'sDn- Darwin, notaly Thoms
8rorrsld (7995) is rrtlthrmplr of this kind of litratur. volutionary idas as a ki
Hirding th tlthr Way ar th n.ranyritis lf volutionary thinking. In n altrnativ idology to
Amri tlda,stlvy aft srvyonfirms thilt most Arrrriansdrl not - to soil rform. Th thir.
liv in volutiln.Ind]'th grat rnajlritylrlivtht th artlr is lssthan of ndlin gntis. I
Th istor of uoltior,Thought 4S
tlr origin of th lrrrntsnd ls- 10,000yars old and that all organisms wr atdin urst of divin n.
on arth. Th volutilrrarypi is rgyin just si dirys, followd som tim latr y thir dstrution in a ni-
ws it ddus hr nd ntw ar - vrsalflood, sav only for thos luky pairs that floatd away in Noah's ark
y provd ttl form a aus.irnd-fft (Rus2005; Numrs 2006|. Amian vanglial hristianity is not our
ial sins,from this wold to ll storyhr,ut th onstant attks on modn Darwinism show larly that
:, and kward throug tinl to th nthusiasts for this kirrd of rligion do not rgard volution as just sintifi
lart of istt-l is onsidrd ttl thoryut as somthing lor' a mtrialisti,sular altrntivto th tradi-
] I1Otrnal ontr<ll.Tlr sintist's tionallif systlllsthat arris lvithirr it opposition to ll dnt and long-
rtion luds th divin spirit and ..ration sintist,''
hldmoral norms. [n th words of th lding th lat
ortantly'w hav om to th ruial Hnry. Morris: ..It is rathr ovious that th modrn opposition to apital
r rligion itsIfis sujtto tlr pla- punishmntfor murdr nd th gnral tndny towrd lniny in punish-
hv trid to show, slioillogyan mntfor othr srious rims r dirtly rlatd to th strong mphasis on
lology b,vth prinipl of lli1turals- volutionarydtrminismthat has haatrizdmuh of this ntury'' (Mor-
lolving matrial strutur tlf th hu- ..notorious Dirrwinian philosop Mihal
ris 1989, 148). Apparntlv' th
Rus'''a wll-krrown ..athisti hltratrist'''has mad rnirjorontibution to
h final disivdg njoyd y si- thmoral rot. Or rntly' th rz.rtlrr ud litrlism of ration sin
's :rpaityto xplain trditionl rli- hasmorphd into th mor sophistiatdanti-Drwinism ot so-alld Intlli-
olly matrialphnomnon.Thology gntDsignThory' whih ssasptsof ogani lif as so ompl as to -
,ndrltintlltuldisiplin' (Wilsrn
quirspial rativ intrvntions y th Dsignr. But still th dat today
is mor than just a onflit of sinvsl'lsrligion and on twn om-
ptingidollgis.Tlr foundr of th lntllignt Dsign movlnnt' formr
s kind of spulatir-r-zrltl-rough h dis- BrklylzrwprotsslrPhiilip Jlhrrsorr,is pliit ir-rsing volutin irr th
rrrlwantd to sustittlris own ids ..Th
oldtrms: hristian philosophy that Was ovthrown in th 1960s was
s SthnJay Gould, partiularly in his an asy targt aus it had om idntifid with Amrian ultur nd
pulishd mlnthly in |JturlHistor. with worldly idas lik human prftiilityand th invitaility of progrss,
:rkind of moral mssagfrlm his writ- whihar atually profoundly un-hristian'' (Johnson 1997,106), Unsurpris-
Liin our soitylr th n]for onsr- ingly,this humanisti mtaphysisis linkd to moral failings-pornography'
nal rligilnmuh mor sop than did gaymarrig,aotiol-l'nd lik transgssionssuh as soialisrl.
Lf:rvordhiidrn of Gol, tht w lru-
ration' ar simply hLrris...Sin di-
rrkedly largr rains, and sin suh a onlusion
is of rptilian dsign , We tnust as-
v volvdon our plant if osmi a- Th history of volutionary thory falls into thr stags.From aout 1700
saurs as vitims. In an ntirly litral until 1859, it was littl mor than a psudosinriding on th ak of th
nd rastlningmamtnls, to our luky idologyof progrss,th notion that humirns an mak ang for th ttr.
J antiiptlth so-lldllW thists' Itrvasopposdto tditional rligion not so r-nuhausit wnt against th
h r l m o f p o p u l : r rs i n w i t i n g . it lft no pla for Prrvidn,th lif
litraltruth of rh Bil ut r-rs
tsitm(206)Dawkins argLlsthat hris- thathango-lsonly through Gol's undsrvdgra. harls Darwin's
t)r soursof humankind's rlls, and that th origi <lfSpeishangd ll tlris. Darwin stlishdth rasonabl
)stitut-a sular philosophy informd truthof volution on nd for all. HoWvr, his mhanism of natural sl-
aks appalsto traditional rligions not tionwas ignord or downplayd, and th lading volutionistswho followd
lsophDanil Dnntt's I)ui's D- Darwin,notaly Thomas Hnry Huxly, Wr muh mor intrstdin using
rr of tis kind of litratur' volutionaryids :rs a kind of popular sin,almost a sulr rIigion' as
'ny itis of volr.rtiorr:rry thinking. In an altrntividol<lgyto th lrristin rligilnthy saw [rlokingthir wy
rrfimsthat most Amrians do not - to soialrform. Th third stagstrtd arond 1930 witlr th inorporation
rjoritylivthat th i.rthis lss than of Mndlian ] ntis. Now thr was th possiility of uilding a nW
46 Tb Histor of uoltiorl, Thought
potssional sin oi volutior.rary lrang. This ourrd, arrd it is this th- Gould, S. J. 1989. Woder
Ory that flourishs today. Yt vllr-rtiorrisrsshould nvr flrgt th past and Nw York: W. !7. Nor
its influ On th prsnt. Alorrg witlr th sin thr is still an ongling Gould, S. J., nd R. C. Lw
dbat about idolog1. nd th tnt to lvhih vrltrtion in som wY ro-
Panglossianpradigm
of th Rol Soit l
vids a sulr altrnatir, to oldr' Ovrtly rligiorrs ways Of viwing th
Grnt, B' R., nd P. R. Gra
world and humankind's status within it. Popultio:h Lrg
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;tn: Houghton ifflin. Ruse,M. |996. ld to :1.he lept oi Prrlgrss i L'llutior Bioltlg.
1de.Nw York: Simn arrd Shustr. ambridg,A: arvrd Univrsity Prss.
.slple.London: NIihalJosph. -. 2005. he uolutio-rtil Struggl. Camidg, MA: Harvard
ri,-s,'
Llndon: ilrl Josph. Univrsity Prss.
rigi lf Spis.Nw Yrk: oiumia Sd[y'D. 2008. rtilisttttlt! Its ritis i tiquit' Brkly: Univrsity of
alifornia Prss.
utd quilibria: An ltrnativ to phylti Spkski, J.J.' J'. 1976. Spisdivrsity in th Phanrozoi-Spis-ra ffts.
ldlsi Plobioklg, 82_1 15. San Iobiolog2, no. 4:298-303.
Spprd, . M. 19.58.Nturl Sltiond 17rdit.London: Huthinson.
,f Nturl Sltit-l,Oford: of<ld
Simpsorr,G. G. 1944. lp d dc i z,rllutitllt,Nw Yk: olrrmia
Univrsity Prss.
don: Mthun. -, |949. Th ig lf t,<llutil.Nw lavn, T: Yal Univrsity Prss.
1996. RsynthsizingvlIutiorrilryand Spnr'H. 1851' Soil Sttis; t, h oditit>sssntil tl Hm Hp|liss
ll Billog17: 57-72. Specifidd th First of Tllt Dulopd. Lndon: J. Chapman.
48 he Histor of ulutior hlltght
How, whr,and whn did lif appar on arth? Although harls Darwin
wasrlutantto addrssthsissusin his ooks, in a lttr snt on Fruary
7, 1877, to his frind Josph Dalton ookr, h wrot in now_famous
paragraphthat ..it is oftn said that ll th onditions for th first podution
of a livirrgbing ar now prsnt,whih ould vr hav n prsnt.But if
(ndoh what a big if) rv ould oniv in som warm littl pond with ll
sortsof ammonia and phosphori salts,-light, hat, ltriityprsnt' that
a protinompound was hmiirlly fornrd, rady to undrgo still mor
omplxhangs,at th psntsuh mattr would instantly dvourd' r
absord,whih wuld not hav n th as for living raturs wr
formd''(Darwirr 1887, 'i, 18). (Th lttr is in th Darwin olltion in
ambridg.It is rfrndas Lttr 747 | in Burkhrt and Smith l994 and is
rprodud in alvin 7969,4_5.)
Dwirrlttr not onlv summrizs in a nutshll his idas on th mrgn
of lif but also provids onsidral insights on th viws aout th molular
naturof th basi biologial prosssthat wr prvalnt at th tim ln mny
sintifiirls.Although Fridrih ishr had disovrd nuli aids (h
alldthm nulin) tn 1869 (Dahm 2005)' th diphing of thir ntral rol
in gntiprossswould rmin unknown for mor than 80 yars. In on.
trast'th rols playd y protirrs in mnifold iologial poSsshad n
firmlyrognizd.qually signifiant, by tlr tim Darwin wrot his lr, ma-
jor advanshad ben mad in th undrstandingof th matrial asis of lif,
whihtr a long tim had bn onsidrd fundarntally diffrnt from ino-
.!76hlr,
ganiompounds.Th primnts of Fridrih Adolph Strekr, and
AlksndrButlrov, who hd dmorrstrtdirrdprrdntlyh fsiility of the
laboratorysynthsis of ura, alnin, and sugars, rsptivly' from simpl
startingmatrials,Wr rognizd as a dmonstration that th hmial gap
spIating organisms fronr th nonliving was not insurrtlr.rntai.
19
F
But how had this gap first n ridgd? Th ida that lif Was an emr- pripittdquartz in th
gnt fatur of natur has n widesprad sin th nintnthntury. Th (Uno t a|' 2006), lthou
major rakthroughs that transformd th origin of lif from pur spula. Collom 2006). Howv
tion into workal and tstabl rsarh modls wr proposals that wr diat iologial sulfat-r
suggstdindpndntly in th 1920s y Alksandr I. oparin and J. B. S. 3.4 billion-yar-old Sout}
Haldan, as wll aS othrs (sBada and Lazano 200). Thy hypothsizd rin nvironmnt inhait
that th first lif forms wr th outom of a slow, multistp pross that nd Low 2004\' Ths r
gan with th aioti synthsisof organi ompounds and th formation of old stromatolits from W
..primordial soup.'' Hr thir proposals divrgd. Whil Haldan argud ida tht th arly Arha
a
that viruss rprsntda primordial stag that had appd prior to lis' origin of lif must hav ta
oprin argud that olloidal, gl.lik systms had formd from th soup, prmit th survival of th
lading to anaroi htrotrophs that ould tak up surrounding organi Th arly Arhan foss
ompounds and us thm dirtly for growth and rprodution. Although quird for th origin and r
th dtails of ths idas hav n suprsdd' th oparin-Haldan hy- ritial fator may hav .
pothsis of hmial volution providd a onptual framwork for th sil s soon as th plan
xprimntal dvlopmnt of th study of th rigin of lif. Laoratory watr. Unfortunatly, tht
xprimnts hav shown how asy it is to produ a varity of organi om- o n d i t i o n so n t h l
pounds, inluding iohmially important monomrs' undr plausibl os- lulr or physil rmni
mi and gohmial onditions. Th roust natur of ths rations has volutionry prosss th
n dmonstratd y th finding that organi ompounds ar uiquitous in ganisms found in the arl
th univrs, as shown by thir prsnin arbon.rih mtorits'omtary ing not only on th o
sptra' and intrstllar lods whr star and plantary formation is taking piod of th origin of lifr
pla. gnral and loal nvlro
impotant for th mrg
dgd?Tlr ida that lit was an mr- pripitatd qtlartz in th ..Sillion-yar-old Drssr Formation in Australia
lrad sin tlr nintnth ntury. Th (Unot al. 2006), although tlris finding has i.rlstln hallngd (Lollar and
d th origin of lif from prrr spula- ollom 2006). Howvr, sulfur isotop invstigationsof th sam sit in-
Lrh modls Wr proposals that wr diatiologialsulfat-rduingtivity (Shn t al.2001), and analyssof
s y A l k s a n d rI . O p r i n n d J . B . S . 3'4 illion-yar_old South Afrian hrts suggstthat thy forrnd ln a ma-
nd Lzano 2003). Thy lrypothsizd rin nvironmntinaitd v anaroi photosynthti pokayots (Ti
om of a slow, multistp pross that and Low 2004). Ths rsults' omind with rports on .43 illion-yr-
4aniomptlundsand th formation of old stromatolitsfrom .JstrnAustralia (Allwood et a|.2007), support th
osals divrgd'Whil Haldan argud ideathat th arl,vArhan arth Was tming rvith prokaryots' and that tlr
stgthat had appared prior to lls, hrvtakn pla as soon as tlr onditions wr suital to
originof lif mr"rst
3 systms lrd formd frorn th soup, prmitth survivl of ths tvps of organisnrs.
at ould tak up surrounding organi h arly Arhan fossil rord spaks for th rlativly short timsal r-
r growth and rprodution. Although quirdfor th origin and arly volution of lif on arth and suggststhat th
suprsdd,tlr Oparin-ldan hy_ ritialfatorma hav rrt Prsnof liquid watr' whih ampos-
idd a onptual framwok for th sibls soon as tlr plant'ssurf finlly oold lorv th oiling point of
'rdy of th origin of lif. Laoratory wtr.Unfortr-rnatiy, thr is no gologial vidn of th nvironmntal
s to produ a varity of organi om- onditionson th arly arth at th tim of tlr origin of lif, nor ar any mo-
)rtant monotnrs,undr plausil os. luiar or physial rmnants psrvd that provid infornration aout th
robust natu of ths rations has volutionaryprosssthat prdd th apparan of th fist llular or-
organi ompounds ar uiquitous in ganismsfound in th arly fossil relrd.Dirt inforratiorris gnrally lk_
i n r o n - r i hm t o r i r s, o m t a r y ing not only on th omposition of th trrstrial atmosphr during th
star and plantary formation is taking priodof th origin f lif ut also on th trnperatur' oan pH, and othr
gnraland lol nvironmntal onditions that may or may not hav n
impotantfor th mrgnof living systms'
of Lif
Th Primitiv arth nvironmnt
sumd that th origin nd arly volu-
a s ( L a z a n o n d i l l r 1 9 9 4 ) , s u h onsidral progrsshs en lrradin our undrstandingof nvironmntl
. t h a t i t i s n o r p o s s i l r o a s s i g n a p r - onditionsof th arly arth nd how tlr transition frorn ioti to ioti
l i f ' u t i n r h p a s t f w . r r s s t i m a t s hmistrymay hav ourrd (for mpl. s Bada 2004). Nvrthlss,
v ourrdhav n onsidraly r- thrar still normous gaps in our dsription of how th simpl Orgnr
al origin of wlrat hav lronsidrd ompoundsassoiatdwith lif s w know it atdto gnatth first liv-
1ontntiousissu (varrZuiln t al. igntitisand how tlrsin trrrn volvd into orgnisms tht lft hind a-
clogial rord of th arly Arhan is tulvidnof thir istnin th rok rrd. o valuat how lif may
|ratar mor than 3.5 illion yars ld. havgunon arth' W must asswhat th plant was lik during its aly
i illion yars that ar prsrvd hav historyand undr what onditions th prosssthought to b involvd in
lhi prsss(van Zuiln et a|.2002|, th origin of lif took pla. Lif as w know it dpnds on th prsn of
;int llf has apparrrtly n |arge| [iquidwatr and ogani polr,mrs suh as rruli aids nd proteins. Th
vilalvidn suggsts tl-ratduring arr arly stag, iologial systems
th mirostrutrrrsprsnt in th 3.5 lakedprotinsand dpnddlargly on atalyti and rpliativ polyrionu-
Australian Narrawoon formation leotids'ut watr was likel1.ssntial fronr th vry ginning, as it pro-
,asirt a\' 20021 Garia-Ruiz t al. vidsth mdium for hmial rations tl tirk pla' Withut ths asi
rgd orr ilrth vry arly in its history. omponntsJ i1sfar as w know, lif is irrrpossibl.
niroilmthanognsis,asd on th It is unlikly that watr md its first apparanon artlr as a liquid. Dur-
ts, has n found in hydrothrmally ing th Hadan priod th volatil omponnts that wr trappd insid th
52 he origi of Lif
for thy wr dstroyd y photohmial rations. During ths Butlrov, who show
pisods highly rduing onditions may hav prvaild. lysts, suh s alil
ars. Th laoratory
to inlud mor or
organi ompounds on th Primitiv arth tnth ntury a lar1
formd (Bada and L
Today organi ompounds ar so prvasiv on th rth's surfa that it is Howvr, ths n
hard to imagin th arth dvoid of organi matrial. Howvr, during th as laoratory simul:
priod immdiatly aftr th arth first fornrd slm 4.5 illion yars ago' to Hookr, ut rath.
thr would hav n no orgni ompounds prsnt on its surfa baus of nitrogn assimila
soon aftr artion th day of rdiotiv lmntshatd th intrior of cano 202a). Th fi
th young rth to th mlting point of roks (\/thrill1990). Volani organi ompounds
ruptions plld moltn rok and hot sorhing gass out of th juvnil Stanly L. ivlillr, w-
arth'sintrior, rating a global infrno. In addition, th arly arth ws also a wk on a mixtur
bing ppprdy mountain-sizdplantsimals,th dris lft ovr aftr th protin amino aids
rtion of th plants. assiv volani onvulsions, oupld with th in- organi moluls (l
tns omardmnt from spa' gnratd surfa tmpraturs so hot that Millr ahivd h
th arth at this point ould vry wll hav had an ..On''of moltn rok simulat th intra
( i . . 'a m a g m a o a n ) . As an nrgy sour
Although tmpraturswould hv slowly drasdas th infali of jts sond-largstnrg
from spa and th irrtnsity of volani ruptions dlind, lvatd tmpr.
turs likly prsistdfor prhaps 100 milliln yars or longr aftr th for.
mation of th arth. During this priod tmpraturswould proaly hav
n too hot for organi omplunds to surviv. !ithout rgani om-
pounds, lif as w know it ould not hav istd. Howvr, on th asis of
data from nint zirons, y approimate| 4 illion years ago (or prhaps
vn arlir) th arth's surf must hav oold to th point that liquid
watr ould ist and gloal oans ould gin tl form (!(zildt al. 2001).
It was during this priod that organi ompounds would hv first startd to
umulat on th arth's surfa,as long as thr wr aioti prosssy
whih thy ould synthsiznd aumulat.!hat Was th natur of ths
prosss?
hmial rations. During ths Butlrov,who showd that th tratmnt of formaldhyd with alkalin at-
hav prvild. alysts'suh as lium hydroid, lads to th synthsisof a varity of sug-
ars.Th laoratory synthsisof iohmial ompounds was soon xtndd
to inlud mor ompl primntal sttings, and y th nd of th nin.
ritivarth tnthntury a larg amount of rsarh on organi synthsis had n pr-
formd (Bada nd Lazcano 2003).
iv on th arth's surfa that it is Howvr, ths nintnth-ntury organi synthss wr not onivd
..warm littl pond'' that Darwin mntiond
rni matrial. Howvr, during th as laoratorysimulations of th
.ormdsom 4.5 illion yafs to Hookr, ut rathr as attmpts to undrstand th autotrophi mhanisms
go'
unds prsnton its surfa aus of nitrogn assimilation and O. fiation in grn plants (s Bada andLaz-
tiv lmntshatd th intrior of ano 2002a).Th first onvining dmonstration of th possibl synthSisof
I roks (.sthrill1990). Volani organiompounds undr prioti onditions Was aomplishd in 1953 by
srhinggass out of th juvnil StanlyL. Millr, who invstigatdth ation of ltri dishargsating for
[n addition,th arly arth was also a wk on a mixtuf of Ha, NH,, LI2' and H,o; rami mixturs of svral
:simals,th dris lft ovr aftr th protinamino aids wr produd, as wll as hydroy aids, ura, and othr
onvulsions,oupld with th in- o r g a n i m o l u l s( i l l r 1 9 5 3 ' 1 9 5 5 ) .
.d surfa tmpraturs so hot that illr ahivd his rsults y mans of an apparatus in whih h ould
v had an .,oan''of moltn rok simulatth intration twnan atmosphrand an oan (sFigur 1).
As an nrgy sour Mill hos a spark disharg, onsidrd to b th
vly drasdas th infall of ojts sond-largst nrgysour' in th form of lightning and oronal dishargs,
ruptions dlind,lvatdtmpr-
illion yars or longr aftr th for-
:mpraturswould proaly hv
o s u r v i v .W i t h o u t o r g a n i o m -
istd.Howvr, on th asis of
ly4 illion yars ago (or prhaps
v oold to th point tht liquid
lgin to form (Wildt al.2001).
-rpoundswould hav first
startd to
; as thrwr aioti prosssy
[at.\ff/hatwas th natur of ths
\r-,0",
Flask
:r anaroihtrotrophsis asd
ompounds Wr a nssary pr-
)oratory synthsis of organi om-
.asfirst ahivdin th 1820s whn .r'
500-
' ould formd in high yild y Flask |10ml
n o n i a n d y h a t i n g m m o n i u m
Figur1. Th apparatususd in th 1953 illr xprimnt.Th 500 flask
y r o g n i z da s s u h , n w r i n was usdto rprsntth lansand th 5-1itrflask was usd to eprsntth
LolphStrkrahivd th laora- atmosphr. A spark disharggnratdarossth ltrodswith a Tsla oil,
lf ataldhyd,ammtrnia, and hy- invntdy Nikola Tsla in 1891' was usdto mimi lightningand orona
1 y th primntsof Alksandr dishargsin th atmosphr.
56 Th rigin of Lif
illr was al to transfornr alrost 50% oI th originl aron (in th form \ n,n
of mthan) into organi ompourrds.Although most of th syrrtlrsizdor-
gani matrial was an irrsolultarlik solid, h was al to isolt amino
aids and othr simpl organi ompounds from th ration mixtur.
Glyin, th simlst amino id, was produd tn 2Y" yild (asd on th R, R'= H or CnH21*1 :-l
original amount of mthan abon)' whras alanin' th simplst mino Retion 1. h Strker-
aid with a hiral ntr' showd a yild of \". illr was al to dmon- hydrlyaids.
strat that th alanin that was produd Was a ratni mixtur (qual
amounts of D- nd L-alnin). This providd onvining vidn that th
amino ids wr produd irr th xprimnt and wr not iologial orr- (O. + N, + H,o). In gn
taminants somhow introdud into the appratus. ward mor rduing ond
Th othr organi lmpoundsthat Millr was al tl idntify mad it pos- aids, purins, pyrimidirri
sil for him to propos a possil ration pathway for th amino aids nonrdun i g rmosphri
(illr 1957). Th proposd synthti mhanism ws in fat th on d- duing or nutral atmospt
sibd by Adolplr Strkr (18.50)whn h rportd th synthsisof alnin. nt mldl for th aly
his invlvs th ration of hydrogn yanid' amtnonia, arrdaonyl om- r|tdth id of nonr
pounds (aldhydso ktons) to fornr amino nitrils' whih th undrgo numr of primnts W(
hydrolysis to form th amino aids (sRtion 1). Dpnding on th on- modl atmosphers of (
ntration of mmoni in t ration mitur' vr1.ingmounts of hydroy found that not only wr
aids ar produd s wll. This is what illr found, with lgr rlativ glyin was asilly th o
amounts of hydroy ids ing forrd in a ration tnixtu ontining lss A gnral trnd showd t
amronia. mor nutral, th yilds o:
It is important to not that nithr purins nor pyrimidins, th nul- lly, although thy wr :
oass that ar part of DNA and RNA, w lookd fo iI-rth miturs of apprd to spially i
th original Millr_Ury xprinrnt. Howvr' in prirnts arrid out a m i n o i d s .T h m a i n p r .
soon aftr illr's prirnty oro and oworkrs' th formation of ad- o l o g i l I yr l v a n to r g a n i
nin fronr ammonium vanid solutions ws dmonstratd (016 1960; 016 a p p r n tI a k o f i o r m t i o
and Kimll 1961). Latr it was shown that th abioti synthsisf purins th Strkr pathway and
nd othr htroyli ompounds lso tks pla undr th sam ondi- of nuloss (Frris t i
tions as in th original illr-Uy prirnr,ut with muh snrallryilds high onntrations of r(
tharr for th amino aids. In additin' it has brrfourrd that guanin an b tions, and irr ths loaiiz'
produd in a dirt ..on-pot''synthsislrlm th polymrization of quous and ktons may hav
solr-rtionsof amronittm yanid (Yusa t al' 1984). ould hav om involv
AlthoLrgh ths rsults r xtrnrly nourging, th atrnosphri ompo- Whth th prinlitir'
sition that formd th asis of th illr.Ur1' primnt is not onsidrd vant with rspt to th r
today to plausil y mny rsrhrs.It is gnrally grd that fr howv. [t has ntly
oygn was absnt fom th primitiv rth, ut thr is no gnral agr- n i f i n ra m o u n t s o [ m i n ,
mnt on th onrposition of th primitiv tmospr;opinions vry from using O,/N'/liquid wat
Strongly rduing (o + N2, NI_l.r+ H,o, or o, + F{2 + N,) to nutral mino ids found prvr
r
'8 The rigi of Lif
produdin tlr rtions,as wll s hydrothrmal vnts r ruh mor fftivirr gulating th onntratiol-t
th lorv pH lrrditionsgnratddur- of ritialorgrrirrrtllulsitr th oans rhirIrirr pl1,irrg signifiantrtll in
1 aront to th system to uffr tlr thirdirts\,'nthsis.
dation inhiitossuh as asori aid Dspitth prolms with staility at higli tmpraturs,th vtr-itl]trd
:h rvryof svral hundrd tims svnthsis thor lrs n hampiond lry Hrrlrr and Wihtshiiusr(2006),
rusly. who hav dsridhav primnts limd to ..narrow th gap btrvn
rimnts,th amounts of fr yanid, iohmistry and volani gohrnistry.,' Usirrg high onntri-rtions
of HN
gstingthat th intrmdiats may (0.2M) and high pssursof Co (7.5ars) in tlr prsnof Ni/F.atlysts,
ids may hav fonrly svral mh- a varityof amino irrrlhydroyl aids ar prt>lul at 100.. Howvr, thr
or th rlatdBtrhrr-Brgspathway. ronrnsaor-rt t rlvanof this typ of primntto th natural gtl-
thir rltivaundnsar si-rilarto hmialnvironnlntstht w.ould ptd to hlrv istd ln th rlv
th slf.orrdnsation of HN in aqu- arth'spiallyth high onntrti()llsand prssursof strtir-rgrllgnts
1. 1978|, th asnof polymri ma- ( B a d a t a | . 2 0 0 7 ) .
:ardthis mhanism. primnts with
Shlsingrand illr 1983) suggst
r H, in nutral atmosphr simulations Did Priotiorgani ompounds om from Spa?
:tion of amino aids.
:iatdwith th dirt illr-Ury typ Th diffiultisstlpposdly involvd with th ndognous, rth-sdsl.rr-
rnt hypothsisfor th abioti synthe- thsisof mino aids nd nulossld to t dvlopmntof altrnatir's'
posd.This stiggstionsultd from In hrly 1990s lr1'airnlSgan (1992| nirlvzdor6's 1961 prlposerl
nal vnts.A group <lfsarhrshzrv on th rol of lrntrynuli as sours oi voltils ftlr th prinririv arth
.s of th hydrthrmal vnt nviron- and proposd tht th xognous dliv1,of <lganimattr y strlids,
rom th harsh onditions ausd y omts'and intrplntarydust partils (IDPs) ould hav plyd a signifi-
an imprtant ol in th origin of lif. antrol in sdingth arly arth with th lmpoundsnssryfor tlr ori-
ommon anstralorganism of all x- gin of lif. This onlusiln was sd on knowldg aout th ogani
svralrsarlrrshav proposd th ompositionof mtorits.It is important tO llot that if this onpt is virlid,
ls nssaryfor th origin of lif wr impatson th aly arth not only ratd dl.llstatingorrditionsthat ma]
o n s ( o l m n lA n d r s s o n1 9 9 5 ) . a . itdiffiultfor lif to originat ut also t th sanl tim prhaps dlivrlth
vrrrt shok nd or,l.orkrs,whr hav rawmatrialnssill.vto st th stagfor th lrigirlof lif. In n vn .ilr
quilirifvo th ormation of om- viwthis lrvpotlrsisould hav prlflr"rrrd irrrpliations|:orth irbtrrr{arr
of
rmal vnt tl-nprturs (Shok 1990; lifin th ttnivrs.Th sour of th ssntialorgani ompoun1srqird
;ynthsisis onsidrdto spially for th origirr of lif is not onstrind r'th onditions on a pz1rtiular
lff-axis systn1s(olm and harlou plant;rathr, svnthsisof organi ompottnds is r"rbiquitouspross llat
th high tmprturs(> 350") asso- takspia on primitiv solr-systmodis suh as astroids and otnts.
no ids nd otl-rriomoluls hav he possiilityof th origin of lif is thus tlnsidraly inrasd,providd
rnhardt t al. 1984; Millr and Bada thssntialrrrgniompounds ar dlivrd intt to a plant tht is suit-
rmino aids ar dstro1.d in timsals ablfor futhrhnrilvolution. This is t to fully drnonstatd.
n 300"' Th rt of hydrolysis for aronousllon]rits,a lass ()f st()nvlItOrits,ar rrnsidrdto
tmratursgivs a hlf-lif of ) mirr- th most prir-rritivljtsin tlr solzrr svstm in trms of tlrir lrrrrrtl
ry phosphodistond; at 350.C th omposition,yt thy hav a high urrdanlf lrgani aron. 1()rthan
Lalf-livsfor dpurination of ah nu- 3% by wiglrt in som ass. Th mtoritslTlosttnsivly analvzd for
s th hydrolsis rats for RNA (Millr organiompounds inlud th Ms urhison (r,vhihfll in 1969 tn Vitl-
l out y Lazcano (1997) that if th ori- ria,Australia)nd Murray (19s0, Kntuk1',Unitd Stats)and th I-lass
y long priod of tim' all th ompl orguil mtorit(1864, Frn). ( and I rfr to typs of hondrits.
vd from whatvr sours' would ar aout 709z"fin-graindmatrial and I ar alrnost lusivly fin-
othrmalvnts.It is thus possil tht graindmatrial' Both typs hv prin.ltrtsivaquOus ltrtit.ltl
60 Th rigi of Lif
rhasis dominatd y an insolul fra- monomrs.Th trm rm littl pod, wih lras long n usd for onv-
r solul organi matt is mad up of nin,rfrsn()t nssrilyto th ntir oa ut to parts of th hydros.
\Hs), lipatihydroarons' aroyli phrwhr th umulation rr.lintration of th produts of prioti
iotta zrrrdBada 2002). h purins ad- synthsismy lrv takrr plzr.lrs inlr"rdoani prtils and sdi-
Lnthirrhav also n dttd,as wll mnts'intrtidalzons, frsh,trshallow plrrdsand laks, lagoons ndr-
.ations<lf20
to 500 parts pr illion goingwt-and-dryyls, and utti nvironmnts assoitdwith glaial
on and Murray and in th I hondrit ponds.
1 9 8 1 ) .I n d d i t i o n , a v a r i t y o f o t h r Simplorgani lmpoundsdissolvd in th prirlritivons o[ othr od-
Llirdingpyidirrs,qirrolins,arrd iso- isof watr wlr'rldlrd to otlntratd sotn mhanislrl in ordr to
th lvIurhison mtorit (Stoks arrd nhanpolyrnizationprosses.onntr:rtioninvrlvingth sltivad-
: i d s ( p o l y o l s )( o o p r e a l . 2 0 0 1 ) a n d sorptionof molulsOnto minral surfashas n suggstdas on mns
l s ( D a m ra n d P a s h l y1 9 8 9 ) . of promotingpolvmrization' and this pross as n dmonstratdin l-
) mtOritssuh as Orguil ontain an oratoryprimntsthat hav usd a varity of simpl ompounds and ati_
m thar of tlr s (hrnfrund t al. vatdmonomrs (ill t l. 1998; Fris 2002). Tlr potntial importan of
'' onsistingof just glvin
nd rami minral-assistd atalysis is drnonstratdy rh montmorillonit-promotl
hondritsis intrsting aus it has polymrizationof ativatd adnosin and uridin drivativsthat produs
arity of amino aids wr rquird fo 25-50 mr oligonulotids(Frris 2002), th gnral lngth rang onsid.
howvr,among th andidats for th rdnssayfor primval iohmial funtilns.
id (PNA)' a nuli ;ridanalogLrin Bausadsoption onto surfrsinvolvs wk nonOvalnt van dr
n sugr or phosphat moitis (glrolm Walsintrations,th minrl-asdonntration pross and susqurrt
hPNA kon,ahiral amino aids polymrizatiorr wluld most ffiintat ool tmpraturs(Liu and orgl
n, possily dlivrd y I-typ ar- 1998;Sowryt al. 2001). This, howvr' prsntssomwhat of a prolm
'th,may hirvn
th lnly amino aids ausas th lr-rgth of polyms fomd on milral surfsitrrass,thy
tndto b mor fimly lund to tlr minral (orgl 7998)' In ordr for ths
polymrsto involvd in sr.rsqunt intrratilns with ltlrrpolymrs or
monomrs'thy nd to rlasd.This oul1 aomplishd y warming
sss th minral,although this would also tnd to hydrolyz th adsord poly-
mrs.A way round this prolm would to rlasth polyrnrs y on-
rth ftl lif xistd, rgadlss of its ntratdsalr sllr'rtions (Hill t al' 1998), a pross that ould t:rk pla in
r of diffrnttyps of sirnpl monomri tidalrgionsduring vaporzrtionor frzillg tlf swtr.
ry a mjor rol in iohmistry today. Th dirtnntrationof dilut soiutins lf monomrs ould also a-
Lstituntswr assmld into polymrs omplishdy vaporation and y utti frzing of dilut aquous solu.
s prsntlth most hallnging area of tions'whih, oupld with othr physiohmial mhanisms suh as th
adhrn of iolrmiallonotrrsto ativ sf;rs, ould lrv risd lo.
rth would nd to urrdrgopolymriza_ l onntr:rtionsnd protrrotd polyrrizzltion.h vllporation of tilal
l pross' It is gnrally assumd tlrat rgiona s n d t h s u s q u n t o n n t r e t i o n< l f t h i r o r g a n i O n s t i t u n r s
)0 monomriunits (20 o 100 mrs) ar havn proposd in th synthsis of a varity of simpl ogani mol-
: and rplition funtions (Joy 2002; u l s( N l s o n t a l . 2 0 0 0 ) . u t t i f r z i n go f d i l u t r a g n t s o l u t i o n s h a s
:ion prrsss must hav n apal of also n found to promot tlr synthsis of k,v iomoluls (Lvy t al.
rimum siz. 2 0 0 0 ) .T h s s a m O n l l t i l t i ( ) n p r o s s s l s o o u l . li r v p l a y d k y
y produd oligopptids or oligontr- rolin th synthsis of polyrrrrs.It hs n shown that tlr frzing of di-
ion retions larly must hav takn lut solutionsof ativatd mino aids at _20.C yilds pptids at highr
. Synon1.moustrms lik ..primordial amountsthan in primnts with highly onntratd solutions at 0o and
l . . h v l d i n s o m a s r s t t - tm a j l l r m i s - 25" (Liu and orgl 7997)' ln alditiln,rDtstudis hav slrown that u_
-i inr:rgof a worllwil oan, rilr in ttifrzingis spiallyfftivin th nor-rrrzymati syllthsisof oiignu-
nparrid y all sorrs of iohmi:rl lotids ( K a r r v r i o ti t a l . 2 0 0 1 ) .
62 Th rigi of I,if
Salt rins mv also hav rrimportnt irr tlr formtiorrof ppids:rnd dns monolayr of ioni'
prhaps othr important iopolymsas ,ll.Aording t<lRod and owork- of pyrit. ln addition,
rs (Rod 1999),..sit-indu] Pptid formirti<lnration'' (SIPF)provi1sa shusr amino aid_ppt
pathway for th ffiintsynthsisof pptids and possilv protins dirtly likly to hiv format
for simpl anrino aids in lnntratdNlll solutilnsotrtaining u(II). fftiv autoatalysis pr
Yilds of di- and tri;lptidsin tl-rrang of 0.4 to 47o l. n rpotdus- ory" (Ross 2008).
ing starting arrrinoaid onntrationsin th 40_50 m N,I ang. lay rl-riner. Non of ths xperill
als suh as montmorillonit aPParntly promot th rtion. Again, th aids ar th volutionar
vaporation of ti1allyflushd ltrgoonsr th frzingof th primitiv oans th rsults ar also oml
ould hav produd th onlltrirtdsirlt-rilrrins nd1 t()promot this primordial souP in whi'
salt-indud pol,vmrizatilt-l
ptlss. sour of ltrons for tI
bl that undr rtain g(
hav rdud not only
Did Pyrit Ply a Rol in th origins f Lif on arth? dp-savnts, and ld l
stan. ould hav takr
.tO?ihtrshlusr
Ft>rsom tim, one of th most srius rivals o th ,.primordial soup'' ht- and 19
rotrophi thory was drivd frm th ids of $7;ihtrshziusr (1988).A- hargs via a N4illr-typ
orling to his ..pionrmtaoli thory,'' lif gan with th apparanf ss did not rmain ol
n aLltoatalytitwo-dimnsiotr:rlhmlithorrophi mtalli systm that surrounding aquous n
took pla on th surfaof th highly insolul minral pvrit in th r,iiniry ioti soup, not of a tw(
of hydrothrmal vnts.Rpliation followd th apparanof this nonorgan. Th ssntial qustior
isml iron sulfid*asdtwo-dimnsi<lnal..lif,''in whih hmoautotrophi not whthr pyrit-mdi
aron fixation ttllk pla y a rdutiv iri aid yl, or rvrsrlrs CO, rdution nd syntl
vl' li rh typ origirrallvdsrid{or th photosnthtigrrrsirlfirrlra. pothtial two.dimnsi<
tIitlm hlorobium limiril' Molular pvlognti trs show that this Proof of Wihtrshius
mod of aron fixatilr-r and its modifiations (suh as th dutivatyl- tight oupling of th r
oA or th rdutiv malonyl.oA pathwa,vs)ar flund in anaroi zlr- t i o n v i a r d u t i v i t r i
ha irnd th mtlst dply divrgrrtutri:r'whih has n intrrtd of homologotls yls th
s vidn o[ its primitiv iratr(aln 1995). But is th rvrsKrs wll as rpliation (a
yl truly primodiirl? imntal sults ahivd
Th rtion FS + H,S = F.Sl+ H, is vry nrgtialli,favoral"It has and Co ar onsistnt !
an irrvrsil,highlv rgoni haratrwitlr a stndard fnrgyharrg th rol of slfur-rih
of (;., = -9.2 ka|lrol, whih orsptrdsto rdtrtion potntial ^ = mulation of organi or
-62m ). Thus th S/H,S omination is a strtlng rduing agnt nd has
shown to provid an ffiint sour tlf ltrons for th rdutiln of or-
gani ompounds undr mild onditions. Pyit formation an produ mo- How Did Rpliati
l t r l a rh y d r o g n : r r r dr d u i r r a t t o a m m < l n i aa, t y l n < l t h y l n ,a n d
thioti i.lttl ati ail, .rs wll as L)olrrotingmor ompl syDthss Th pimordial roth r
(adn 1995), inluding pptld lnds tht sult fronr th ativatio of drland in whih a wid
amino aids with aron monid and nikl/iron sulphids (Hur and sizd' dstroyd, or in
\/dlrtrshiusr1998). Although pyrit-mdiirtdO, rdLrtionto orgtri Rgardlss of th orp
ompour-rdshas not n ahivl,th fiatil undr plzrusilepi<lti ould not hav volvd
onditions of aron monoi]into ativat]ati aid l.lva mitur f o. guarant th maintnl
.Wihtrsh;ir-lsr
pripitatd NiS/FS/S has n rportd (u and Ponnts undr th at
1 9 9 8 ) . H o w v r , i r r t h s p r i m n t st h r i o n so t l r i t l n e q u o u s n . molular ntitis a]
viltlnrntto whih powdrd pr,rit has rrddd; thr, do not form a would hv markd tl
:
5
h rigi of Life o.)
otntin th formation lf pptids and dnsmonolayr of ionially ound moluls or tak pl on th surf
rs wll. Aording to Rod and owork- of pyrit.In addition' a arful aminatiorr of th Hur and !zihtr-
formatin rtion'' (SIPF)provides a shiusramino aid-pptid synthsisshm hs found that it is highly un.
pptidsand possily protins dirtly likly to ahiv formation of oligomrs that had th aility to promot
..pionrmtaoli th-
ltd Nal solutions ontaining u(II). fftiv utoatalysisprosssassoiatdwith th
g of 0.4 to 4o ae n rportd us- o r y " ( R o s s2 0 0 8 ) .
; in th 40_50m rang. lay minr- Non of thsxprimnts provs y itslf that oth nzyms and nuli
rtly promot th ration. Again, th aidsar th volutionary outom of surfa-ounddmtaolism. In fat,
o r t h f z i n go f t h p i m i t i v o a n 5 th rsults ar also omptibl with a mor gnral' modifid modl of th
salt.rihrinsnddto promot this primodial soup in whih pyit formation is rognizd as an important
sourof ltronsfor th rdution of organi ompounds. It is thus possi-
ltht undr rtain gologial onditions th FS/H,S ombination ould
havrdud not only o ut also O. rlasd from moltn mgn in
)riginsof Lif n arth? dp-savnts, and ld to iohmial monomfs. Pptid synthsis' for in-
stan,ould hav takn pla in an iron and nikl sulfid systm (Hur
rs rivals to th ..primordial soup'' ht- and Wihtrsh:iusr 1998) involving amino ids formd y ltri dis-
. i d a so f W i h t r s h i u s r ( . l 9 8 8 ) .A - hargsvia a Millr-typ synthsis.If th ompounds synthsizdy this pro.
lry,'' lif gan,ith th appran of ssdid not rmain ound to th pyrit surfa ut driftd away into th
molithotrophi mtoli systm that surroundingaquous nvironmnt, thn thy would om part of th pr-
insolulminral pyrit in th viinit iotisoup, not of a two-dimnsional autoatalyti ration shm.
owd th apparanof this nonorgan- Th ssntial qustion in diding twn ths two diffrnt thoris is
nal ..lif,''in whih hmoutotrophi not whthr pyrit-mdiatd organi synthsis an oLlr' ut whthr dirt
:tivitri aid yl, or rvrs Krs o, rdutionand synthsisof organi ompounds n ahivd by a hy-
Or th photosynthti grn sulfr.rra- pothtial two-dimnsional living systm that laks gnti information.
lar phyiogntitrs show that this Proof of !7zihtrshiusr's hyplthsisrquirs dmonstration not only of th
fiations(suh s th rdutiv atyl- tightoupling of th rations nSsryto driv autoatalyti O, assimila-
)athways)r found in anroi ar- tionvia a rdutivitri aid yl ut also of th intrwavingof a ntwork
:utria'whih has n intrprtd of homologousyls that, it is assumd,ld to all th anaoli pthways' as
Madn 1995\. But is th rvrs Krs wll as rpliation (adn 1995). This has not n ahivd.ln fat, pr-
imntalrsults ahivd so far with th FS/H,S omination in duing N.
, is vry nrgtiallyfavorl. It has and o ar onsistnt with a htrotrophi origin of lif tht aknowldgs
]trWith stndard fr nrgy hang throl of sulfur-rih minrals nd othr atalysts in th synthsisand au-
sponds to a rdution potntial ., = mulationof organi ompounds.
tion is a strongrduing agnt and has
:of ltronsfor th rdution of or-
ns. Pyrit formtion n produ mo- How Did Rpliation First originat?
l ammonia, atylnto thyln, and
ts promoting mor ompl synthss Th primordial roth must hav n a wildring organi hmial won-
rds that rsult from th ativation of drlandin whih a wid array of diffrnt moluls Wr onstantly synth-
rnd nikl/iron sulphids (Hur and sizd,dstroyd' or inorporatd into yls of hmial transformations.
t-mdiatdo, rdution to organi Rgardlssof th omplity of th prioti nvironmnt' lif as w know it
h fiation und plausil prioti ouldnot hav vlvd in th asnof a gnti rpliating mhanism to
:tivatdarti aid y a mitur of o- guarantth maintnan,staility' and divrsifiation of lif's asi om-
lortd (Hur and $7ihtrsh:iusr ponntsundr th ation of natural sltion. Th apparan of th first
th rationslur in aquOus n- molulr ntitis pal of rpliation, atalysis' and multipliation
.What
has n ddd: thy d<l not f()rm a would hav markd th orisin of oth lif and volution. wr th
64 The (.)rigiof Lif
irst molulr living ntitis that distin- th PNA-RNA and PNA-DNA hlis ar both right-handd. Although ap-
ry? palingas possilandidatsfor th first slf-rpliatingmolular living n-
rrs nd smali oligomrs to living nti- titis,oth PNA and TNA hav positiv and ngativ aspts. Th main
th uiquit<lus gntisystm of tant ngativfators ar th lak of ny dmonstratd oligomrization pross
; on of th major unsolvd prolms in that would produ ths nuli aid analgus undr plausil prioti
. of atalytialiytiv RNA moluls onditionsand th lak of any dmonstratd atalyti proprtis.
suggstionsthat th first living ntitis PNA is attrativ aus its akon is ahiral (lking handdnss),
noluls(riozynrs),in a hypothtial whih liminatsth nd for th sltion of hirality for th tim of th
d t h R N A w o r l d ( G i l r t1 9 8 6 ;J o y origin of lif. Its omponnts, AG and nuloass linkd to ati aid'
aptd,ut th limitd staility of th havbnprodud ndr simulatd prioti onditions (Nlson t al. 2000).
is molul was proaly not a dirt Howvr'PNA is stlsptilto an N-ayl migration ration that produs a
rarrangdPNA. This prolm ould minimizd, howvr, y loking
.ry unstal,whih maks its prsen thN-trminal position y tylation' for xampl.
l d t a l . 1 9 9 5 ) .B o t h l a d h y d r o i d on th asis of an tnsivstudy of sugar-asdnuli aids' TNA ap-
)parntlystailizrios (Riardo t al. pafsto b suprior to othr possibl sugar-asd nuli aids with rspt
at with riostl fom a stal iyli to its as-pairingattriuts,spiallywith RNA (shnmosr1999, 20041
Howvr, in ordr to involvd in th Shningt al. 2000). Th ttrossugar in TNA ould hav n synthsizd
l RNA, rios would likly hav to duringth ration asad that taks pla during th formos ration first
) pn t domposition. orovr, dsridy Butlrov in 1861. Th 4-aron sugars thros and rythros
r ombinations of drivativs of nul- ould hav n radily synthsizd y th dimrization of glyolaldhyd
ny hav n prsnt in th prioti (sRation2), whih in turn ould hav n produd from th dimriza-
molulapl of atalyzing its own tion f formaldhyd. vn though th prsn of a 4-aron sugr in TNA
Joy 2002). Ths diffiultis hav ld rdusthis prolm to 2 sugars and 4 stroisomrs' to dmonstrat how
whih informationl mromoluls oligonulotidsomposd only of L-thros ould prfrntially synth-
of tant nuli aids may also hav sizdundr prbioti nditions is a formidal hallng.
, that is, with phnotyp and gnotyp It is possil that PNA prdd TNA and in fat assistd in th transition
hus a simplr slf-rpliatrmust hav to th first TNA moluls. As statd arlir, th sltion of th hirl sugar
ndrshav n suggstd.Th RNA omponntof TNA would hav rquird som sort of sltion proess to
:y to atalvzrations and to stor in- in opration. orgl and oworkrs hav suggstd that th inorporation of
ruloass nd th akon that hld hiral sugar dinulotids at th nd of a PNA hain' whih ould hav o-
rrily th sam as thos in modrn RNA urdsimply y han, an indu hirality into a nuli aid produd y
polymrs and th atlyti :1gntstht PNA.indud oligomrization (Kozlov t al. 2000). PNA ould thus hav
unknown. providda mans of onvying th ritially imprtant iologial proprty f
rpls of molular gnti systms that hiralityinto polymrs nar th tim of th origin of lif. This possiility po-
nd thsprovid ampls of th typs tntiallysolvs an ongoing dilmma whthr th origin of hiality ourrd
givn ris to arly slf-rpliatingnti. forth origin of lif r during th volution of arly living ntitis.
idats inlud nuli aid analogus
s h n m o s r1 9 9 9 , 2 0 4 ; N l s o n t a l .
]NA), whr th akon onsists of
HCo'o?-Ho 9Sr91o Io2-FIoH.Co-CH2oH
N-(2-aminlthyl)glyin o (AG) (th tlo
id linkag to th amino group of
Formaldehyde Glyolaldhyd
IA), whr th akon is mad up of
,distrorrds.Bth of thsandidats
Reation2. Th formation of ttrossugars from th dimrization of
l !atson.rikas pairing with om- formaldhyd and glyolaldhyd.This is nly part of the ovrll formos sris
jA, and DNA. With PNA-PNA hlis and
of reations, ot hxossnd pntosswould hav n produd y
ar produd in qual amounts' whil subsquent rations.
h ()rigitt f 1,if
Rgardlssof th typ of analogu simiIar to nuli aid or othr typ of th asi mahinry
rpliator sstm that ws usd y tl-rfirst self-rpliatingntitis, polvmr .
partmntalizd into
staility and survival would hav n of ritial importan. Nuli aidsin Dspit th susl
..with
grrralhv vry shrtsurvival tims at lvatdtllpraturs. RNA is r,r lms with this
unstabl ausof th prsnof th phlsphldist ond involvirtgth 2'- ordr to maintain a s
ydroyl group of ris. Th staiIiryof TNA is unkrrorvnat prsnt' ut it ating systm insid
is proaly somwht similar to th staility of DNA. It is possiltlrat otlrr layr barrir. This
ttros sugar-asdnuli aids wr lss stal thirn TNA usof thir as amino aids. As a
tndny ttl for lss stal hlial stfrrtul]s(snmtlsr|L)99|rrd tlrt sid th vsil'' (Gr
th sltion of TNA Was th rsult of this fatlr,althotlgh this is also not surfs ould hav
known. Tlr stility of PNA lrs n prtly invstigatd,and providd th ules that usd th z
N-ayl migration ration an rninimizd, th mid linkag in PNA volvd in polymriz
rvould xptdto hav a stailit at nutral pH similar to that of pptid omplxity, synrgis
honds in pr<ltins(.Wng1998). This suggststht in nr,ir<lnmnts rvith tm- of th omponnts il
pratursof around 25", its survival tim wotrld in th rang of 104-105 portd into th vsi
,vars.Salry'rins trray lrav playd a rol in ariy nuli aid suvival - formd a singl om
aus high salt onntrations hav n found to nhn nuli aid st- rsults showing that
bility. For mpl, th staiiitis lf svral transfr RNAs (tRNAs) wr om polymrizd
signifiantl.vinrasdin |-2 Nl soluti<lnin omplrison rvith staility pidi arri may no
in pur water (Thit al. 2002). Th staility of DNA also inrasswith in-
rasil-lgsalt onntration (fo ampl, sBda 2004).If thsrsultsr,ver
ppliale to othr nr'rliaid analogus suh as PNA nd TNA, thrrsalt Th Transition l
solutions ould hav providd a prortiv nviront-trnt that ould hav n-
h:rndth survival oi arl1'slf-plitingmolular ntitis. Th volution of th
-h
$rmmrns rquird for th mt:gnf rpliating systms? tirly on RNA (i..
mrgnof lif m-v st irndrstoodin trt-t-ts of th dynaIlis and vo- volution toward m<
lution of sts of hmial rpliirtingntitis.\7hthr suh ntitis Wrn- on molul that
..t
llsdwithin mmrns is not yt lar, ut givn rh priori availallity Laoatory-basd
of amphiphili ompounds, this rn,vhav wll n th s. Mmrn- atalyti RNA mol
forming lipidi ompounds may hav n providd to th primitiv rth rang of important
from xtrtrstrials()Llrs(Dmr and Pashl 1989). This sotlr ould RNA fragmnts (lig
potntially supply a wid varity of aliphati and aromati hydroarons,al- 1999|. Th list of dt
lhols, and r;rnhd arrd straight fattv aids, inluding som that ar that th RNA world
nrman-formingompunds. Prioti lipidi moluls ma1,lso hav r. uls, prhaps fun
sultd frorlr aioti synthsis, ars shtlwn y th formatiln rrf nornral ftty 2002). Th omplx
ids, glyrol. glyl phosphat, and tlrhr tipids (iherg t a|. t977\. ti transfr, nd vaI
Also, primntshav shown that simpl miturs of fatty aids and glyrol dmonstratd in th
form fro.l miturs <lfmt>no.,di-' nd triglridsur-rdrild lnditions immatuity of tlris
(Hargravst l. 1977)' It is vry atrrativtO assul tht orpartmntal- Ch 2002; Joe2|
ization within liposot-llsformd y amphiphili molulsof prioti origin Th invntion of
\vs ssntiillfor th mrgnof lif. RNA nr<lltrls lsordorlt las hiny ndd for l
suh as montmorillonit' whih an atalyZ th formation <lf RNA Four of th bsi r
oligoms, an e napsulatdinto fattl, aid vsilsrvhos forlltion in ozyms, and it has
turn is alratd y th lay. By inorporating additiorral fatty aid rni- tions is not likly a
lls' ths vsils an gow and divid whil still rtaining a portiln o{ oigin, most pra
tlrir ontnts ndd o sLlpprt RNA rplition. In this mannr' som of th as' thn th r
Th rigin of Life 67
similar to nuli ail or othr typ of thasimahinndd for RNA slf-rpliatilnould hv Irom-
6rst slf-pliringntitis' p<llyrrrr prtmntalizd irrto poto-typils (rizy t :rl.200; hn 2006).
lfritial imp<lrtarr. Nuli aids in Dspitth sussof th RNA napsulatioIrprimnts,thr ar pro-
at lvatdtmpratus.RNA is vy ..rvithin
lmswith this th vsil'' snario (Griffiths 2007). For mpl, in
: phosphodistrond involving th 2.- ordrto maintain a stdy supply of moluls ndd for survival of a rpli-
g of TNA is unknowtl t prsnt, ut it atingsystminsid a vsil, moluls would nd to ross a lipid-lik i-
ailitv of DNA. It is possil that t>thr layrarrir'This rrlay hav n mirjor ostal fo hargd spis suh
lss staltharrNA ausof rhir ..out-
as aminoaids.As a rsult' prlrirpsth first rpliating systm volvd
rllturs(shnmosr1999) and tlrat sid th vsil'' (Griffiths 2007). In this as, vsils attahd on minral
lf this fator, altough this is also not surfasould hv n in los proimity to sirnpl slf-rpliating mol-
l ptly invstigatd,and providd th ulsthat usd th atalyti proprtisof th minrl to nhan rations in-
'inirnizd' th amid lirrkag in PNA volvdin polymrizationand rplition. As thstlr two systmsvolvd in
tt nutrlpH sinrilar to that of pptid omplity,synrgistiintrirtiotrsould irav dvlod thar lllwd som
uggststhat in nvionmntswith tm- of thomponntsinvolvd in th rpliating systm to om frly trans-
til would in th rang of 101_105 portdinto th r,sil.At this point th two systms ould hav rrrrgdand
rol in arly nuli aid survival - formda singl omartmntalizd, slf-rpliation systm. Howvr, rnt
n found to nhan nuli aid st- rsultsshowing tht ativatdnulotidesan diffrrsinto liposoms and -
svrltransfr RNAs (tRNAs) wr ompolymrizdwithin thm suggsttht undr rtain onditios tl-rli-
solution in ompzrris<ln with sterility pidibarrirmy not hav n irnpntral(nsy t al. 2008).
;tirbiIityof DNA also inrasswith in-
I 's B d 2 0 4 l . l ft h s r s u i sw r
gussuh as PNA and TNA, thn salt Th Transition toward a DNA/RNA/Protin World
tiv nvironmnttl]t ould hav err-
rting molulrnritis. Th voiutionof th first livirrg rrrolularlir,ing rrtitisinto ons basd n-
:mrgnof rpliating systms? Th tirlyon RNA (i..,th RNA rvrld) would lrav n th nt stp in th
lod in trms of th dynamis and vo- volutiontoward modrn ihrnistry.RNA has n found to an all-in-
n t i t i s .W h t h r s u l l n t i t i sW r n - onmolulthat an not only stor information ut also atalyz rations.
:ar, ut givn th prioti vailaility ..tst-tu-volution''primtrtshav dmonstatdthat
Laoratory-asd
hv wll n th ils. mran- atlytiRNA moluls (riozyrns) hav th pitv to ary Out a wid
n providd to th primitiv arth rangof irportant iohnrial ations,irrluding th joining togthr of
n d P a s h l y1 9 8 9 ) .T h i s s o u r o u l d RNA fragmnts (ligation) and pptid-ond formation (Bartl and Unrau
phati and romati hydroaons, l- 1999).Telist of dmonstratdatalyti rations is tnsiv ad suggsts
[amy aids' inludig som that ar thatth RNA world ould hav had a lrg rprtoirof atalyti RNA mol.
>tlliidi molr-rls mv als<lave r- uls,prhps fr"rtining in onrt with on an()thr(Doudna irnd Ch
vrr y th fornl:rtitlt-tof ntlrrrlal fatty 2002).Th otrrplsrisof rtionsnddto prr-r.rit rnultiplition'gn-
L d o t h r l i p i d s ( i h r ge t a l . 1 9 7 7 ) . ti transfr,and variation rqr.lirdin th RNA wrld has so filr not n
pl miturs of fatty ids and glyrol dmonstratd in th laoratlry,ut optimism rlrrainsbausof th rlativ
rd triglyridsundr mild tlnditions immaturityof tis ara of rsrh (Bartl nd Unrau 1999; Doudna and
ativ to assumthat ompartmntal- h2002;Joy 2002).
p h i p h i l i m o l u l so f p i o t i o r i g i n Th invntion lf protin synthsisand th npsulation of rtioll ma-
RNA molulsdsod onto lilys hinryndd for rpliation n1y have takn p[ during th RNA world.
rr atalyze th f<lr:mationof RNA Fourofth asi ratitlnsinvolvd in protin ilsynthsisar atlyzdy ri-
ltty aid vsilswhos formation in ozyms,and it has n notd that th omplmntary natur of ths ra-
otporating additional fatty aid mi- tions is not liklv aidntal ut rathr suggstiv that thy had a ommon
lid whil still rtaining a portion of origin,most proably in th RNA wlrld (Kumar atr]Yarus 2001)' If this was
. rpliation.In tliis mannr, sonr of thas,thrrtlr primitiv nltloasod usd tor protin iosyntlrsislrad
Th rigi lf l,if
its origin in th RNA worll, lthough th b:rssusd in th early od ould Prthrmophili organt
hv en diffrrrtforn tll ons usJtody (Kol t l. 1994). modrn iology was a
It is possil tht y th tim RNA-asd lif appad<lnarth, th sup- DNA/protin-asd lift
plis of simpl ioti orgalli omportndsdrivd from th soursdisussd At first glan, oth
arlir would hav n grtly lirinishd.nr. of th lrponntsof th PPr to support a }
primordial soup had likly n tnsivl1.onvrtd into poiymrs, inlud- arly. Larg-sal anal
irrgthos ssoitdr,r,ithliving rrtitis,nd thtrsth rrv nl:ltrialsnddt of primitiv arth was
sustain lif had om largly hr.rstd. This inlplis tht simpl mtoli- rrmaind moltn for
lik pthwys must hav rrin plir in ordr t() nsur a supply of th in- (Nfthrill 1990), but n
grdintsnddto sustain th istnof th primitiv living ntitis.In this hydrosphr implis th
as som mtrllipathways ndd to produ sslltial omPonnts Howvr, thr is th
rquird y primitiv living ntitisWr prhaps origirrallynonnzynllttior wnt lat rtion im.
sminzymatiautoatalyti prosssthat ltr brfin tund as iozy- may hav oild th o
mati nd protir.r-sd nzmti poessing rltlmi rrnt. Could modrn bioh
ore than with rlier living rnolulrntitis,th main limittion in th ditions? Th proposals
RNA rvrld would hv n th xtrm instllilitv of RNA. This i turn lms, inluding th h
implis that RNA m<llulsmust hav n vry effiint in arrying out ompounds suh as ar
slf-lpliatiOnationsin ord tl t-nintainarr adquat invntory of mol- moluls, whos ha]
uls ndd ftrr survival. Th instiIity of RNA otlld hv n th pri- 250_350. ar at th n
mary rson ftl th transition to th DNA/prtltin world, tvhr, usof It is tru, of ours,
rh inrsdstility of th gntimoluls,srrrvivalwould hr,nlss fstr. Primitiv nzyI
dpndnton polym staility.Aording rlJoy (2002)' it is possilthat nd thus th rat nha:
irr th RNA rvorlliozyms ros tht ould talyz th pllyrnriztionof on way to ovrome
DNA, nd in this mannr information stord in RNA ould tansfrdto rizd lswhr (Islasi
th mor stal DNA. Anothr rsotl for DNA takovr ould hav n
1. rdud onntr
that aus of inrsd staility muh longr oligomrs ould hav au-
and NH.;
r-rruiatd.his rvould hav providd n nlrandstor:rgapaity fo infor-
2. lowr Stady.stat
mation that ould passd on to th nt gnraion lf living niris.
whih at tmpra
Befor long, RNA, whih had on playd tlr singular rol tlf rpliatitrnand
frmamid and tl
atalysis,ws rplad y th mlffiintnd rousr DNA/prtin .orld,
3. instability of ra
rvhrinRNA wrrsdrnltdto a rol of mssngr/transrir of DNA-stord (RFIo(Nt{,)
irrformtion ndd for rtltin iosvnthsis.
o f a m i n o a i d s ;a
4. loss of organi
s t a i l i t yo f g n t
Did Lif Aris in a High-Tmpraturnvironmnt?
Th rognition thz
vn tlrouglrDNA is nror st:rlthan RNA, it is still ridly dgrddat l- ar oupid y hyp
r,atd tmpraturs.In addition, Protin nzyms dnatur rapidly at lvatd itslf onlusiv pro
tmpraturs.This rtrsthr,at lst irtitillylimitd th nr,ironmntswh lif (Brohir and Phi
DNA/protin-asd lif ould surviv for anr. signifizrntprid of tim, nd as sriptions of th vo
was th as for ll othr arli living ntitissd tln nuli aid, sr.lrvivl biohmia1ompoul
would hav n most likly und ool onditiorrs(Bda and [,azno2002). bly niv to attmpt
Nvrthlss, sevral rsarhrslrav advoatd high t1praturs,spill living systms from
thos zrssoiatdrvith [rydrlthrnralvrrt systnrs,as th nr,ironmnt whr rl gntransfrof.
DNA/protin_asd ntitis first ros. Prlponntsfor high.tmprturOrt- misd th gnti r
gin it th f:tttht th trnivrsltr lf tI1tlif irpprst<l[.l<ltdin hv- onlusions aout th.
Tbe rigi of Li|. 69
r th assusd in th arly od oLlld prthrmophiliorganisms. Thus if th lst om]on anstor (LA) of all
ltody (Kolt aI.1994). modrnilogy was a hyprthrmphil, thn it is onluded that tlr first
-sdlif irppardOn art' th sup- DNA/protin_asdlif nrust hav arisn in a similar typ of nvironmnt.
rnds drivdfrom t sours disussd At first glan' oth th molular and th palontologial fossil rords
ishd.:rrrylf th omporrnts of th apparto support a lryprthrrophiliorigin of lif. Lit on arth aros
sivly onvrtdintrl polymrs, inlud- arly.Larg-sal analysis suggsts that son aftr its formation th surfa
s, and thus th rw mtrials nedd to of primitivrth was xtrml hot. Th plant is gnrlly thouglrt to hav
td.This irrrplisthat simpl mtoli- rmaindmoltn for som tim aftr its formation 4.6 illion yars ago
in ordr tO nsur a supply of th in- (Wthrill 1990|, ut minralogial vidneof a 4'- to 4'4-illion- yar-old
oi th prinlitivlir.ingntitis.In this hydrosphr lmplis that its surfa apidly oold down (\r'ild t al. 2001,).
id to produ ssntial tlnrponnts Howvr,thr is thortial and mpirial vidn that th plant und-
- ' r 1 . l r l i pt lqr i g i n l I l r n o n n z v m a t i o r Wntlat artion impts (Byrly et a|.202; Sl-ronrg t al. 2002) that
rlratiatr amfin urrds riozy- mayhavoild th oans as lat as 3.8 illion yas ago (Slp t aL.|989).
ossingamdominnt. ould modrn ihmistry hav arisn undr ths high tmprtur on-
ul ntities,th main limittin irr th ditions?Th proposls of a high-tmprtur origin of lif fa major pro-
rrninstaility of RNA. Tlris in tr'rrn lms'inluding tlr hnrial domposition of prsumd ssntial iohmial
r v n v r y f i i n ri n r r y i n g < l u t ompoundssuh as amino aids, nuloass, RNA, and othr thrmllail
rintainan lquatitlvntorr.of mol- moluls,whos half-livs for domposition at tmPraturs twn
'ility of RNA ould hav n th pi- 250-350"ar at th most a fw minuts (!(/hit1984;Millr and Bada 1988).
)NA/potirr world, whr, aus of It is tru' of ours' tht high tmpratursllow hmial rations to go
oluls,sul,ir'aIlr'luldh:rl'lln lss fasrr.Pimitiv nzyms' on tlry appared, ould hv n inffiint
dirrgto.|oy(2002), it is p<lssiltlrat ndthus th rat nhanmnt ssoiatd with highr tmpraturs wouid b
lt ould t|yzth polvmization of onWay to ovrom this limitation (Harvy 1924), Howvr, s summa-
stord in RNA lr"rld trzlstrrdto rizdlswhr(Islaset a|.207)' high-tmpraturrgil-tls would lad to:
n for DNA takovr ould hav n
1.rdudonntratiOnsof volatil intrmdiats'suh as HN, H2O,
rh long oligomrs ould v tt-
and NH,;
n nhan{ storagapaitr,for infor_
h nt gnration of living ntitis. 2. lowr stady-statOnntrationsof prbiti prursors lik HN'
yd th singulirrrllof rplition nd whih at tmpratursa littl aov 100. undrgoshydrolysis to first
forrnamidnd thn formi id arrd ammoni;
intand roust DNA/prorin world,
{ mssngr/transritrr 3. instabilityof rativ hmial intrnrdiatslik amino nitrils
of DNA-stord
(RHo(NH')N), whih play a ntal rol in th Strkr syr-rthsis
rhsis.
of amino aids; and
4. loss of rgani ompounds y thrmal domposition and diminishd
stailityof gnti polymrs.
rturnvironmnt?
Th rognitionthat th dpstranhs in rootd univrsal phylognis
RNA, it is still rapidl1'dgraldat l- aroupidy lryprthrnophilsis ontrovrsil and dos not provid y
n nzymsdnturapidly t lvatd itslfonlusivproof of a higlr.tmpraturorigin of DNA/protin-asd
ritially limitd th nvironmntswhr lif(Brohirand hilipp 2002). Givn th hug gap xisting in urrnt d-
l any signifintpriod of til, and s sriptionsof th voltrtionarytransition twn th prioti syr-rthsis of
ntitisasdon nr.rliaid, survival iohmialorrrpounds and th LA lf ll tarrt living ings, it is pro-
onditions(Badaand I,zno20O2). ly naiv to attrmpt to dsri th origin of lif nd th natur of th first
dvotdlriglr tnlpraturs,spi:rlly livingsvstemsfronr nrolularplrylognis(Islaset aI. 2007).In addition, lt-
nt systms'as th nvironmnt whr ralgntransfr of thrmoadaptativtrits has apparntly grtly ompro-
P r o p n n t s o rl h i g , h - t m p t urrlr i - misdth gnti rord prsnt in modrn organisms, r,vhih maks any
tnt lit appars to ro<rtdin lry- olusions aot th nvironnrntwhr th DNA/protin world origirrtd
70 hc rigilt of I'if
B r r a t l . 2 0 0 7 ) . A n a n a l y s i s o f tantforms of lif ut also imply that th sts of gns that nod th om-
: nzm'[vrsg},rs'that is sifi ponntsof ths omplx traits wr frozen a long tim ago (i.e., major
]r apprntlynot nstralto ths or- hangsin thm r vry stongly sltdagainst and ar lthal). It is tru
J-adaptdvrsions of thos prsnt in that no anint inipint Stagsor volutionary intrmdiats of ths mol-
r 2 0 0 2 | . v n i f t l r L A w a s a h y p r - ular struturshav n dttd, ut th istn of gradd intrmdiats
,uyand haussilorr2008), thr ar ill- an b ddud'and thrfor th suprnatural origin advoatd y oth old-
l distriution,inluding th possiility fashiondand ntmporary rtionists is rndrd unnssary.
hanhigh-trnpratur rgimsthat may Fo instan,th fat tht RNA moluls y thmslvsar apl of pr-
l (Slpt al. 1989; Gogrrl-r-Bokels formingll th rtions involvd in pptid_ondformtiorr suggsts's statd
ia to trm nr,ionmntsy latrl arlir,that protin biosynthsis volvd in an RNA world (Zhang and h
al. 2000) and otr thrmoadptativ 1998|,that' is, that th first iosom lakd protins and ws produd only by
(3) outornptionof oldr nrsophils RNA' This possiility is supportd y rystllogphi data that hav shown
)tdto strss-indr-ring onditi<lnsothr thatth ribosom'satalyti sit, wlrr pptid-bond formation taks pla, is
a z a n o! 9 9 5 \ . omposdof RNA (Ban t al. 2000; Nissn t al. 2000). lursto th gnti or-
ganizationof primitiv forms of translation ar also providd y paralogous
gns,whih ar squnsthat divrg, not through spition, ut ftr a du-
nthsisto th Last pliationvnt.For instan' th prsnin all known lls of pairs of homolo-
gous gns that nod two longation fatrs, whih r GTP (guanosin
triphosphat)-dpndnt nzyms that assist in protin iosynthsis, provids
oaly RNA-asd lif as wll, anr vidnof th xistnof a mo primitiv, lss rgulatd vrsion of protin
tln, th way aron was produd and synthsisthat took pla with only on longation fator. In fat, th xpri-
s forvrhangd,spilly rftrpho- mntalvidnof in vitro tanslation systms with modifid ationi onn-
organi aron prdud y arly lif trations lking both longation fators and othr protin omponnts
Ho, and simpl organi tlmpounds (Gavrilovaet aI' I976; Spirin 1986) strongly supports th possiility of an oldr
] hav f ddth amoutts of or- nstralprtin synthsisapparatus for th mrgnof longation fators.
irrgsynthsizl'fom ithr horn. Bioinformati nlysis of sqund llular gnoms from th thr major
ours undr th st onditions. Th domainsan usd to dfin th st of th most onsrvd protin.noding
on th rth thn shiftd frorn on ini. squnes to haratriz th gn omplmnt of th LCA of xtant lif. Th
aioti origin to t>nmad tlp ntirly rsultingst is dominatd y diffrnt putativ ATPass' and y moluls in.
volvdin gnprssion and RNA nrtaolisr.Th so-alld DAD gyrass'
ltin world that iohmial mhinry whih unwind RNA moluls and lav thm rady for dgradation, ar s
ran strttturs'omparal to thos onsrvds many transription and translation gns.This suggststh arly
sprd.This ls tlrus th first tim that volutionof a ontrol rnhanism for gn xprssion at th RNA lvl, pro-
righthav nprsrvdin th form of vidingdditional support to th hypothsis that RNA moluls playd a mor
lir stagswould hv lft .hind only prominntrl during arly llular volution. onsrvd squnslatd to
ot sttrvir.dth rar':rgsof gohmil iosynthtipathwys inlud thos noding putativ phosphoriosyl py_
: tim. Although tras of hydroaron rophosphatsynthas and thiordoin, whilr partiipat in nulotid m-
billion-r.ar-oldsdimntaryroks, th taolism. Although th information ontaind in th availal dataass
s found tl dt (Broks et tl|. 1999), orrspondsonly to minor portion of biologial divrsity, th squnsre-
lation,nd post-dpositionalmtamor. portdhr ar likly to part lf an ssntil and highly onsrvd pool of
rlir rord of ioti organi hm. protindomains ommon to all orgarrisms (Brra et a|' 2007a).
.ithprimitivlif. Th high lvls of gnti rdundany dttdin all sqund]noms
l sslltialfattlrsof gn<lmrpli. implyot only tht dupliation has plyd a major rll in th artion of
) rati()ns'nd mmrn-assiatd th ompl gnoms fond in tant lls, ut also tlrat for th arly
h molui:rrdtails of ths univrsl dupliationvnts rvald y th larg protin familis, aus th duplia'
lnof th mlnlph1.lti origin of all tion is usd to uild mor mplity, simplr living systms xistd that
72 Th rigi of Lif
lutionary transitiorr twn th prioti synthsis of iohmial om- Burkhadt' F., and S. Smi
rlas Dri, 182
pounds and th last ommon anstorof ll tirntliving ings' ut attmpts
Univsitl' Prss.
to rdu it hav llowd mor pris dsription of th ginning of lif,
Bvrlv' . R.' ). R. Lorr,
Nowadays ntral issu in origin-f-litrsarh is to undrstnd th abioti l:r}'fnl th Pilr
synthsisof an nestrlgntipolymr ndowd with atalyti ativity and its C a l r , i n ,M . l 9 6 9 . h m i ,
furthr volution to an RNA world and ultimatly to th DNA/RNA/protin LiL,ig Sstms o tb
world haratristi of all lif on arth. ! fa major unsolvd prolms, but C s t r s a n a ,J . 2 0 0 1 . o l
Bilpbsi t 150
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fossilsand th rlyrisf ukarvotes. Si285: 1033-1036. F i s ,J . P . 2 0 0 f . o n t
h ()rigi l| Lif 7.t
oti synthsisof iohmial om- Burkhardt,F', nd S. Smith, ]s.|994 . ldr of tbc tlrrsprlltdtlf
all tantliving ings,ut attmpts hrlsDru'i, 1821-1 I]82, \Yith Supltt. ztt|lridg:()rridg
UnivrsitPrss.
: dsription of th ginning of lif. .Woodll,
Bvrlv,G. R., D. R. l-lw,J. L. and . i. 2002. An Arhan irrrpt
sarhis to undrstand th ioti
lafrorrrth il nd K;rpvaIr.tolls.Si 297: |32.5_127.
lndowdwith atlyti ativiy and its alvin' . 1969. hmil u<irttiott:Mlttlr F'ulitttitl t(')|Urdsth rigi tlf
ultimatly to th DNA/RNA/protin Liuing Sstmso th rth d F'Ishr.oxflrd: C]l:rrlldr>n Prss.
/ fa ma|or unsolvd prlms, ut strsna,J' 2001. ornparativt.gllolllis and lrionrgtis.IJirlt-llimit
ry' unsolval,or unknowabl. Biop|lsir 1.506:|47_162.
hn,I. A. 2006. Th mrgof lls 1uringtl.rtlrigin of |if.Si 374
1558-1-5-59.
hvba,., d . Sgn. l992. rrdgnOusprodutioll. Ogl)tlllsdlivrv rrrrd
impashoksynthsisof organi loluls:An ilrvntOrVfl th origins lf
n d B ' S . a m 2. 0 0 7 . . 4 3 i l l i o n - |ife.Ntt,tr 355: 12.5_12.
Austrlia: osystm-sal insightsto i v sH, . . l . ' J . . h l r r s . . [ , a z n t lS' . L . N , I i l l ra, r l d J . L . t ] d . 2 0 0 8 .
h 158 198-227. rassssmnt of pritltiorg:rrrisynthsisin rrutrlp1antirvetmlsplrrs.
statusrort.rth PItrSienc rigisof Lif d urllttti,f th Birlsphr 8 l0.5_11.5.
ooper,C., N. Kimmih, W. Blisl'.].Srilrr.rlr' K. Brelrarrr,aIll L. GrrI.2001.
4. Impatmltingof froznoanson aronaousrtoritss a sour of sugr-rlirtl lrgrrilmpouridsfor
in of lif. Prlt:digs
of th Ntiol th rly ea|t|1. Ndtur 4l4: 879_88.
;0. Dahm,R. 200.5.Frilrihishr nrl tlr disovrv f D.A. Dcrll1lllttll
o, H.J. lavs,R. . Hazn,nd Blolo3r'2-8:2-4-28.
rr thoriginof lif on eart.Sir Darwln,F. ]887. hr Lif d Lttrs rlf hrls l)ui, ]lttdis lt
ut o bi ogr 1lh i l (',hp tr, Lorrdoll : M urrir1.'
:vldnw waysto studyth rigins of Damr,D. W., arrlR. M. l)irshly.l9tt9. Arnlriphili tlnlpurrlslf tlr
urhison aronaoushondrit: SurftrropItisan1mrr:rtr
firstimlu|es.
Sie296: formation.()rigis of Lif d utlltion rl|-thBirlsllr 19 21_8.
d Duv,. l995. \/itlDLrst:I-it'. s smi Imprtit'c.N, York: Birsi Books.
illr primnt.
Si3:74 5-7 46. -. 200. A rsrhproosl o th ()riginof |if. rigills o| I-ifc d
Th stilityof aminoidsat Luolullo1l | I 1 5 l | ) s D l J , 1 '. i i : ) . ) y _ ) , / Z t .
.l
955. Produtiln of som <rrganiompounds undr possilrimitiv S l p ,N . . , K . J . Z a h r c
arth onditions. llrl lf th mric hmil Soit 77:251_261 . osystls by lrg ar
19 57 . The mhnism of synthsislf amino aids y ltri dishags. S o w r l ' y .s . . I . ' . M . r
Biohimi t Bklphsi t 23 48()489. dsptior.rof adnin
1997. Peptid nuli aids and prioti hmistr. Ntur Strutrl Spirin,A. s. l986. Rlbos
B i o l o g 4 | | 7 - | 6 9 . Bn jamin/ummings'
illr, S. L., and J. L. Bada. 1988. Sumrin hlt springs and th origin of lif. Springstn, G., and (]. F.
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RNA may hv n th first gnti molul. Prodigs of th NtioI l982. Bsinitrog
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of riosom tivity in pptid ond synthsis.Si 289 92_93. Glyooll homlllg
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th natur of trial innovtion. Nture 405 299_304. 2002. Th sarh foi
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Bklphsil Rsrh ommuitios 2: 407_412. i, . M., r.rdD. R. l-r
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primitiv eart|1.Ntr 19 442-44. Uno, Y., K. Yilml,N.
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onditions. I. Adnin from hydogn ynid. rhius of Bhmistr d Ntltr 440:5l6-.5 l9
tsiopbsis94 217-227' van Zuiln, N4. A.' A. Lp
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76-82. R. H. 1984' ydr
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isotop vidnfrom 3.8-Gyr mtmorphosdsdimntsfor arly mt()rit \}/ild, S. A.' J. \W.Vlly,
omardmnt of th eaffh, N(ltur 418: 403-405. dtritl zirons fo th
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Shopf, J. \).199. irofossils of th arly Arhn Ap hrt: Nw vidn llleailr ullutiolt
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Shk, . L,. |990. Gohmil onstraintson th origin of orgni ompounds in rilzyms. Ntur 39
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Th rigi of Life 79
80
Plotolrg1'
d th istlr of Lif 8,l
Narativ
.hr
r many ways tO r()untth strrytlf th history of lif. As <lnof
manytypialapls,Bnton anl Harpr (|997,2008) psntd squn
8. Primt socitis, bum soitis, d th origi of lguge. The yars' thn living sp
disiv stp in th trnsition from ap to human soity was proly th past 600 million
th origin tlf languag.In many wys human languag is lik th gn- 2. Th sond argumn
ti od; information is stord and transmittd,with modifiatron, past 450 million ya
down th gnrations.ommuniation holds soitistosthr and 1 0 m i l l i o n y a r s ,a n d
allows hunrans to sapvolution. trrstrial spis tha
aynard Smith and Szthmiry (1995) argu that all ut two of ths ight 3 . h t h i d r g u m n ri
transitions wr uniqu, ourring just on in a singl linag.Th twl - s p i so f o s s i l m r i
ptions ar th oigir-rsof multillular organisms, whih happnd thr numr of known m;
tims, and of olonial animals with stril asts'whih has happnd many Allowing for nonprs
losss,this ould als
tims. Had any of th othr si trnsitions not happnd, and tht inluds
th origin of lif itslf (numr 1), thn w would not hr. Tht only two spis (Spkoski 19!
of th ight hav dmonstraly happnd mor than on spks against
In ths disussions it has
Simon onway Morris's (2003) thsis that ()nvrgntn natur is so pr-
valnt that humanity and mny othr tnt styls of lif r virtually tion fo mrosopi o
invitl. ros ar ludd u
and vry littl is known of
vr istd ar aliv tod
Divrsifiation to 750 million spis rn t
NURs
oDLs
Lif is astonishingly divers today, with stimtsrnging from 5 milliln to
Biodivrsity, thn, has p
100 million spis, ut prhaps' 1Orsorly, homing in on a 6gur -
twn 10 million and 15 million spis(sth min ssay ..Th Pattrn and tody. But how? Thr ar
Pross of Spiation'' y argart B. Ptk and Shala J. Hankison in this ths an prssdin t
vllum).Of ths,fwr than 2 million hv n rognizd and dsrid a straight lin, an ponI
ruptd inras (Figur 1;
so far.
i m p o s d ( F i g u 1 B ) .
It is ommonly assumd that iif today is mor divrs than it has v
n.This smsin som Ways ovilus,ut it ould aIso onstrud as x- The lir modl rprs
traordinary vanity, somhow akin ttl th viw that all volution was plannd r of nw spis in h
o t h r s i s a n t i n r a s ,i .
to lad to human iggs, nd that somhow this instant in th vast span of
tim is th most importnt of ll. Howvr, it is vidnt that all living organ. volutionay ,rnhing m
isms and ali organisms known as fossils driv fom singl ommon ans- tim spiation ats hav
larly at a rat suffiint to
tor (on th asis of th vidn of shrd ompl haratrs,sr-rhas th
DNA-RNA systm of inhritan,homoo gns,and th lik). That om. in rh t of volution in
mon anstor, th singl spis that gv ris tl all of lif, istd som t o t l n u m r o f s p i si s
.5_.8 illion yars ago. But how many spishav vr istd? t h o r d m a i n s f i x d ;h
Biologists nd palontologistshav trid rrumrof lins of rsonlng to o r p r o a i l i yo f s p l i t t i n g
stimat th totl numr of spis tht hav vr istd. Prhaps living ally n rjtd as impr
iodivrsity is only 24Y" of th total that hs vr istd' as is suggstdy The epoetil model i
thr lins of rasoning (Spkoski 19921a 1994): tion. If spiation and xt
will rgular douling <
1. Th first argumnt is asd on an assumd pattrn f spisdivrsity of volution at th lvl of
inras nd th known avrag duratiln of a spis for it oms nlltial rat of inas ol
xtint. If spisdivrsity inrasdroughly linarly (additiv modl) modl has n applid to
through th Phanrozoi' nd vagspisduration is 5_10 million th divsifiation of lif ir
Plotolog d th Histor of Lif 8.'
MoDLs
)Stimats ranging from 5 million to
Biodivrsity,thn' has pandd from on spisto many millions of spis
iorly, homing in on figur -
today.But how? hr ar many Ways to go from n spisto many' and
th main ssay..Th Pattrn and
thsan pssdin trms of thr mathmatial modls, rprsntdy
ak and Shala J. ankison in this
a straightlin, an xponntial urv' and a logisti urv' first as n unlntr-
tavn rognizdand dsrid
ruptdinras(Figur 1A), and sond with som mass tintions supr-
imposd(Figur 1B).
'y is mor divrs than it has vr
e liner modl rprsnts additiv inras' th addition of a fid num-
rut it ould also onstrud as x-
rof nw spisin h unit of tim. (Th inrasin this ampl and th
r,iwthat all volution ws plannd
othrsis a nt inras,i.., tru ins minus xtintions.)In trnrs of an
tow this instant in th vast span of
volutionryranhing modl, additiv inras would man that though
:r, it is vidntthat all living orgn-
tim spiationrats hav dlind, or tintion rats hav inrasdrgu-
|rivfom a singl ommon ans-
larly at a rat suffiintto mop up th ssspiations.Th implid dlin
:d omplx haratrs'suh as th
in th rat of volution in th linar modl oms aout simply usth
bo gns,and th lik). That om-
total numr of spisis inrasingrgularly, yt th rt of inrasaross
v ris to ll of lif, istd som
th oard rmains fid; hn,for any individual volutionary lin, th rat
spishav vr xistd?
or proabilityof splitting (spiatin)must dlin. Suh a modl has gnr-
d a n u m o f l i n so f r e s n i n gt o
ally nrjtdas improal.
t hav vr istd. Prhaps living
The pontilmodl is mor onsistntwith a ranhing mod of volu-
' has vristd,as is suggstdy
tion. If spiatilnand tintion rats rmain roughly onstant' thn thr
tla |994):
will rgular douling of divrsity within fid units of tim. A stady rat
;umdpattrnof spisdivrsity of volutiont th lvl of individul volutionary lins salsup to n xpo-
rtion of a spisfor it oms nntialrat of inras ovrall sin total divrsity is vr inrasing. This
roughly linarly (additivmodl) modlhas n applid to th divrsifiationrats of individual lads and to
;spisduration is 5_10 million thdivrsifiationof lif in gnral (Bnton 1995; Hwzulla t a|. 1999|.
86 Plrltllog1'
d th Histor of Lif
B PIaozoi
G e o | o g i |t i m e
r lt i m e
200
'r00
Figur . Pattrns of th divrsifiation oi lif thrlugh tim in trms of hangs
in numrs of fmilis tant r strtigraphi stag, plttd for all organisms
(A), ontinntal orgnisms (B), nd marin organisms (). In ah grph a
mximunr nd minimum ar slrown, asd on a ot-r-rination of strtigraPhi
and haitt.prfrninforn-ratin.Th minimum masur inluds or-rl
familisrordd as dfinitly prsnt within ah stratigrphi stag or as
definitlyspnning tat stag, and only fmilis dsignatd as rstritd solly to
th marin or ontiDlltl rlm. Th miInr.rmmasllr inluds trlso all
doutfLrlstratigraphi attriutions of familis and ll quivoal nd shard
haitatdsignations.Th sum of minimum masurs fr ontinntal nd rnain
organismsis qual to th minimllm masll for all taxa togthr. Th sum of
mimur Ilasllrs'h<lw.vr, dtls not qul th mimum m-asufor ll
Jur ret txa usfamilis with qtrivol nvirnmnta[ ssignmnts and thos that
our in oth marin and ntinntl sttings ar ountd as oth marin and
ontinntl.Stratigrerphiarviations as in Figur 2 with th ddition of n,
nozti;P' PrirmriaIr.(Bsd lr.rBrrton l99.5.)
e2 lotrllog1'd th L{istorof Lif
Until 1970 most volutionists assumd tlrt fossils <luld s,v r,y littl Th puntuatd-q
original about spiation' Tn ldrdg and Gould (197z) hallngd th 1970s and 1980s was
onsnsuswith thir thory of volution y puntuatd qtriliria.ldrdg tagonists kpt shifting
and Gould trmd th standard viwpoint phlti grdulism. It assumd a s do n t h i d a s i
that volving linags Wr hanging at vail rats (Figur 4A) ut wr rl supports of pun.
hanging mor o lss ontinuos| (gsls/.Spiationr'vassn as a y- ida of speies sltio
podut of this pross lf hang: sointi,slinagsha.lom so diffr- and' in som modls,
nt from thir starting points that thy ha] volvd into ir nw spis (th ordr haratrs. Sp<
hroosp:iesonpt). Spiation y splitting (ldogesi.s/ould lstl hp- and vn in oppositio
pn, ut th pross Was no slowr or fastr than normal rats of linag ida of an ..pansior
volutin. (Gould 1990), with h
Th opposing viwpoint' volution puttttdquilibritl (Figr-rr48), Darwinian viw. Suh
proposd that rapid morphologial harrgt ladognsis,rathr than alr- most nthusiasts, and
gnsis,Was th most important pross.Tlr normal stat Of a linag was natural sltion duri
s/asls (no hang),nd fronr tim to tim spitionwould hpprr,ut suh ally onsidrd to t,
spiation vnts wr rapid and rvolutionarY.Th puntutd-quilirium to rsult in signifiant
modl maks two laims: tht rts of hang ;rlonga lingr'plrntuatd' tion of individuals. H
nd that rapid hang is orrlatd witlr s;liationvnts.Th lattr lairn dn pointd to its ir
was th most ontrovrsial,ut it has n rgardd as th ky, dfining l- to xplain it (Land 19
mnt of th puntuatd-qullrbriummodl L.'yits suppotrs. A plthora of as
ldrdg and Gould asd thir nw modl on two osrvations: |99 that purpotd
1. Stsis is ommon in th fossil rord. Fossils an min onstant ln quiliriumviws..ar
not provide nough v
apParanthrtlr'rghmny mtrs of siimnt(i..'thotrsndsor
millior-rsof yars),and thn vrythingsmsto hang' of migrtionsof tx i
2. If most spiation happns ording to Mayr's lloptri (gographi- wht was a gradul i
stppd, puntuationa
splitting) modl, thn it would appar as a rapid vnt in rh fossil
rord. Th dtail of th gadual divrgrrof two is<llatl 1981; Shldon 1987) t
populations
intns fous on fin-s
would not sen.
thy wr still sujt
triloits showd a gr
ags just did not sp
mntd a numr of pu
intrprtd as mor lik
nonBnti hang in sl
h fossil rord dr
and Ansty (1995) sur
tns in th fossil r.
angd from radiolri
stratigraphi ags rang
iority onntrating in
of th arth. of th .5
|\,4ophoIo9y with anagnsis (15
6 3 % ) . I t s m s l a , t
F-igur4. ntrastingxptatiorrs of spis.lvl
r.llutionill th [lsi and it had not n pr
plrylti-grlr'talisn-r
modl (A) and tl-rpuntutd-quiliiu modl 1B). Wht thn of th pr
Dgrof nltlrphologiallrngis inditlon th -is, tit-t.l th 1,-1y1,.
t.lt.t at? As vr. th Dr
Pleotolognd th Histor of Lif 95
] thirt fossils ould say vry littl Th puntuatd-quiliium vrsus pylti-gdualism dat of th
n d G o u i d \ \ 9 7 2 ) h a l l n g dt h |970s and 1980s was hug fun for all onrnd' nt last austh pro-
)y puntuatdquiliria. ldrdg tagonistskpt shifting ground and rvising thir laims. Th first phas ws
ft p1,ltigrtdulisnt.It assumd asdon th idas in lddg and Gould's ssays' From 1975 onward sv-
arialrats (Figur 4A) ut wr ralsupportrsof puntuatdquiliria xtnddth modl to spousa nw
est/.SpiationWas sn as a y- ida of spissltion, a highr_ordr pross in whih spis wr sortd
ns li1s had om so diffr- and' in som modls, ould evn undrgo sltion of ..emrgnt'' highr-
rd volvd into a nw spis (th ordr haratrs. Spis sltion ould, in thory, our indpndnt of,
.ting (ldogezesls/ould aisl hap- and vn in opposition to' natural sltion (Stanly 1975). This ld to th
lstr than nomal rats of linag ida of an ..pnsion'' of volr.rtionarythory to many hirarhial lvls
(Gould \990), with hints of an apparnt rjtin of aspts of th no-
plttutdquilibri (Figur 48), Darwinian viw. Suh xtrm positions w andond fairly rapidly y
at ladognsis,rilthr th:rn na- most nthusiasts,and paloiologistsontinud thir staunh adhnto
Th normal stat Of a linag was naturalsltionduring th lat 1980s and 1990s. Spissltion is gnr-
;pi:rtion.tluldhappn. ut suh ally onsidrdto thotiallpossil ut trmly fl and unlikly
'nary. Th puntuatd-quilibrium to rsult in signifiant adaptations of th typ that our in th natural sl-
rg along a linag punttlatd' don of individuals. owvr, th notion of stasis stuk: too muh fossil vi-
;pitionvnts.lr latrr laim dnpointd to its imp<lrtn,nd qllantitativ gnti modis wr found
r rgadds th k' dfinir.rg l- to plain it (Land 1986).
t y its supportrs. A plthora of s studis ws ssmld and pulishd from 1975 to
ldl on tr,oosrvatios: |99 that purportd to tst th phyleti-gradualism vsus puntuatd_
quilirium viws. arly fforts wr oftn inadquatly doumntd and did
F o s s i l s l n n l i n t r t l 5 1 1n','
not provid nough vidnaout th auray of dating and th possiility
dir-t-rr-rt (i.., tlrotlslrrrdslr
of migrationsof taa in and out of th study ara. It soon am lar that
g smsto hang.
what was a gradual angnti pattn to on prson was n oviously
t ayr's lloptri (gograhi- .Williamson
. a s a r a p i d v n ti n stppd,puntuationl pattrn tO anot. Som studis (.g.,
th fossil
1981;Shldo |987) involvd hundrds of thousnds of spimns and an
rgnof two isolatdpopulations
intnsfous on fin-sal dating and statistial analysis of hug sampls' ut
thy wr still suit to itiisnr. For mpl, Shldn (1987) study of
triloits showd a grat dl of anagnsis and stasis' ut his sampld lin-
a g sj u s t d i d n o t s p i a t . . W i l l i a m s o n( i 9 8 1 ) l i v dt h a t h h a d d o u -
mntda numr of puntuational spiation vnts)ut thy wr gnrlly
interprtdas mor likly to xampls of ecopheotpic hang, that is,
nongntihang in shp rsulting from tmporary nvironmntal strsss.
Th fossil rord dmonstrats th widsprad ourrn of stasis. rwin
and Ansty (1995) summarizd th rsults of 58 studis of spiation pat-
trns in th fossil ord pulishd twn 1972 and 1995. orgnisms
rangd from radiolaria and foraminifra to ammonits and mammals' and
stratigraphi ags rngd from th ambrian to th Nogn, with th ma-
jrity onntratingin th Nogn' th past 25 million ys of th history
of th arth. of th 58 studis, 41 (71%) showed stasis, assoiatd ithr
l\,4orphology
with anagnsis(15 ase 7o) or with puntuatd pattrns (26 ass;
63%),|t smslar, thn, that stasis is ommon in speis_lvlvolution,
-lvlvolutionirrth lssi and it had not bn prditd from modrn gnti studis.
u t d - q u i l i r i rmrmo d l( B 1 . What thn of th puntuatd-quiliriumvrsus phylti-gradualism d-
t l r lt h - i s .t i m t r nt h y - . r l s . bat? As vr' th potagonists on ithr sid sought to dmonstrt th
Pleotolog nd the Ilistor of Lif
trm thr ar organisms suh as planktoni protists that liv in hug plpu-
4.0
lations that sldom nountr arrirs to disprsal, and that do not possss
omplx bhaviors assoiatedwith rprodution. For ths,gnti isolation 3.0
of populations is a rar vnt, and spiation,whih prhaps oLlrsrarly' is
2.O
probaly gnrally long trm and gradual, lasting prhaps half million
yars. At th othr xtrm ar organisms suh as frshwatr fishs that liv
in spatially struturd and oftn-transintnvironmnts. r spiation might
b so ommon that vry lak and rivr has its own rprodutivly isolatd
population of a partiular typ <lffish, ut thspopultions might nithr 2.8
:talthings.Whih Dvonin o. Bnton,M. J., and F. |. Ayal. 2003. Dating th tr of |ife.Si30: 1698_\70.
limdinststhat rpt round th Bnton,N,I.J., and D. A. T. arpr. 1997. tssiPltolog. arlow, ss' U.K.:
l,ongmn.
th air? \hih Jurassi osrvr
2008. Itrodl|tio to Plobiiog d th Histor of Life, Oxford:
rw flitting and daning in th sun- Blakwll.
rrs would volv soiality and thn Bnton' M. J., and P. N. Parson ' 2001 , Spiation in th fossil rord. rds i
to untold lvls?urrnt iodivr- F'olog d urlltkl 16: 4 5-41 1'.
.W.
night lif volv nxt? Thr r in- Bnton,. J., and G. Strrs. 1994. Tsting th quality of th fossil rord:
Plontologilknowldg is improving. Geolog 22z |t|_1\4'
Loiling Watrs' and som tht liv
Bnttln,.J., . A. Wills, nd R. Hithin. 2000. Quality of th fossil rord
roks bnath th sa floor. Ths
through ttm.Ntur 4: 54_57.
p p n da n d a r h a p p n i n g . Brmham, [-.D., and . D. Hndy. 2000. an fst arly rats ronil molular
y of lif, though, than mr dou- dts tl th amrian plosin? Prodigs lf th Rol Soit of Lodo,
int groups. Dinosaurs annot Sris B 267: "l'041-|047 .
l and show us rganisms that did htham'A. H., and J. B. .Jaksn. 1995. Pross {rom pattrn:Tsts for
sltinvrsus random hng in puntuatd ryozoan spiation.In D. H.
LrrsWr 10 tims th mass of th
rwir"rand R' I,. Ansty, ds.' Ne, p1lrhs to Spcitk i th Fossil
rad wingspansthr or four tims Rlrd,\84_207. Nw York: oluIia Univrsity Prss.
tint arthropds wr largr than nway orris, s. 2003. Lif's Solutio: Iuitbl Hms i Lol Uiurse.
r u | y w o n d r f u l .a n d r h y p o s i n - mridg: mridg Univrsity Prss.
lphysiologists. opr, A., nd D. Pnny. 1997. \4ass survival of irds aross th rtaous-
Trtir or-rnday:Molulr vidn.Siee275: 1109-1'113.
n prditd(rtroditd?)from
Darrvin,. 1tls9' th rigi of Sples.London: John urray.
ails hav always rmaind mystri-
astal,S. 1999. iVlolulrvidnfor th arly divrgnof plantal mammals.
ur of ing al to disst suh Bbl.ssays 21 :'l 0.52-10-58.
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lprov' th lvl of undrstanding grdualism.In T..|. M. Shopf, d.,odIs i Pleobiology, 82_115. San
Franiso:Frman, oopr.
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nd R. I-. Ansty, ds., N prlhs fu Spitil i tb Fossil Rcord'
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ry, and W ar ginningto s for ]lttrIof the }ologiclSlit, Lldo 159:489491'.
Gould, S. .|. 1990.Is nw and gnral thory of volution mrging?Pleobiolog
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nw insights into iomhanis,
Gould, S.J., and . B. alloway. 1980. lms and rahiopods-ships that pass in
) vn mor imprssiv.Th study
th night. PIobiolog6: 8-396.
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dgs'S. B., and S. umar.2004. Prision of molular tim stimts.Trndsin
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Kumar, S., nd S. B. Hdgs' 1998. A molulr timsai{or vrtratvolution.
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Laandira'C. ., and J. J. Spkoski lr. 1993. Inst divrsity in th fossil rord.
|lfe' rds i oklg d S i 2 6 1 : 3 1 0 - 3 1 5
Lnd, R. 19B6. Th dynamis of pak shifts and th pttrn of morphologial
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Mwll' W. D., and . J. Bnton. 1990. Historial tstsof th absolut
tn P. . J. Donoghu nd . P. ompltnssof th fossil rord of ttrpods.Plobiolog 16:322-35.
: Iulr Ilcks d th Fossil May' R. \4. 1994. nptual spts lf th quantifiation of th tnt of iologial
is. divrsity. l,hillsophil rstit-lslf th Rol Soiet, Sris B 345:13-2.
104 Pleontolog d the Histor of Life
Adaptation
dn for vrtrathistory. Si
105
106 Adpttil
fatur of a plant or animal. But thr wr ovious limits to mallaility- th gns rsponsil fo
hikns nnot b rd to ly two ggs pr day evry day of tlrir livs-and thy wr in th prviotr
natur smdal to distingurshonly sustantialdiffrsamong spis dvlopmnt did with .
or stoks (Provin 1971; Rus 1979)' Ths substantial diffrnssmd quny of th favord gt
apal of ing produd only y sltivrdingof fam stok (if natur ltion to inras;mor
Was sorting among varitis of donrsti goats introdud to an isl:rnd) or avrag th nt gnrt
original ration (if natur was sorting among spis). ltion ours onsistnt
This xplanation of adaptiltion am inrasingly umrsome s mor ing sltd h:rngsu
and mor fossil organisms wl.disovrd.Fo on thing, layrs of rok that tiotr ats on it vrit
inludd fossilizd spis did not inlud fossilizd matrial from xtant g r a t r h u m r r l e t i v
spis;if natur wr mrly filtring spis'whr Ws th vidn that An daptation is any
xtant spis had vr oistd with tint ons? For anothr' although nti rsons to spi
many fossil organisms or liml rsrblanto xtant ons (.g.'dirrosaurs), pl' w all rpir'1dvlo
othrs smdvry similar to xtant spis(.g.'fossil floundrs).It was un- tmporary ponds. Adap
lar why on spisshould hav gon xtint wlril anothr with vy simi. aus it xplins th
lar fatursshould prsist. v o k i n g p u r p t l s f u ld s i g
Th sintifi rvolution in th stud of adptation is th ida of natural s- mnt for how nw sp
ltion' as suggstdy harls Darwin and Alfrd Russl walla. Although popultions down div1
both mn offd th ida, it ws Dwin who artiulatd it ror ompltly om so tnsivth
and intgratd it mor thoroughly into a largr thory of iologial volu- Sparlrtspis.
tion. As a rsult, w us th trm Drii uolutil to rfr to th pross Darwin's ritial argt
through whih adaptations mrg'a prossalso lld uolutio b trl ould irrdd disringurs
seltiotl or dptiue euolutil. t r a v l s h h a d s n 't l
In Darwinian volution, natural sltion, upld with inhritan of tions of th sam plan
small trit diffrns'is th gnt of adptation. Natual sltionis a statis- ass ths diffrnsr
tial filtring Pross; individuals with partiular haratristis r mor Arlriplago; in oths l
likly to surviv or produ mor offspring than individuals with othr har- i n t h s a m i r t ls p i
atristis.A mor formal dfinition of natural sltionis th ausal ssoi- mad rhs kinds of o
ation of a fatur or trait with fitnss. B fitss w man th numr of H saw all ths 1iffr
offspring that an individual lavshind; individuals that di without rpro- n t u a s f i I t r i l l g i s t
duing hav zro fitnss. The usl ssoitio of feture or trit tuith fit- individul diffrns t
ness mans' oprationally, that a fatur or vlu of rrait is dirtly sustntilulultiv
rsponsil for whthr an individul livs or dis or why som irrdividuals a m o n g p o p u l a t i o n s . }
lav mor offspring than othrs. tion of whthr natural
For xampl' many frogs brd in tmprary plnds tht hold watr for a whethr trait diffrn
rif priod and dry vry sason.Th tadpls in thsponds must orplt and ultirnatly wht
thir dvlopmnt into juvnil frogs for th pond dris and thy ar i n t h i r s | t i r 'r g i m
killd. Tadpols with vry slow rats of dvlopmntdi austhy annot a half.
omplt mtamphosisfor th pond dris. Ntural sltinfiltrs indi-
viduals with diffrnt rats of dvlopmnt so that only thos with mor
rapid dvlopmnt mrg from th pond. vidn for Adap
This filtring would hav no lasting efft if irrdividuals who dvlopd
rapidly wr not gntially distint from thos that dvloped slowly. Th Biologists hirv onttn
fftivnss of natural sltion mgs from th hrital natur of trait mnt and hav tn
variation; in th nt gnration th gnti varints rsponsilfor th fa- lvl. Thy hav als
voral trzritvalus will inras in rlativ aundan. In tlr frog arpl, mk sintifi xpla
dDttio 107
obvious limits to mallaility- th gnsrsponsil for mor rapid dvlopmnt ar mor ommon than
dy vry day of thir livs-and thywr in th prvious gnrationausth gnsrsponsilfor slowr
;tantialdiffrnsamong spis dvlopmntdid with th tadpols that arrid thm. Th inras irr fr-
s substantial diffrns smd qunyof th favlrdgnsauss th avragvalus of th traits r.rndrs-
rdingof farm stok (if natur ltionto inras;mor opis of gnsfor rapid dvlopmntman that on
.Whn
;oatsintrodud to an island) or avragth nxt gnrtionof tadpols will dvlop fastr. ntural s-
,ngspis). ltionours onsistntlyovr tim, th avragvalus of th traits that ar
nrasinglyumrsom as mor bing sltdhang sustantilly.Th long and mor onsistntl sl.
For on thing' layrs of rok that tion ats or-rtrait varition and th mor gnti vaitior-rin th trait, th
: fossilizd matrial from xrant gratrth umulativ hang in trait valus.
:is,whr was th vidn that An adaptation is any fatur of an organism that has vtllvd through g-
int ons? For anothr, although ntirsponsto a spifiologial agnt of natural sltiln.ln our m.
:to xtantons (.g.,dinosaurs), pl' w all rapid dvlopmntrat an adaptation to th risk of dsitionin
(.g.,fossil floundrs).It was un- tmporaryponds. Adaptation is ntral to Darwin's argumnt for volution
n r w h i I a n o t h w i t h v r y s i m i - bausit plins th maths twnfaturand nvironmnt withut in.
voking purposful dsign or spial ration. It also provids th vital argu-
laptationis th ida of natural s- mnt for how nw spis aris; whn adaptiv volution taks diffrnt
.!7alla.
Alfrd Russl Although populationsdown divrgntpaths, th diffrnstwnthos populations
ho rtiulatd it mor ompltly omso tnsivthat thy no longr an intrrd,and ah oms a
argr thory of iologial volu- sparatsp1s.
l uolution to rfr to th pross Darwin's ritial ergumnt Was that natur's filtr, his natul sltion,
;s also alld uoltio b turl ould indd distinguish srall diffrns among individuals. Dr"rringhis
trvls h had sn, tim and again, diffrns in fatus mong popula-
n, oupld with inhritan of tions of th sm plant or animl spis in diffrnt loirtions. ln som
. t i o n .N a t u r l s l t i o ni s s t a t i s - assths diffrnswr larg and dramati, as h sw in th Galipagos
rtiulr haratristisr mor Arhiplago; in lthrsthy wr smallr and mor sutl, s in diffrns
than individuals with othr har- in th sam ird spisin diffrnt loations in South Amria. )thrshad
rral sltionis th ausal ssoi- mad ths kinds of osrvations, ut Darwin intrprtd thm diffrntly.
fitss we man th numr of saw all ths diffrnsas points along a ontinuum; rathr than srng
rdividualsthat di without rpro- natur as filtring xisting larg diffrns,h saw natur s filtring small
tion of feture lr trit tuitb fit- individual diffrns that, lupld with inhritan and tim, produd
or valu of a trait is dirtly sustantial umulativ hangs in fturs and sustarrtil divrgn
or dis or why som individuals a m o n g p p u l a t i o n s .T h t s t i n g o f D a r w i n ' s a f ! ] u m n t ,t h a t i s , t h s o l u .
tion of whthr natural sltion ould as powrful as artifiial siti<rtl,
rary ponds that hold watr for a whthr trait diffrns that wr undr sltiorrwould prov hritabl,
lls in thseponds must omplt and ultimatly whther populations would dir,rg through diffrns
'r th pnd dris and thy ar in thir sltiv rgims, has oupid iologists for naly i1 ntuy and
:lopmntdi austhy annot a half.
:is.Natural sltion filtrs indi-
rt so that only thos with mor
vidnfor Adaptation through Sltion
:t if individuals who dvlopd
hos that dvlopd slowly. Th Biologists hav ontinud to dsri th math of tlrganism to nvlron-
om th hritabl natur of trit mnt and hv tndd our knowldg of that math to th molul
varints rsponsil for th fa- l v l .T h y h a v a l s o u n o v r d s t r i k i n g p a t t r n s i n t h o s m a t h s t h t
aundan.In th frog ampl, mak sintifi xplanations tlth than l)arwinian volution untnill.
1 0 8 Adpttiotl
rtal studisthy hav providd th a diffrnthmogloin gn whos protin havslik most vrtrath-
prsould nly pondr. nroglobirr(Fdr and Brrrggn 1992).
Som f th most striking patrns of adaptation involv onvrgnt or
parallladaptation.onvrgnt a1aptationours whn th sam adaptation
is found in unrltdorganis-rs;paralll adaptation ours whn rlatd or-
:ions.arnivorous plants our in ganismsdisplay th sam adaptatin through indpndnt origins. In oth
:apturand digstinststo btain assadaptationsrfltrpatalmaths of rg:rnismto nvironmnt.For
ough thir rtlots. rtain fish and xampl,two unrlatd frogs that liv undr similaly drrrandinghot dsrt
ds and floodplains av ggs that onditions on two diffrnt ontinnts' Phlllmds suugii in South
in th soil until floodwatrs rek Amri ad birlmntisrmpili in South Afria ah ert uri aid
. Svralspisof fish liv in polar athrthan ura' as do othr adLrltfrogs; uri aid xrtion rqr-rirsaout
i, rlying on a spializdantifrz on-twntithof th watr rquird for ura rtion and rprsnts an
':ithinthir odis (ohahka d adaptivsolution t th nd for wtr onsrvation (Shmidt-Nilsn1990).
A drmati arpl of paralll adaptation is th ontrast twn th lif
vritis.Dmslfly spisin th historisof guppis from upstram and downstram loations in Trinidad
rflis as tlrir prinrary prdators that is rpatdin svrl drairrgsfrom on muntitl ang to anothr
havior than spis that our arossth island. Guppis from upstram loations om rprodutivly
haviors that prott against drag- matur at latr ags nd largr sizs than guppis from downstrm loa-
ld vi vrsa' so th haviors ar tions, lnd upstram fr-rrals
produ fwr, largr ais in ah rood.
or (MPk 1995I' Th fungal Ths diffrns ar mathd to thir diffrnt nvironmnts; downstram
m grows on two losly rltd guppissuffr mu high prdation rats, hav lowr poprrlation dnsities,
am loations, Phseols ulgris and, in gnral,must omplt thir lif yl quikly or not t all. Guppis
;ions and sirrringon ah spis. from diffrnt drainags ar muh lss losly rlatd than upstram and
1 t sa s p i l i z d. l t s u r l . I 1 [ l n t i n g downstrmguppis in tlr sanr ivr, rvhilr indiats that th distintions
as diffrnt spisof damslflis twnth upstram arrd downstram popultions hav volvd tim and
' d a t o r s( S i a r d
t al. 2007). Pop- again (Rznik and Trvis 1996).
zs from mor northrn latituds
: dehydrognasthan populations
DoNsRA|oNs oF NAURAL sL|oN
,u:rnd th rgulation of prs-
g thrmal rgimsto produ sim- any iology ttooks still rploy . B. D. Kttlwll's lassi study of in.
t tmpraturs'whih allows th dustrialmlanism to illustrat th powr of natural sltion (.g.,Futuyma
f thrml nvironmnts (Shult 2005). As tr trunks in Britain turnd dark fror-nir pollrrtion, th ol.
oration of th ppprd moth' Bistl btulri, hangd from mottld to
rrly striking austhy oftn in- lak ovr a larg ara. Th ttooks usully dsriKttlwll's osrv-
s y th sarrrindividual at diffr. tions of prdation on tthrd moths of diffrnt olors on diffrnt ak_
n of hmogloinin frogs. In most grounds;prdatsfound it asy to dttnrottld moths ln dark tr trunks
: lood that arrisoygn to lls and lak moths on lihn-llor]trunks in unpollutd forsts.Although th
lnditions. This proprty nlrans valu of th osrvationson ttlrrdroths has n qustiond y ritis'
n ar aidi nvironmnts,and h- thos osrvations W not th ritiirl vidn. Th onvining data
to thos lls. But mny tadpols mrgd from Kttlwll's mtiulous mark-nd-raptur studis; h
rmogloirrmolul with th r- markd larg nums of moths of diffrnt <llors,rlasd thm, irnd
tdr mor aidi onditions.Ths ountdth numr of raptursof ah typ of moth in diffrnt loations.
hmogloinfailitatsthir ility lris work demotrstrtdtt th diffrnt olor typs had diffrnt nrortlity
'hn thr is littl of
it availal. IatS'and that th math twn motlr olor and bakground was ing
) oygn is adilyrlasdwhr ..Industrial
moldd by sltion(sth alphatialntry mlanisrn'' in this
r f r o g s ,w h i h r . r t h l r , x p r s s volum).
1 1 0 dpttil
ns of naturl sltion in tion. A In th most thorough studis iologists hav r-ratdth prsumd vo-
iffnsin -rortalit1,rts o lvls lutionarytrajtory of th diffrns.Th Triniddian guppis off on of
rls with ditlrnttr:lit valus tlr dif- th most thoroughly studid xarnpls.Rall that upstrm guppis, rgard-
datorsand pathogtlsto oli strss lssof stram or mountin rang, liv llngr,matur ltr and largr, and
,ririslrforsts to Dilrwin 6r"rhsirr havfwr' largr offspring at ah or-rtof rprodutiorrthan thi los rl_
ition th rality of natural slti<ln ativsdownstram. Th magnitud of ths diffrns is not trivial; fmals
in upstam loations ar-r mor than 20" |argr at mturity, hav fwr
ut an it b a drir,ing fo of vl- than half as many offspring as downstram guppis, and produ individual
turl sltionthat ar ruiill to s- nwornsthat ar 50_75% largr than tos produd y downstram gup-
I u t i o n .F i r t . e n . l t i t l n i l l n O n l pis' Th ar a numr of upstram loations aov sustantial watrfalls
nsistntto mold th faturs of o- in whih guppis ar asnt.Jolrn ndlr nd Dvid Rznik ploitd this
: variation flr traits undr sltion opportunity by introduing downstram guppis into som of ths loa.
ftus?r'vtl lins of l,idrran- tions,in fftrplitingtlr likly ours of guppy oloniztion of upstram
habitat(Rznik et a|.1.997).Th introdud guppis followd an volution-
nvironmtalaltrationsmird y ary path tht in lss than si yars produd trait valus omparal with
) thosltratilns. h dvIopmrrr thos sn in naturl upstrarn popultions. Natural sltiorrin th up-
owd humns to rat novl nvi. strmnvironrnt' ating on th gnti variation that was produing small
ltrium' n arrtiiotiis toin; to diffrnsamog individual downstram guppis trnsplantd upstram'
it poisonsth ll tlr dnaturit and moldd nw populations of upstram gtrppisirr vry short ordr. [t was th
a powrful sltiv for for altr- primntthat Darwin would hav likd to do.
;ll; individrralsthat <llddlryrlr
d rprodu mor otn tharr thos
lratingp' spisaftr spis How D.W RonstrutAdaptiv volution?
a rrldtlrnrto tk on o an-
of antibiotis.Although th lls Although studis of sltion and rspons dmonstrat that Darwintn vo-
id in th l 90s, thir iohmistry lution ours,th full pitur of adaptation mrgswhn stdisof sltion
( D z i d i t l . 2 0 0 8 ) . ar joind with painstaking work in grrtis,dvlopmnt, arrd plontol.
s p n s t t l n l t l d r n h m i s t r yi s n t l t ogy. of ours' the pitur is nithr omplt nor prft; iologists hav
lf instsar.voIv]iohn-ril not studidvrv striking faturof vry organism, and vn for som of th
nzte |987):rnd wlyplant spis most srutinizdass' thr is still muh to larn.
is, vn glyphsat,th ..lnin a Th tatursthat dfin th modrn quids, horss and zbras, offr a par-
r aquatl oganisms hv r,olvd tiulrly strong mple (MFaddn 1992). Palontology has rvld th
:d havy lntals in watr (Klks squnof fatur apparan from th longation of th lowr lims, th
nts hav unlrgontrrsivil- rdution of tos, and th mrgnof hoovs to th inrasing hight of
rsponsto th i]rsnof havy th rnolars.BionrhniaIstudis hav pinted to th dvntagsoffrd y
f rining,lddgas<riin, nd vn partiularfaturs;longatd lims failitat rapid running for xtndd p-
9871. riods, and highr molars ar fiial for grazing touglr, silia-ontaining
; h a v s r u d i dh u r r . l r dtsl i a s si n vgtationlik grass. Plaing ths hangs in th ontt of a hanging li_
r diffrntllationshiit radily mat and th mrgnof savatrnasand plains, w an ronstrutth vo-
lholog,lif history, o ftursof lutionary pathwy through whih small anstal rowsrs produd
rplof intrr1ing. Thv hav family of graztng dsndants mtlldd y living irr opn Ountry nd ing at
and, through a varity of pri- risk from a varity f larg prdators.
ing idividuIs trvnJoatior.rs), Suh onstrutionpsumstlrat th transitionsw ronstrut,lik th
,n in th diffrnt lotions is r_ lngationof th lim and th rdution in th numr of digits in th foot,
; ( R z n i ka n d T r v i s l 9 9 6 ; T r a v i s ar dvlpmntallypossil.That is, th rtlnstrutionPrsums that a d-
r r2 0 0 1 ) . vlopmntainrhanis-rftlr produing suh hngs n dsrid, and
1 1 2 dpttil
th th gnti vri:rtions in tlr lrnntshrlr-rglr wlrih thr mhtlism qustions has inspird
tltrld t r'"'tluldhv n radily vailal. Although W anllor tst ths ian volution' vn a
prsumptionsirr tint spis,W-ilIl quir insights frtlm str-rdis of ist-
ing spis.
A4lsttrrstrialvrtrtshav fil, digits ln th flt at tlr nd of ah ontrovrsis in l
limb, ut th position of thos digits on th fo.tt,thir lativsiz, and vn
|s HE RAW
thir nr-nr ,.1nviv. Stu1islf ftlltand ligit dvltlpr-nrrt
il] tratal -
READILY AVAII
primnt:rlnimals lik salamandrsnd hikns hav shown that th num-
r, siz, and position of digits ar dtrmind},moltrlrsigr-rals harrgd At first glan th ans'
1,mryoni lls ]urirrgal,vlim dvl<lprnt (Shtrin2002)' Rlativly ttion is rdily avail
small liffnsin rh timing, inttrsity,and pattrrringof thos signals guppis offrs ompt
hrrgth sr.rs1ullt pattrn of digit dvloprrlnt.Dlays in signaling lvill sstantial adaptiv
it-llpdth forr;rtilnof th third, fotrth,and fifth digits, and if thy dl d- x p t i o n a l s :h u .
vllp, tlros digits an quit srll. Dlys irr signaling, opld with ellular onntrtio
gr()\^,,thin th foot itslf, will ltr th ositi<lno digits on th flot.Thr"rsl- h i n d l i m l n g t ho f t r
thtlr'lghW nn()trtath prist1vltlpnlntal pth that ld to hrofin found apprial lv
fossil quids' W l111lffr :r plausil rnodl through whilr umulativ viduals within a singl
hllrtgsin signaling rrd gn prssilttor'rlll-ravtrltrrsformda fiv_t()d B u t h i s r n o l
ftlot inttl :r sillgl-todhoof. that natural sltin
\zilullglrti vrition f<lr t ky dvl<lpmntlimnts hav lln shar dvlopmntal
vilirl?T.hst llalog,v()lllsfrtllrl stulisof rlativ lrindllnl lngth.in valus to inhritd
frogs. Frtlg spisvary normously in th lngtlr tlf th hidlil lmpard to sltion. In svr
with th:rtlf th ody; rikt frogs hr,hindlims sustantiall1,longr tlran lis, swordtails, and
ti tld1.lrlgths,r'r,hrsspadflor-ttladlrindlims ar raly mo than an ndorin pathw1
:rout lrlf th ldy lngtlr.vn losly rlatd spislik th arking tr opmnt (Kllmnl9t
frog nd gl1tr frog lf th slutlrlrstrn Urritd Sttsan hr,sustn- soition btwn r.t
tiallr- diffrltlativ hindllm lngths.Ths diffrt-ls ris from diffr- h i s a s s o i a t i o n r a
ns in th timing of lim-ud initition and th rat of lim.ud growth in largr ody sizs at
th tir1p<lls (Blouin |99 I). Gnti vlrriatiorrtilr th rlativ lrrgthof th If slowr maturation l
hi]lim app:rrsradily l'irilal \, in sitlgl frog popultions (Blluin solution; tlr gnti
1992I. This work suggststhat th raw matrial for hindlim volutitln in maturation is not imr
fr<lgsrr.sponlrlv not liffiultfo sltilnto find in tl-rpast arrd, ,- - Th ky point v
tnsitln,suggsts silnila lrrlusionflr othr anin-rals. Straint hold in th l
lt is sy to ritiiz lnlttsionslrout adptation in htlrss whos pr- suffiintly long, wor
sllnlpti()nsl111 tstdonl\, tlrr<lugpri-ltltl studis o irds or fogs. ths traits? If sl
Suh trapoltion u'ill alw1,s nssarywlrn w attmpt t() ronstrut m u t t i o n s f i l i t a t t
adaptation orr a larg sal, suh as might f<lundin th fossil rord. It is and rapid maturatio
alstl asy rt>ritiiz't gnrality rlf onlrrsionsillut ada1rtatilndivd though it ay wll h
frll study on a snrall srl,suh as th as tlf industrial lnisrn. l a m aris, it is imp
riti, th rid lrIrgin th frqunr,of altrrativforms of :r singl gn wld favor suh
ftl lllratilnis lradly vidn tht naturi-llsltilt-l an div profound T h p o s s i i l i t yo f .
transformtionsin orgnismi f:lturs-l(lharrgingollr pttrnsin moths of adaptationsthat ;
tll rrs ntluglr tl undrstand h<lwvolrrtilnmol]sfish musls ir-rtolrri xampl' most asS
()rgns<lrtnsfln]splant lavs itlto taps ftl atlring and digstig in- h a v si n a p a r t i u
sts? h diffiulty of alaning what an ahivd xprimrrtallvat sum a partiular p
smallr sls ig:rillsr th l:rrgr.sals of th rrlost importnt volutilrlary oloration ar filrr
dpttil r1)
gits on th foot at th nd f ah
ontrovrsisin th Study of Adaptation
t foot, tlrirlirtivsiz' and vn
i digit dvlomntin tratal - |s H RAW MARIAL FoR ADAPA|oN
:hiknshar'shlwntht tlr num- RADlLY AVAlLAL?
ld y mtllrrlrsignlslrngd
At first gln th answr to th qustion whthr th rw nrtril for dp.
l t l p m n t( S h u i n2 o o 2 I . R l a t i v l y
tation is radily availal would sm ovious. Th study f th transplantd
, :rnd pattrning lf thtls sigI-rals guppis offrs omplling vidn that th grrivariation rquird for
.lopmtrt.Dlays in signalirrgr,l.ill
sustantiladaptiv volution ws avilal. This dos not appar to an
nd fifth digits, and if thy do d-
ptionalas;hundrds of studisof a varity of haratrs,ranging from
)lys irr signaiing, oupld u,ith
llular onntrations of alohol dhydrognas in flis and th rlativ
tion of digits on tlr foot. Thus al-
hindlim lngth of fogs to th olor of swallowtail uttrfly pupa, hav
pmntirlpath that ld to a hoof in
-rrodl found appriallvls of gntivariation for individual traits among indi.
through ,lrih umulativ vidulswithin a singl population.
:ould hav trnsflrml fiv-tod
But th istnof hrital variation in singl haatsdos not imply
that natural sltionan mold fatrsaritrarily. Diffrnt haratrs an
vlopmntIlmnts hav rr
shar dvlopmr.rtal pathways that aus partiulr omintions of trait
-rdisof rlativhindlim lngth in
vlusto inhritd togthrand thry onstrain th immdiat rspons
. lngth of tlr hirldlir
mpard to sltion.In svral spisof pliliid fish-th fmily of guppis, rnol-
i n d l i m ss u s t a n t i i r l livo n g t a )
iies,swordtails' arrd mosquito fish-th pross of mturation is govnd y
d h i n d l i m sa r r a r l y m o r t h a n
an ndorinpathwy that rdusgrowth rat as it alratssual dvl-
:latdspislik th arking tr
opmnt (Kallman 1989). Th gntivariation in this pathway ratsan as-
LjnitdStatsan hv sLlstlr-
.hs soiation twn rapid nraturation rat an] sm:rll ody siz at mturity.
diffrnsis from diffr- This assoiation mans that th rspons to a sltion pfssur that favors
nd tlr rt tlf linr_dgrowth in
largr ody sizs at maturity will invitly inlud slowr maturation rts.
tion for tlr rlativ lrrgthof t
If slowr maturation rats ar dtrimntal for an rason' thr is no ovious
n singl frlg popultions (Blouin
slution; th gnti variation for th omination of larg siz and rapid
rtril ftlr hirrdiim vllrtion irr
maturation is not inrrnditlyvailal.
on to find in th past lrnd, y -
Th ky point rvolvs around th word immedit'l; dos su a on-
t h ra n i m a l s .
straint hold in th long run? /r w to wath suffiir-rtlylosly and wait
adaptation i hoss rvhtls pr-
suffiintlylong, would W se mtlttios in individual gns that dissoit
rimntalstudison irds or frogs.
thstraits? If sltion atd with suffiint Onsistny'would ths nW
y rvhnW attmptto rostrut
mutations failitat th vntual mrgn of th orninatiorr of larg siz
flundin th ftlssil ror]'It is .W
and rapid maturtion? hv no simpl answrs to ths qustions; al-
l u s i o n sa o u t d a p t a t i o nd i v d
though it may wll that givn nough tim, a mutation of th right kind
s of industrial mlnisnl. Tl a
may aris, it is impossil to prdit if th ologial irumstans that
altrnativftlrms of singl gn
would favor su a mutation would still prvil whn it finally aros.
rral sltionan driv prlfound
Th possiility of onstraind gntivariation provoks a ronsidration
hrrgingollrpttrllsin moths
of adaptations that a uilt on assoi:rtionstwn individLrl trits. For
n rolds fish musls irrto ltri
xampl,most ass of rypti oloratiln in animals work only if th animal
ps fo athing nd digsting irr-
bhavsin a partiular fashion, wlrthr it is to main motionlss or to as-
n alrir,dprinrrrtallyt
sum partiula postur or intatio.isrnaths twn bhvior and
th most importnt volutionarv
oloration ar failurs. T adnt advoat of Dar,inian volution might
1 1 4 dpttio
argu that natural sltion hs assmldth omponllts of suh adapta. possibl solution ontl
tions via mtiulous molding of individual fi1tursuntil th st omina- s t i l l u s t r t i o n so f t h i
tion mrgd from all th possiilitis.At th othr trl n riglrt gu frnt poptrltins of t
-
that th ontd inhritan tlf th onrponntsoffrd natural sltiona ologial hallng.
limitd rng of rarv mtrial frorr-rwlrih only a fw good maths ttr ir- ids; diffrnt popul
umstanswr pssil. i o h m i l n d p h y s
Th sam argumnt n prov<>kd y onsidring th volution of ny (atsumur and Voss
puttiv rldapt:rtionthat involr.s rultipl haratrs, it ;l iohmil ltri fishs o-pt
patWay or a suit of assoiatdhavitlrs.Th ontrovrsy is not Whthr o g n s ( B n n t t1 9 7 1 )
suh fturs rprsnt adptlltirns;th ontrovrs is vr h xtnt t Th othr part { th
whih adapttion is onstraind y th natur of th vailal raw matrial. ural slti<ln filtrs r,
This is L.linglrpprolrd fr-l sval prsptivs, fror rinirrg th na- prior volution. As a l
tur of nW mutations for ky haratrsto disstingth ssoietionsamong tion, arrd history an g
individual fatursof an adaptation inttl thir gntiontrrland sard d. proably not what al
vlpmntal pthwavs. wr bing dsigrrdd
lind spot ausd y t
i n f f i i n ts t r u t t t r i s
ls ADAPATIoN PRF?
organ and susqunt
Th rrrarvlousmaths of orgnisrrrto nvirtlnmnt offr a strong t]pta- tion. An asir nd bt
tion to onlud that adptation is prft. Indd, whn ontmplting in- not through th rtin
triat adaptatins lik ag-spifihmogloin gn prssion,th amzd Thr is anothr Wi
sintist is rr<ltvr,v diffrnt frorn th awstruk plt or th inspird rd by volutionary
thologin. Brrt th sintist must tak this qustion es tstbl id. faturs of organisms
Thr ar two oprationl wys in whilr w rnight us th tert prft; it prforrn a spifi funt
ould rf to whthr vry fatur of an lrganism hs adptiv signifi- that aptur and fous
an nd/or wlrthr t matl-rrf orgirnism ttl tlvirtltlIntis as gool as molular signls intt
possil. bn onsidrd am
Nlt vry fatur of an organism is adaptivly signifiant.F.or olr thing, of ths omplx trait
onrtd inhritanof haratrs,as dsrilprviousl1,,mns tht som of Darwiniirn vo|uti
faturswith no adptiv signifianmay prsist austhy ar inhritd vry thm in th stu
throtrgh shrd dvlopmntl pathrvays rvith othr fturs that rrsnt
an adaptation. onrtd inhritann also pr()ttftursthat r dtri.
AN DARWlNI
mntal y thn-rslr,s if thy ar inhritd alorrg wit adVanta1]rous ha.
xPLA|N oM
trs and th nt fitnss fft of tlr mination is plsitiv. This is th
ssIlof rnthtlr for th volution of snsn;gnswith dtrimntal Darwin himslf appl
ffts at latr ags. aftr a sustantial rprodutiv priod, rmairr in th dilmma is tht all sr
population austhy onf a larg fitnssdvatagarly in lif, for boli pathway lik tht
or rly in an <lrgnism'srprodr-rtiv priod. psit f rany indil
Is th mt of fatur to nvironmnt as good as possil?Oviously th proprly intgratd f
answr to this qustion dpnds on lrorn,w dfirrptssible.Ad tlrrin lis ring so lany things
th nswr, at last in part. Ntural sltilndos not dsign an orgnism or ntural thologian's z
its fatus;it mrly filtrs xisting variation. Th nd produt of Drwirrian sam position tirkn l
volution is lways :ls good s possil, ut hr plssiblis dfind as th tnt thsis offrd y
st of th availal options' whih ar dtrmind in tun by th gnti th y or th Krs l
varition that is availal nd .hat th onsturintson that vaiati<lr-r might ltitln. Chnging th
. Put anothr way, adaptation is ontingnt pross;it onstrtttsth st nothr (Aala 2006
dpttion , t.)
But is it? Drwin argud that ni]turalsletiolroul1produ tlrrrpladap- hagfish). A sond intr
tations if thy volvd through a sris of srall stps, ilh of whih had sm ians, rptils, nd irds t
daptiv advantag ov its prdssor.Th v<llr.ltitln of th 1-ltrldonlv fish, and th dditilrra
valdy ronstutingth hist<lrialslutlof thssmll hangs,and from ons assoiatdW
an undrstandingf lvhy an y lllksas it dos rquirsan ttndrst:rnding of Ths would rmark;
tlrat sar.nsignatur of history (.Wlrrand Dpw 2004). To th modrn volu- tion drov th wholsal
tionary iologist,th ferilurto appriatth onringntand historilnaturs shaps and rting ntl
lf adaptation is th signal w,knssin th ar]ulnntstilr IntlligntDsign. z 0o7l .
Darwin illustratd lris argmntflr th v 1,pointing otrt th divrsity of But th diffrns rn
potosnsitivstruturs in diffrnt spis tlf gastropods (snaiIs,russls, st. Gill arhs ar a (
squid, and otpuss).Ths struturs reng frot.nlr-rstrs of lls in sor s a llows. Fish us t
lams that an disrimirrt twn light lld dak to t spistiatd, avity. .Whn a fish opl
mra-lik ys of an Otopus' with a rng of phltosnsitir, ogns of in- oral avity inrass,an'
rasing mplxitv in twn. Drr,vin grrdL.l1. nlogv, ssrting that bllows ontrts and r(
tlr modrn divrsity of phtlsnsitivtlrgans in gastropods illustrats th watr through th gill sl
stpsthr ould hv orlfror ansrort() dsndantto uild tlr most stors of all iawd vrl
omplietd of ths orgns' th otopus y, with a omparal plssiility trorne prtils into t
for uilding th vrtrat }l.In this r]umlltsn-rallhangs agrinstth in th oral vity, whi
akground of what hs ll for n aumult into a rrrrarkalrl also funtions as a rsp
trirnsforrrrationof lls nd tissus. wll supplid with blo
Dar."vin'sargumrlt,whil rvalinghis grat knowldg of anirl divrsity thm, and th thin tissu
and his rathtakingrtivity' did rrltnvin th ritis d, in f;.rt,dls from th watr into th
ntlt lrrd itslf to a rodrn vliuti<;naryirnalysis.Th spisinvllvdin the drivd from th gill a
argumnt ar too distantly rlatdtO on nothr for skptialt:rsts; th fssil gasp and pross fod
ord arrd th inrpliatinsf omplrrativr-nolulardta slrgstthat it hs n h a n st h t r n s m i s s
n tns of millions of yars sin many of thm shard ommon anstor on th outr surfa of 1
(prsumirrgthr on vn gir'srlno th ida f vltrtion).Th vllu. gion of our skulls'
ti>naryPathway to th Otoplls y has long sin n lost. ]Wwould hv t-to ow an suh divrs
bttr luk trair-rgth pthwirv to tlr vrtrt1';ntrnlrv tlffrs larg rats, volutionaril
group of organismstlrat all lravwll-dvlopdyspt fo thos living in fish ar prt Of a rp
vry drk nvir<lnmtrts that sl to hav son]rilvolv]rdud ys. musls that opn arrd
vn in vtetsth progrssiolrfrom simpl to trtllplys ourrd irr that pand and ontra
som unknown st of ilnsttlrsthat livd ovr 400 milliln yrs ag(). arhshav th funrio
Th lilrnrafor rrndrstarrdingth vtllutiol-lof tllplity is that it o- grasping and grinding
urs on a timsal that is vry long onrpard rvith th timsal on whih t h t r a r l s f o r m a t i o nf o
mutationl vrition irl sirrglhirratrriss or natLrralsltionhausts so dramati as to s
th vailal variation in a singl haratr. But vn whn w rsort to hangs.
brad ornprisons <l indit infrns,it is oft tlr s tt w arl Th trnsformation ]
mak slmvry strong infrnsin partiular ss.!7 rviw two lf thos is lss dramti than it
ass :rusthy illtlstrt htlw ,an omin dt fr<lnldiffrrrtlif- spis.Th fossil ro
sin disiplins to dviop insights int<lth tlrigin of or-nplstuturs' isms that rprsnta tl
vn thLlghthis origin is not dirtly osval. th rptils tl-ratpp
omposit of svn ol
mpl 1': Fro gill rhes to er ossiles d js skull was formd by t
An old infr frtlm th ftssiltlrd is tht vrtIit jr'vswr drivd on on th skull (Figr
from th gill rhs tlf j:rwlss fish (rprsntdtoly y lan-rprysnd dntary, and artiula
dpttion ft/
tion ould produ onrpl adap- hagfish).A sond infrn is that th singl middl-ar ossil of amphi-
;rnallstps,h t>fwhih hd sllr.r ians' ptils,nd irds rvas drir,dtrom an lmntof th jaw suspnsionof
r volutionof th y ould only fish, and th additional twtl middl-ar ossils of mammals wr drivd
]qunlf rhssmtrll hnges, lrd fronronsssoitdwith th |aw joint of tir rptilian anstors(F-igurl )'
t dos rquirsan undrstandingof hswould rmarkal voltionary vntsin whih mutation and sl-
Dw2004)' To th modn vrllu. tion drov th wlrolsaltransformation of haratrs' molding ntirly nw
.htlrrtingrrt
and lristoril naturs shapsand rating ntirly nw funtions (!7ak 1979; Radinsky 1987; Luo
fgumntsft, IntlligntDsign' 2007\.
y y pointing ollt th divrsity lf But th diffrns in funtion rrrak sh a progrssion sm fniful at
- . i so f g s t r o p o d s( s n a i l s 'm u s s l s , st.Gill ahsar omposit of ons, musls, and nrvs that funtion
ang from lustrsf lls in som as a llows. Fish us thm to xpand and ontrat th volum of th oral
ht nd drk to th spistitd, .!7hn
vity. a fish opns its mouth, th llows xpands, th volum of th
n g o f p h o r o n s i t i vor g l r n s l i i n - oral vity inrass,and wat is drawn irr. !7hn a fish loses its rrrouth,th
argued y analog1',asstin]tht bllowsontrats and rdusth volum of th oral avity as th fish pls
rgans in gstop()dsillustrats th Wtrthrough tlr gill slits. ln living iawlss fishs(and prsumaly in th n-
tor to dsndantt uild th most storsof all jawd vrtrats),this pump is a fding dvi' drwing wa-
y, rvith a omprrlpossiiliy trornpartilsinto th mouth that adhr to a stiky sustanprodud
.gumntsrnall
hrrgsagainst th. in th oral avity, whih is thn onvyd to th gut. Th ntir apparatus
n aumult ir-rtoa rmarkrl also funtions s a rspiratory orgn. Th tissusthat lin th gill ahs ar
wll supplid with lood; th pump passs frsh, oxygntd watr ovr
3raknowldgof aninrzrldivrsity thm,and th tlrin tissstlrt lin th arlrsallw oygn to trrrsportd
nvin th ritis and, in fat' dos from th watr into th lood vssls.By ontrast, th struturssupposdly
rrlysis.Th spis irrvllvdin tlr drivd frol th gill rhs lrav mrrr iumsrid funtions. Th jaws
nothrfor skptiaItasts;th fossil grasp and pross food. Th first rniddl-arossil in trrstrialvrtrats
nrtllulrdt sr-Iggst tht it hs nhansth transmission of th nrgy in sound wavs fronr th ardrum,
f thm shad a ommo anstor on th outr surfaof th ody, to th innr ar, whih is in th tmporal r-
l th ida lf l.oltrtitln).
Th volu- gionof our skulls.
g sine nlost. ! would hav no L{ow an suh divrs struturs'not all of whih ar prsnt in all vrt-
'rtl.;nttlr nlu,offs
a lag rats, volutionarily linkd to ol1anothr? Th iaws in th aly jawd
iopdyspt1thos living in fish ar prt of a rpatd sris of ons that inluds th gill arhs. Th
' slndariiyvolvlrdued
ys. rnuslsthat opn and los th jaws funtion in a similar fashitln to thos
iinrpl to ornpl .ysourrd in that pand and ontrat th gill arhs.In som living spisof fish, th gill
'vr400 million yarsi1go. arhshav th funtiollal quivallrtof tth and funtion as do our jaws y
lluti<lnof omplity is that it tl- grspingand grinding food. Givrr th pattrns of musl nd innrvrion,
rard rvith t timsalln whih th transformationfrom gill arh tO jaW suggstdy th fossil rord is not
rrissor ntural sltiorrxhausts so dramti as to sm an unlikly urrrulativfft of small, attain:rl
:tr. But vn whn w sot to hangs.
, it is oftrrth lls tht w n Th transformation from jw support in rptilsto r ossii in mar.rals
ulr :rss.\/rviw two of thos is iss dramati than it might appr from mparisorrsof only th isting
on-rindilta flr diffrnt lif- spis.h fossil rord offrs striking pattrns of lrng aross th organ-
' th origin of ompl srruturs' isms that rprsnta transition twnrptils and mammals (Figur 1). In
rval' th rptils that appar to anstrl to rammals' tlr lowr jw ws a
ompositof svn ons and th artiulation twnth lowr jaw and th
ls lld itus skull was formd by th artiular bon on th lowr jaw and th qudrt
that vrtratjaws wr. drivd onon th skull (Figur 1A). In mammals th lowr jaw is a singl bn, th
sntdtoda v lmpys irrrd dntary, arrd rtiulation of th j. tlss th dntary on itslf nd th
1 1 8 dpttitl
D{ondy| sqrramosal on on th
a0uI
delhmeil
t/-
sids (th sintifi trm
jaw hings that onsist
surangu|r
squmos|
I
middle
th lowr jaw and th q
on!at
Adutt
ear
1B_). This dual artiu
quadrtojugaI
. Monodlphis faturs of th sklton,
mbryoni rptils and mammals. I
and squamosal ons fil
{= rplad th artiular ar
=_ on th artiular an
o
of jaw and skull, thy v
fossils nd from th ml
mals indiats that th
and orintation' and
l<
l
I
| I
) o () Brsilith rim rep
.
; Triassi through th lat
3s.*. Pchgnlus (B) but rt
D
(D) Sinocoldon was '
from th arly Jurassi,tl
q though th individualo
f
GI (Lft) orguo
late Triassi thrugh th
a 8 r a t l yr d u dp r t i
To th right of ah ol
tht th stapdialpros
prossrtiulatswith t
( ) ( R i g h t ) T h i si s a v
D.ond|e
orgucodoz, illustrt
SO-glenoid
srisof ons inluding
quaoratolug|absenl
bons hav a diffrntn
animals and thy an ls
Figr-rr1. This figur, modifid from Luo 2007, illustrats th volution tlf th
totympaniis a ring o1
manrmlian jaw, livlfrom :r sris of ftlssils in th synpsid linag (th linag irl
twn th outsid wor
wih mmm:rls mrgd) nd fro.l th mrylni dvltlpmnt of t living marals.
(F) Ths ar adult an
Th lrawings on tl-rlft (A-D) rsnt th l<lwrlfi vrrtral (Ottom) suri of tlr
rnitb orhchus. |n te
aniu.t,as illustrtd in th drawing in th low-lft orn at th otttlrn of th fisur.
mdial surfaf th lov
Th figurs tln th right (_H) rprsnt th mdial (irtnr)nd vrrtral surfas of lr.
totympani,mallus'
ltlu.r jw. h numrs rprsllt rltiv ags, wit tlr lowr numrs indiaring oldr
onfiguration sn in th
flssils and th highr numrs inditing rnt mammals.
(G) Th lowr drawin
() Prlirlgthlls ws iat Triassi Synapsid tht is r.rotin tl-rdit
anstral lin adult jaw of Ylood
to mammls; thr was a tl.rplartiulatitlr.rof ltlwr jirw nl rniunr involving th
.,mllus'' Th ondition of th
(tl-rrodifid artiLllr orr) ln th low jaw and tlr ..inus'' (th rnodifid
duk-illd platypus.
quirirt orr),quirdatojrrgal,d squamosl ons of th uppr jw,.
(H) Th lowr drwin
(I\| h1'gelsrpsnts a difrrrtrnh of th Syri;lsids tht dats tl th lt
oPossum' ol delph us
Trissi_rlJurassi priol. This group ht-rd duill rtiultion in whih th ..n-ra]lus',
mryoni jw is vry sr
On th loWr jirrv artiultd with th ..inus'' on th Llpr jaw and th dntary on th
Ynoonldoz, with th
lowr jaw artirrlat1with th squarosal on th upp jw. Thr Wils l1o 1uadrtljugal
th lowr jaw, whil th
on.
kl lmntsing
dpttio 119
into what is now alld th rallus and th quadrat into what is rrow alld m u s l s a s s o i t dw i
th inus. drivd from what is
Th drivr for this transformation would hav n natural sltiln'sfa- nrvs that srv ths
voring nhandaility to dttvirations. Rsarhsuggststhat th jaw in nrvs.
ths transitionai animals transmittd virations from th ground to th mid. All ths osrvation
dl ar, as it dos in som modrn snaks-think of a small, nlturnalanimal r h sa n d t h t t h m l
that pikd up virations y touhing its jaw to th ground and transmitting mutation irnd sltion
virations from th jaw to th staps.Ths virations would hav travld ponnts. But hw nd
through th artiular (mallus) and quadrat (inus) ons to th staps.In \|977) on dsri
two linags of nonmammalian Synapsids, nithr of whih is anstral ttl nring; a tinkr uss v
mammals, th inus has an outgrowth alld a stapdial pross' whih is a tions, whil an ngin
lvr arm that transmits th for of th sound wavs from th inus to th that adapttion is a hrs
staps. Its prsn in ths two linags stlggststhat oth of thm wr s t u t u r sf o r n w f t l n
volving an nhand paity to transmit sound from th jaw to th staps (ffiint feding) and fl
and thn to th innr ar. into largr, mor ativ
Th arrangmnt of btlns in th linag that inluds mamrnals is diffrnt tant funtion of th ph
than in ths othr linags ut dos inlud n inus with a stapdial pro- a n a s s o i r i o nw i t h f .
ss.Tlr final stagof ours is th sparationof th ossiIsfrom th jaw it- pharyngal pump hav
slf. \/hnth ossils am sparatdfrom th jaw' thy ould transmit los th jws, so thy
sound viations ffiintly from th adrum to th middl ar. Th om- funtion.
plt fossil rord rvls that, rathr than thr ing a dirt transforma- Although th transit
tion from the primitiv synapsid lndition to th mammal ondition, thr drstand, th assoiatt
was a divrsity of arrangmnts lf ns in th jaw-ar rgion and a divrsity trat.Stik your fillg
of rarrangmntsovr tim. Th anatomy f living rptils nd mammals will s tht th jaw j
rprsnts two ndpoints of transformation that followd divrs paths in you an fl th joint r
twn,with most of th variants having gon tlnt. v r t r t s .i n | u d i n g:
Although th fossils Suggstth progrssionof fatursin this transforma- p tlpik up vi
tion, dvlopmntal iology offrs us twl important additional lirrs of vi- and thn to transmit th
dn. First, whn th ossilsfirst ppar in mammalian mryls,thy ar snaks and small mamr
part of th jaw assmly.During dvlopmnt'thy sparatand f<lrmth fa- t i o n s a r n o t u r n |, r n
miliar struturs of th middl r (igur 1F-H). Whil th notion tht an m n so f t h j w s t o l
organism's dvlopmnt rapitulatsits vrlutionaryhistory has n dis- l a r g l ye o n s q u n
rditd in gnral, this is a as in whih dvlopmnt offrs a lu to an th prior rl tht th j
volutionary history that is othrwis traalin th fossils. vironmnt to th innr
Sond, gill arhs, jaws, and ar ossils hav a ritil rlationship to
on nothr that sets thm apart from othr faturs of th skull. Th vr- rple 2: FIorI ul
trat ody is a omposit of thr diffrnt layrs of tissu, th todr- our sond ampl, v
mal (outr), msodrmal (middl), and ndodrmal (innr) layrs. Ths nti analyss ontribu
t h r l a y r s a r a s i l y d i s t i n g u i s h df r o m o n a n o t h r a r l y i n d v l o p m n t t a t i O n s( S h m s k n d
ut om lndd in many parts of th ody as dvltlpmnt progrsss. w fous on shp and
Most of th vrtratskltlnis drivd frtlm th middl, Or msodrml' rs flr thir auty n<
lyr of tissu. Howvr, th ons assoiatdwith tlr gill rhs, jew, and rprodutiv strutur
middl-ar ossils hav a omlnon dvlopmntal origin diffrnt from polln (th mal gam
that of most of th sklton;ths ons ar drivd from spializdlls of mal gamt).
th outr, or todrnral,layr of tissu. Likwis, th musls nd nrvs Th llowrs that insp
assoiatd with ths ons ar rlatd to on anothr in thir origin; th animal pollintors likr
dpttion 121
h quadratinto what is now alld
m u s l sa s s o i a t dw i t h t h g i l l a r h s ' j a w s , a n d m i d d l - a ro s s i l s r a l l
drivdfrom what is alld te brhiomric sris of musls, and th
rld hv n naturl sletion,sfa.
nrvsthat srv ths musls all dvlop from th sam group of ranial
ns. Rsarhsuggststhat th jaw in
nrvs.
rtionsfrom th ground to th mid-
All thsosrvationssupport th rgumnt tht jaws ar drivd from gill
-think of a small, noturnal animal
rhsand that th middl-ar ossils ar drivd from jaws, and thus tht
j w t o t h g o u n de n d r e n s m i t i n g
mutationnd sltionan indd m<lldompl fatursfrom simpl om-
rsviratiorrswould hav trvld
ponnts.But how and why would suh a transition our? Frangois Jao
lrat (inus)ons ttl th staps.In (1977\ one dsrid volution as a pross of tinkring rathr thn ngi-
Js, nithr of whih is anstral to
nring;a tinkr uss whatvr is at hand and modifis it to srv nw fun-
Lllda stapdialpross,whih is a
tions,whil an nginr dsigns somthing nw from srath. Jao mant
sound-avsfrom th inus to th
that adaptation is a historial pross ausit involvs modifying xisting
s suggststhat blth f thm wr
strutursfor nw funtions. In this as gill rhs gn with on funtion
it sound from th jaw to th sraps (ffiint fding)nd failitatda sond (rspiration).As vrtratsvolvd
into largr, mor ativ animals' rspiration am an inrasingly impor-
that inludsmammals is diffrnt
tant luntion of th pharyngal pump. Th antriormost gill rhs rtaind
lud n inus with a stapdial pro-
n assoiationwith fding. Th ons' musls, and nrvs that pow th
ation of th ossilsfrom th jaw it-
phangalpump hav th sam ation pattrn as thos that opn and
from th jaw, thy ould transmit
losth jaws, s thy wr rdily modifid to spializin on or th othr
drum to th middl ar. Th om-
funtion.
ln thre ing a dirt transform-
Although th transition fr<rmjaws to middl.r ossils is harder to un.
)n to th mammal ondition, thr
drstand,th assoiation twn th jaw joint and haring is asy to illus-
n th jaw.arrgion and a divrsity
trat.Stik your fingr in your r and thn opn and los your mouth. You
ny of living rptils and mammals
will s that th jaw joint is right nt to your outr nd middl ar so tht
Ltionthat followd divrs paths in
yoll an fl th joint mov s you opn and los your mouth. Som living
] gon xtint. vrtrats,inluding small nturnal mammals, taans,and snaks, all
;sionof ftursin this transforma-
apparto pik up virations from th trnal nvironmnt through th jaw
l important additionl lins of vi-
and thn to transmit thm through th jaw t th middl and innr ars. Th
r in mammalin rryos,thy ar
snaksand small mammals that rly on this mhanism for piking up vira-
nnt'thy sparateand form th f-
tions r noturnal and do so through th tip of th jaw. Th ryling of l-
1F_H). \Whilth notion that an
mntsof th jaws to om sound-transmissionons in th middl ar is
volutionaryhistory has n dis.
larglya onsqunof th proximity of th jaw joint to th innr r and
:h dvlopmnt offrs a lu to an
th prior rl that th jw playd in transmitting sound from th trnal n-
lalin th fossils.
vironmntto th innr r.
ls hav a ritial rlationship to
thr fatursof th skull. Th vr-
mple 2: Florl euolution in moke flors (Ivimulus)
rnt layrs of tissu' th todr-
our sndxampI,whil diddly lss swpingin sop, rvalshow g-
ndodrmal (innr) layrs. Ths
ntianalyssontriut to undrstndingth arhitturof ompl adap-
on anothr arly in dvlopmnt
tations(Shmskand Badshaw 19991'Bradshaw and Shmsk2003). Hr
: ody as dvlopmntprogrsss.
w fous n shap and olor of flowrs. Although vryonappriatsflow-
l from th middl, or msodrmal,
rs for thir auty and fragran, from an volutionary prsptiv thy ar
iatd with th gill arhs, jaw' and
rprodutiv struturs that ar spializd to produ and dissminat
llopmntal origin diffrnt from
polln (th mal gamt)and riv and dlivr polln to th ovul (th f-
Lrdrivdfrom spializdlls of
malgamt).
Likwis' th musls and nrvs
Th flowrs that inspir our gratstafftion ar thos spilizd to attrat
:o on anothr in thir origin; th
animai pollinators lik s o hummingirds. Ths flowrs ar atually
122 dpttitl
ompl ominations of haratrsthat togthr ar an adaptation for dis- the flowr. Th flowrs
prsing and ollting polln. Flowrs pollinatd y animals are ompara- produtiv organs onta
tivly larg' olorful, and shapd in ways that failitatth transfr of polln. thy ar nountrd y
Som flowrs ar highly spializd to attrat spifi pollinators, usually y of th insid of th flow
a olor and/or fragran that r partiularly attrativ to a pollinator. Th flowrs that lak th hol
sugary ntar is th rwrd offrd y th flowr to th pollinat,or, a rward y hummingirds, whil
that draws th pollinator's intrstto th insid of th flowr and ststh pol- du larg quillltitis Of
linator in pla to pik up and dposit polln. any flwrs hav mrkings nthrs (ml org:rns)t
that dit th visitor toward th ntar,som of whih a visil only to an- that polln is dpositd.
imals lik insts that s in th ultraviolt (lak-light)frqunis.othrs stamns (fmal ()rgan
hav modifid ptals or spals tht srv as a soft of landing strip on whih t h r nn s r h i r d f d r .
th pollinator an st' nd som spishv spializdstrutursto attah ah flow is wll m:
th polln to th pollinator. diffrns twn tlr
Two losly rlatd spis of monky flowrs, imulus letaisii and . pl alptations for p
rdilis, our along riparian hitats in th wstrn Unitd Stats (Figur 2). math of florvr tl polli
Thy sgrgt y lvation, with . ltuisii ourring at highr lvations, Organs' and ntar pro
and thr is narrow rang of lvation in whih th spis ovrlap. Gnti and sltion ould hv
analyss show that ah spiesis mor losly rlatd to th othr than i- a Ommon nstor. O]
thr is to any othr spis of monky flowr, so any diffrns etwn ussd arlir tht gn
thm volvd aftr thy last shrd a ommon anstor. diffrns rprsntth(
Th two spies hav quit diffrnt flowrs. Th spis at th highr l- hratrs. lf this wr
vations' . lisii, is pollinatd primarily y s. It has pink flowrs that plind vry simply
ar rlativly wid and whos ptals form horizontal landing strip; yllow t th two flowrs; shi
struturson th landing strip appar to guid sto th appropriat part of nario, not srrstantially
mottld to lak. At t
mati diffrnsar a
ah of whih has vry l
or lngth of th anthr
n v y m t i u l o u sa l
th floral traits. Of or
trm vlWs.
O u r k r l o w l d g f p l
p t i n g h y p o t h s s .A l t t
of spals, ptals, st:rm
g n s 'i t i s u n l a rh o w
of th rprodutiv str
dotion rat.
Th rout to th ans
nd thir ollagus y
Figur 2. Studisof th olor, siz,shap,and fatursof th flowrsof two
lular mthods (Sh
loslyrlatdspisof imulus hv illustratdth gntintrol of
2 0 0 3 ) . S h m s k a n d i
rmpladptations.Lft: Te lu flow of . letuisii,whih ftursa
..landing pltform'' usd ,v Mndl whn l
for th inststht transfr polln twnflowrs and
..guids,' tnth ntury. Thy pr
that dirt th inststoward th ntarand th rprodutiv
strutursof th flowr. Right:Th rd flowr ot . rdilis,whih is a long, rids rarly our in n:
narrow' tuular flowr with ntat th as,rahaly th tongusof th ar quit hardy in gr
hummingirdstht srvas th pollinators. lagus nt rlssd h
lza Ad|lttil
sond gnration of so-alld krossd plants with a varil mix of A fundamntal tnt of I
gnsfrom ah original spis.Thy thn turnd to molular iology' m- trs that nhan th fitn
ploying molular markrs tht enald thm to ssoiatptiular varia- wll-ing of spis.I:
tions in floral lraratrsas sn in th vrious orninations of plants with in prtiul:rr for th g<l
diffrnsin molular maks on spifi lotions of hromosoms. Th gon from sintifi dis
diffrnson th hromosoms rfltth loations of gnsthat ontriut nfit to th individu:rl.
to th vition in th flowrs' But altht>ughthis may
Twlv individual haratrswr idntifidarrd studid with this mthod, planatit-rfor all fatu
whih rvald that th truth rvas indd somrvlrr twn th wo x- dividual sltilnoms
trm ids :rout gnti ontrol. A singl grr dtrmind whthr th m i t s ,i n w h i l r n a r l ya l |
flowr was d or pink. By plaing f.lowringhyrids and akrosssin na- It is diffiult to onstu
tur, Shmsk and ollagusfound that th olor diffrnlon had an individu| with :r fitnss.
normous impat on whthr th flowr would visitd y a or a hum- not prolu.
-I.h Th ky to rsolving th
mingird. sintists also found that whil anothr singl gn mad a
larg ontir.rtionto th diffrnsin 9 of th 12 floral traits, thr wr that th ol<lrryworkrs l
many gns tlrt mad smllr ontributions to on or a small sust of th th srifi in individua
traits. rais th offspring oi .l l
From thsrsultsW an offr som plausil ronstrutionsof this om- ommitt-t-lllt'Thr ar
pl adaptation. Th fft of flowr o1or o visitations suggststhat a from th ooprtiv ra
hang in a singl 8n' that fr olor, ould initiat th volution of a sp- providd v oldr offspr
ialized flowr. Diffrnt pollirrators attratd to th diffrnt olors would hirs n <lsrvdin sv
rt diffrnt sltionpressurson th rmainirrgtrzrits.Although it is un- Ths ass :rramp
lar whtlrr th larg sirrgl-gnefft on nin traits prddo followd tiorr. In kin sltilnindi
th hangs in th many othr gns that afftd on or a fw traits' th dution lf rlatd individ
ovrall pitur is of a singl, simpl gnti hang (olor) inititing a stp- omination trf numr
.his
wis pross of modifiation and rfinmnt whos nd produts ar th two th srifi. is a por,
..SlialBh
omplx adptations that w now s.Th math of fatur to irumstan main ssy
..
was not built qrritas asily as hanging th oloration of moths ut did not th alphatial ntris
rqui so mtiulous a molding as to strain duliry' I n f l r t ,k i r r s l t i l l rils
A finl not out th historial natur of irdapttion is that it dos not for th lvl t whih th
sm pssil that th hangs in floral shp and rpodutiv haratrs thr is i1 usal rlations
.!ithout
ould hav prdd th olor hang. variation in olor nd th r- suh as th family lvl,
sultant diffrnin spisof pollinators, thr would hav en no onsis- ilis or lrgr groups f
tnt sltion prssurto mold divrgnt vlus of th othr htrs.But goup sltior-rto our
on th olor diffrnSppard,tlr divrgnt foraging of s and hum_ phistitd thoris, n
mingirds proaly ata|zeda rpid divrgn'This ampl illustratsth to diagntls how oftn t]
ontingny inhrnt in th volution of omplx faturs in whi a singl thn that of th individu
hang in a fatur an promot asad of susqunt hangs that rfin A wid varity of {a
its adaptation to nw irumstans.To rturn to th ritiqu bf Intllignt othr thn that of th rr
Dsign, wihout appriatingth rol of history, w would hrd prssdto rsarh.Bsidsth ml
plain how n{why ths flowrs divrgd so sulrstntially. of grup living arrd
h a v i o r r lS o i o i l o g
..Group sltion'' in th
W H A A N I N D l V | D U A Ls L | o N N o X P L A | N ?
al rol in undrstandin
Our disussion of adaptation hs mphasizdhow th ftursof individr'rals a n d h l s t s( s t h m a i l
ar moidd y th ation of sltion on th avirilal gnti V;rrii1tion. winin diin'' y \
dpttio I !.\
d plants with a varial mix of A fundmntltnt of Dwinian vtllution is that sltion favors hara-
l turndto molular iology, m- trstht nhanth fitrrssof individuais; it dos not nssarilnhan th
hm to assoiatpartiular varia- wll-ingof a spis.Indd,th notion thilt volution hs d<lnanything
rious omintionsof plants with in prtiul:rrfor th good of th spisis, as fundrntlpinipl, long
fi loationsof hromosoms. Th gonfrlrnsirrtifidis<lurs; arry nfitto th spisis silfftof th
: ioationsf gnsthat ontriut nfitto th individual.
But ltlrtlr"rgh this may .a usful first prirlipl, it annot th Orrt
ifidand studidwith this mthod' planationftlr all Aatursof organisms. Tlr mtlst striking lrzrllngttt ill-
somwhr twn tlr two - dividual sltitlnoms from th s<lialinsts' ants' Wasps' s, and tr-
gl gn dtrmind whethr th mits,in rvhih nlrrly all mmrsof th olony forgo prodution ntirly.
n g h i d s n d k r o s s si n n - It is diffiult to rOnstruthoW sltioll 11l-t1()tl] individuals wluld far,or n
th olor differnalon had an indivrdulwith a fitnsslf aso|utzro, whih is wht happns if on los
'ould visitd y a not rprodu.
or a hum-
whil anothr singl gIrmad a Th ky t<lrsolvirlgtlris parado fr s'ats, and wsps is to apprit
of th 12 florl traits, thr wr that th olony w<lrksar raising thir nis and nphws. It appars that
|nsto on or a small suset of th th sarifiin in]ividualfitnss is ltpnsatdy th nhnd ailit to
ristlr offspring f siling, providd that rlstindil.iduls sustain this
usilronstrutionsof this om- ommitmnt.Thr ar many anrpls of lss trllforms of ltruism,
lor olr visitatirlns suggsts that a from th ooprtiv raring of offspring y sistrs in lion prids to th id
uld initit th voiutirnf a sp_ providdy oldr offspring tl thir pntsirr raising y()ungroffspring, as
td to th diffrnt olors would has rrosrvlin svrlspisof irds.
mainingtraits. Although it is un- Thsssar ampls <lfsltiont th lvl of th family, or kin sel.
,n nin traits prdd or fl]lowd tion. In kin sltiorrindividul sirritiis firr,ordrvhrrit nhnsth po-
' afftd on o a fw traits, th dutionlf rlatd individtra[s'with th dgrof saifiproportional to th
i h n n g ( o I o )i n i t i a t i n g s t p - omintionof numr and rlatdr-rss of th individrralsthat nfit from
rt whos rrd pduts irr th two thsarifi.This a polr'rfulplanation filr nrirn1'fornrsof soiaIity(sth
is
: math f fatur to irumstan main ssav ..Soil Bhvior and S<lioiollgy''y Danil Runstin and
l oloration of moths ut did not th alphatialntris ..Altruism'' and ...W.D. Hmilton'' in this volum).
n dulity. In fat, kin sltion is or-rlf svrl vaitisof sltion tht ar nnrd
, of adaptation is that it dos not for th lvl at rt,hih tlry our. T'hsvaritis of sltion mrg whn
hap and rpodutivharatrs thris us:rlrlationship twll fatursprssdat a partiuiar lvl,
horrt vaiation irr olor nd th r- suhas tlr familv lvl,anl tlr sussof th units ar tht lvl. thv fnr-
thr would hv n no onsis- ilis or lrgr groups of r-rnrlatdindividuls. Th onditions for kin nd
,alusof th otlrr haatrs.But group sltiont() our hv n xplord xtnsivly in a v:rrityof so.
rgntforging of s rrd hum- phistitdthl>ris, nl h ritizrlissu in nlodrn volrrtionaryiology is
gn.This ampl illustratsth to digrroshow <lftnth nditiolls that promot sltion t lvls thr
omplx faturs in whih a singl than tht of th individual irr mt.
of susqunthangstht rfin A w i d v r i t y t l f f t l r sa p p r l i a l o n l v 1 , s l t i o I - lt l v l s
turn to th ritiqu f Intllignt o t h rt h n t h a t O f t h i r r d i v i d u l ,i r r r dt h i s i s v r y a t i v t o p i o f m o d r n
;tory' W would hard lrssdto r s a r hB. s i d st h m a n y a s s o f r p r d u t i v l t r u i s m , t h r r : r s s
.d so sustntirrlly. o f g r o r ' r pl i v i n g a r - r d o o p a t i v h v i o r ( s t m i n s s a . . S o i l B -
h v i o ri r n d S o i < l i o l o g y '' y D a n i l R u n s t i n r r d t h l p h a t i a l r r t r y
. . G r o u ps l t i o n ' 'i n t h i s v o l u m ) .
S l t i o na t s v r a ll v l s p l a y s i t i -
oN No PLAIN?
a l r o l i n r t n d r s t n d i n gt l - r v o l u t i r l r r a r i n t r t i o n s t w l -pl a t h l g n s
d how th ftursof individuls a n d h o s t s ( s t l r m a i n s s a y , . v < l l u t i l n a rByi o l o g v o f D i s s a n d D a r -
l t h a v a i l a | g n t i v r i a t i o n . r v i n i a nM d i i n ' ' y i h a l A n t o l i n n d t h a l p h a t i l n t r y . . H o s t
1 2 6 dDttil
parsit volutiot-t''in this volum); a pathogn tlrat nrultiplis too quikly s that thr ar long
nd kills its lrost tlo quikl for it an b trr-rsmittdto nothr host not at ll, thn rif in
loss out aus sltion among familis Or groups (rprsntdy th drdg dsrid th l<
pathogen popultiorr within an individual host) ovrwlrlms sltion short intvals as ptrnl
a m o n g i n d i v i d u a l s ( p r s n t d y t h s t r u g g l f o r p r o l i f r i r t i o nw i t h i n n ssoition twn pu
individual host). th trm ptltuted q
Gotrld rgrrd that ot
with nturl sltiona
AR sUDls oF MlRoVoLUloN RLEVAN o
on th prmis that if s
U N D R s A N D IN G M A R o v o L U | o N A R Y P A R N s ?
a prd<rminantPattrn
[t is onvnint to hartrizth produts of evolution as ithr mlrovo- haratrs suggsts rs
lution or marovrlutirln.iroeuoLutioll rfrs ttl small-sirlhangs that suggstdto thm that l
ar dirtly osval, lik many o{ thos distrssdlir (mlanism irr a susqunt puls lf a
moths, disas rsistanin atri, or instiidrsistan).As w hav Although Gould nd
disussd, mirovolution has n vry thorouglrly dourntd and has that nsud aftr thir {
n studid irr ways tht mt any tst of sintifirigi<lr.As rsltlt,it is nr First, neturl sltion
ontrovrsial,vn among n-ranyfaith-asdritis of volution (sth main isms to small-salhar
ssay ..Amian Antivolutilnism: Rtospt and Prospt,' y ugni tlran natural sletion
Sottin this vlum). qunt assoiation tw
crleuollttio rfers to th Iargr-salvnts tl,ratw attriut to volu- suggststo Gould and
tion, sulr as tlr origin of nw spis,th diffrntialprtllifrarion or diffr- phnomna, ut thy h
ntial tintion of rtain typs of spis, and th mrgn of highr d u r i n g t h 3 6 - y a rh i s t
lvls of th txonomi hirahy.This is th aspt of volution that is on- Th ountrargrrm
rovrsialusit rprst-tts vnts that otrr on a timSllltht is -ruh hs thr parts. First, v
longr than our livs :lnd hn not dirtly osrval. As w hav dis- tion y ntural sltr
ussd, sintistsmust turn to indirt mthods to draw infrensaout th thn th rats i]ssolat
undlying mhanismsthat aus marvolution. G16pags finhs an
Th most rvolutionary ftur of Darwin's th rigi of Spcies i n a p l p u l t i o ni n s i r
(1859) was to popos naturl sltion as th singl unif1'ing mhanisrn fall. In dought yrrs
tht auss oth miro- and marovolutiln.Drwin argud that mrovo. sdswith thik sl<
lution is just mirovolution writ larg, or that th possW s nd study ftiv at harvsting su
as th aus of mirovolution will, givn suffiit.lttitn, lso aus vry- Baus tlrr is a gn(
thing that w attriut to marovlution. H ergud thi.rtnrural sltion, finhs xhiits largr,
whih alrssth volution of th ad:rptationsdisussdthroughout this s- to put this rat of vol
say, is also rsponsilfor th origin of :r1llvls of biologial orrrplityand sil rrd' it is l0,000
for th origin of iologial divrsity, or all th spisthat hav n found tprt as puntuation
on th arth thoughout its history. inonsistnt witlr tlr r
otrr inaility to osrvmrovoltltionin tion has inspid ontrovrsy tion' [f anything,w r
ovr whthr ntural sltionrally is th unifying mlrnism of all volu- fossil rord.
tionry hang. StphnJ. Gould and Nils ldrdg ar th uthors of wht S o n d , s l t i o l li s
is urenly th most prominent sintifirgulnnt against ntural slti<ln This, of ottrs, is tlr.
as a singl, unifying aus of miro- and nrrovolution (ldrdg and ut hr it is importan
Gould 1972; Gou\d and ldrdge 1977). Ths authors Wr motivatd y tion an onl mak fi
th pttrrrof hang rordd in th fossil rord. If w look t th fossil his. tn to hang, mltst
tory of th volution of most orgnisms, suh s th inrasof ody siz in Darwin's lins of argu
horss or rain siz in th hominid linag lading to our spis,w o[tr-l t that domsti dtlg
dpttkl 127
argumnt in road outlin, it fll to his sussors to disovr whih lmnts aFddn, B. J. |992. l"rl
the Fmil quidc. N
of his argumnt Wr possil in thory, whih wr dmonstral in pra-
N{Nair,. R. 1987.Ha
ti, and whih wr roust to varitions among spis and irumstans.
sstm' rtJs ilt tll
T'hat so muh of his original rgumnt mains standing is a triut to his g- tsurnura, I]', and G. Vss
nius; that so muh rmains to disovrd is a triut to th mrvls that i n h r r , r ' t l - s p t t sJp i t
adaptation offrs and th divrsity of natur th:rt it has atzrlyzd. tomtllog 57: 9| |-9
N,lPk,. ,\. l99.5. or1
damslflis rf tw<ltltlt
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rd is a tribllt to th marvls that in th rl-sptlttdspidr mtt trln't.hus rti. !lLtrl of L.('|lo||1i
-urthat it has irtlyzd. tomologl' 57 : 9 | |_9 |7 .
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R z n i kD
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P U r p Ibe t e i
FrnisoJ. Al Flavobatria
Thermotog
|es
Figur 1. Th univrsaltr
Molular biology hs mad it possil to ronstrutth ontinuity of su- Batria,Arhaa, and uk
ssion from th original form of lif, anstral to all living organlsms' to oganismsar rlatdy o
vry spisnow living on arth. Th univrsal tr of life, whih mrs y th tr's..trurrk''(thst
using slw-volvingrilso
all known sorts of organisms, inluds thr sts of ranhs' Batia, Ar.
haa' and ukarya' whih all mrgd from on form of lif, LUA (Last
Univrsal ommon Anstor; Figur l). Al1 living organisms ar rlatd y
This has mad it possil l
ommon dsnt from a singl form of lif, rprsrrtdy th tr trunk in
ously unknown and to or
th figur. Lif may hav originatd mor than on on our plant-w
Th prision with whih l
prsntly do not know-ut only on form of lif prosprd and lft dsn-
vidn from molular b
dants, th organisms that nw populat th arth.
h molular <rmpo
Batria and Arhaa ar prokaryoti mirosopi lrganisms.ukarya ar
natur of th omponnts
organisms with omplx lls that ontin svral orgnlls,on of whih'
n d u s d .I n a I | a t r i , r
th nulus, inluds th hrditry DNA. Most ukaryoti organisms r
sists of a diffnt squn
singl lld and mirosopi. Animals, plants, and fungi ar multillular or-
sntd as A' , G, and T; Fl
ganisms, thr of th many rnhsof th ukary st.
ontaind in th DNA of t
h main ranhs of th univrsl tr of lif from LUA, svral illion
th sam in all organisms
yars ago' to th prsnt hav n ronstrutdon th whol and in many
sorts of organisms ar syntl
dtails. Mor dtails out mor and mor ranhsar pulishd in sors
varial lngth, of th sam
of sintifiartils vry month. Th virtually unlimitd volutionary infor.
a l h u n d r d o r h a m i n i
mation nodd in th DNA squnof living organisms allows volution-
of biohmial rations-
ists to ronstrutany volutionary rlationshipsthat hav ld to prsnt-day
nrgy and to mak up th
organisms, or among thm, with as muh dtail as wntd. Invst th ns-
This unity rvals th g
sary rsous(tim and loratory pnss),and you an hv th answr
ganisms. Thr is no oth
to any qury with as muh prision as you want.
formity whn numrous
likly. Th gnti od st
thr nulotids in th u
olular volution: Uniformity and Divrsity atly th sam amino z
t h a n i t i s f o r | a n g u g t t
In its unviling of th natur of DNA and th workings lf organisms at th snt patiu|a objr.
lvl of nzyms and othr protin moluls, molular iology hs shown plnt, tr' woman-ar(
th unity of lif. orovr, molular iology has shown that ths mol- diffrnt oks, on an b
uls, DNA and protins, hold informtion aout an organism's anstry. of ommon origin.
t.)z
olulr uolutio l
Pyoditium
F|g||ates
Iavobatria
rihomonads
Thrmotg|s |\4irosporidi
DipIomonds
.''io
8roov
*j U
Ph
UU'l U
c
U
*^.l Leu
U
UUGJ U
o
o
! l phogPht str
t*r
U |
o *^l
id N ln ss
I Leu
L
*"1
AUUl
y)
AU l
I lle
I
^u^
AUG et
GUUI tl
GU l
G I val
GUA
I
.u"J
igur 2. Th doul.stranddhlialnfigurationof th DNA molul.Th
Figur 3. The gntiod:.
molulonsistslf two polynultidhains.Th outward akonof th
mssngrRNA and th am
molulis mad tlf altrnatingdoyriossugrs(S)and phosphtgroups
nitrogn as thymin dos
(P).Nitrlgnassonntdto th sugrsprojttowrd th ntrof th
th othr thr nitrogn as
molul.Th two nulotidhains r hld togthry hydrognonds
adnin (A)' ytosine (), aI
btwnomplmntirrypurin-pyrimidinss.Adnin (A) nd thymin (T)
protins (with th thr-lt
flrmtwo hydrognonds;ytosin() and guanin(G) form thr.
(Ala, A), arginin (Arg, R), '
(ys, ), glyin (Gly' G), g
( H i s , H ) ' i s o l u i n ( I l ,I ) ,l
)' phnylalanin(Ph,F),
tyrosin (Tyr' Y), tryptoph
Gns nd protins ar long moluls that ontin information in th s-
qun of thir olponnts in muh th sam way in whih sntnsof th
nglish languag ntain information in th squn of thir lttrs nd
words. Th squnstht mak up th gnsar passd on from parnts to th rational for th ror
offspring and ar idntial from gnration to gnrtion' xpt for o. nd outomswiIl rvi
sionl hangs introdud y mutations. losly rlatd spies hav vry As an illustation, lt ul
similar DNA squns;th fw diffrnsrflt mutations tht ourrd ooks are 2oo pags long
sin thir last lmmon nstor. Spisthat ar lss losly rlatd to on amination rvals that t
anothr hiit mor diffrnsin thir DNA than thos mor losly r- for word, xt that an r
latd ausmor tim has lapsd sin thir last mmln nstor.This is two ooks annot hav l
olulr uolutil 1''
*l
U A G
= UUl uu1 U
-".::&
UGUl
Phe
,,-- | I Tyr | y,
UU l ULL
I
UAl UG J
l&'
....c-
uu^l U UAA Stop UGA StoP
lu |
*ur
.".1
UUC J UG- UAG Stap UGG p G
'*-l
^,,
-" t*-l U
Fl
U | |
t
J
*^l l Lu l Po
*l
:::J
-r..4, A
l
J
Gln
- UGJ Gl Gl G
-l-
^Ul *ul ^"u-l
tW
AUU.l U
;i
U
I lle A | AAJ
Asn
AG J
Ser
|
l
^u^ ^.^
|
t^'
*".l Lys
Ac"l
Arg
A
UG et AGJ CJ AGGJ
G
GU l
cuA I val "tt I ^"
G
GAJ
GAAI
l 'p
""'
GGI",,
A
I | I clu l
P.ent GUG- GG- GJ G6GJ G
opid fror th othr, lr <lthhav n opid, ditly or indirtly, from stalishd whn th squt
th sam oiginal ook. ln living ings, if ah omPoilnt nulotid of diffrnt organisms. Th s
DNA is rprsntdy on lttr,th omplt sqtlnof nr-rlotids in th vn twn vy diffrr
DNA of a lrighr organisnr ,,orrldrquir svrl lrudrd llks,ah with partiv anatomy' for
.Whn
hundrds of pags, witl.r svral tlrlusnd lttrs tln irh pg. th pin trs, and humn i
..pags''(or squnsof nulotids)in ths ..oks'' (gnoms)ar ant-
protin squns that ar
ind on on, th orrspondnin th..lttrs'' (nulotids)givs un- multipliity. ah oganis
rnistakalvidnof ot-nlrronorigirr. all rflt th sam evoluti
olulr biology offrs two kinds <lfargumnts flr voltrti<ln, irsdon gne or protin does not si
two diffrnt grounds. Using th alphat arralogy,th first rgumnt sa}'s a st of spis, addition
that languagsthat us th sam alphat (th sam hrditry molul,th mattr has n sttld to
DNA md r"rpof th snrfour nultltids,and tlr sanr20 atlrino aids in oov, th widly d
thi protins),as wll s th sam litionary(tlrsm glltiod),arrrrot opn up th opportunity
of indpndntorigin. Th sond argumnt ()nrllssimilarity in th s- nt dgrs of rsolution il
qun of nulotids in th DNA (and thus th squnof amino aids i volving gnS to reonst
th protins);it says tht ooks with vry similar tts annot of indpn- fastr-volvin$ $ns to r
dnt lrigirr. divrgd oganisms.
Gns that nod th l
volving gnS.Thy hav
Informationalaromoluls ships among groups of or
among tria, rha,;
Nuli aids and prorins hav n a||ed i|'ormtiol romoltlls ing world), whih divrg
as thy ar long linar moluls mad up tlf squns of units- among th mirosopi pl
nulotidsin th as of nuli aids, amino ids in th s lf prltins_ ompard with plants and
tirat rtin oirsidr.rlmoLlnts of volutionary itrflrmirtion.omparisor-l of orgnisms that divg
of th sqn of th omponnts in two momoluls stalishsth tiond alir, whih volv
numrs that ar diffrnt.Bausvolution usually ous y hanging on gns' is usd to diphr l
unit at a til' th numr of diffrr-rs is an ir-rdiationof tlr rny of suh as among humans, fi
omon nsty. th fiinopptids involv
Th dgr of similarity in th squnof nlotidsor tlf amino ids ing th volution of los
an prisly quantifid. For ampl, in humans and lripanzsth maaqus' himpanzs,a
protin molul alld ytohrom_.whih srvs vitl funtion in rspi-
r a t i o n w i t h i n l l s ' o n s i s t so f t h s a m ] 0 4 m i n o i d s i n a t l vt h s a m
ordr. It liffrs, horvvr, frm th ytohro-t_of rlrsr.rsrnonk1.s y olular Phylogni
1 amino aid, from that of horss y 11 additinl amino aids, and from
that of tuna y 21 additional amino lrils. DNA and protins providt
Th dgrof similarit rfltsth nylf ommon nsty.Thus in- ags from ommon nCeS
frnsfrom omp.riltiventrr'nd othr lisiplinslnrtlir-rg vllu. nti hang that hs o
tionary history an tstd in molular studis of DNA nd protins by sm at first that quantifyi
xamining thir squnsof nulrltidsand amin aids. T authority of impossiI ausit w
this kind of test is ovrwlrlming:irl-rof th thor"rsands of gns irnd thorr- isms that livd in th prst
snds of protins ontind in an orgnism pvids an indpndrrttst of past ar somtimes prs
that organism's volutionary histor1'. largly disintgratd. Nev
Molular volutionrystudishav thr notal advntagsovompar- vid informatin about a
iltiv anatoly and th tltlrrlassial lisiplins.on is that th informtion As a spifi ampl'
is mor radily quntifial.Th nr-rrrrr of Lrnitstht ar diftrlltis rdil hrom-, whih is involv
olulr uolution 17
opid' dirtly or indirtly, from stalishdwhn th squnof units is known for a givn maromolul in
,, if ah omponnt nulotid of diffrntorganisms. Th sond advantag is that omparisons an mad
rpltsqunof nr-llotids in th vntwnvry diffrnt sorts of organisms. Thr is vry littl that om-
svralhundrd ooks, ah with parativanatomy, for xampl, an say whn organisms as divrs as yasts'
.Whn
rd lttrs on h pag. th pintrs,and human ings ar ompard, ut thr ar numrous DNA and
. . o o k s ' (' g n o m s )
'hs ar am. protin squnsthat n ompard in all thr. Th third advantag is
h ,.lttrs''(nulotids)givs un- multipliity.ah organism posssssthousands of gns and prtins, whih
all rfltth sam volutionary history. If th invstigation of on partiular
argumntsfor volution, asd on gnor protin dos not satisfatorily rsolv th volutionary rlationship of
t analogy, th first rgumnt says a st of spis, additional gns and protins an invstigatd until th
: (thsam hrditarymolui, th mattrhas n sttld to th satisfation of th invstigator.
ls,nd th sanr20 amino aids in orovr, th widly diffrnt rats of volution of diffrnt sts of gns
rary (thsamgntiod), annot opn up th opportunity of invstigating diffrnt gns for ahiving diffr-
ul-nntonrns similarity in th s- nt dgrsof rsolution in the tr of volution. volutionists rly on slowly
hus th squnof amino aids in volving gns to ronstrut rmot volutionary vnts but inrasingly
similar tts annot of indpn- fastr-volvinggns to ronstrut th volutionary history of mor rntly
divrgdorganisms.
Gns that nod th riosomal RNA moluls ar among th slowst-
volvinggns.Thy hav n usd to ronstrt th volutionary rlation-
ships among groups of organisms tht divrgd vry long ago: for xampl,
among atria, arhaa, and ukaryots (th thr major divisions of th liv.
||ed iformtiol mromoluls ingworld), whih divrgd mor tan2 illion yars ago (s Figur 1); or
mad up of sqnsof units- among th mirosopi protozoa (e.g., Plsmodim, wih auss malaria)
nino ids in th s of protins- ompard with plants and animals, th thr of whih ar ukaryoti groups
utilnary infornration. lmprisln of organisms that divrgd aout 1 illion yars ago. ytohrom-' mn.
Wo maromolulsstalishsth tiond arlir' whih volvs slowly, ut not as slowly as th riosomal RNA
tion usually Oursy hanging on gns,is usd to diphr th rlationships within larg groups of organisms,
; is an indiirtion of th rny of suh as among humans' fishs, and insts. Fst-volving moluls, suh as
th firinopptids involvd in lood lotting, ar appropriat for invstigat-
; o f n u l o t i d so r o f a m i n o a i d s ing th volution of losly rlatd animals, for xampl, th primats:
, in humans and himpanzs th maaqus'himpanzs, and humans.
l i h s r v s v i |f u n r i ni n s p i -
04 amino aids in atly th sam
tohrom-of rhsus monkys y olular Phylognisof organisms
additional mino aids, and from
i. DNA and protins provid information not only aout th ranhing of lin-
ny of ommon anstry.Thus in- agsfrom ommon anstors (ladognsis)ut also about th amount of g-
othr disiplinslnrningvolu- nti hang that has ourrd in any givn linag (anagnsis).It might
lr studisof DNA and protins y smat first that quantifying anagnsis for protins and nuli aids would
and amino aids. Th authority of impossil aus it would rquir omparison of moluls from organ-
f th thousandsof gns and thou- ismsthat livd in th past with thos from living organisms. Organisms of th
sm providsn indpndnttst of past ar somtims prsrvd as fossils, bt thir DNA and protins hav
larglydisintgratd. Nvrthlss, omparisons twn living spis pro-
:notal advantagsovr ompar- v i d i n f o m a t i o na o u t a n g n s i s .
:iplins.on is that th informatin As a spifi ampl, onsidr th prviously mntiond protin yto-
of units that ar diffrnt is radily hrom-,whih is involvd in ll rspiration. Th squnof amino aids in
I8 oleulr uolution
this protin is krrown for many rganisms, fom tria and ysts to insts volutionary trs aI
and hunrans;in animals' ytohrom_ollsistsof 104 amio aids. Whn th volutionary histoy of
amino aid squnsof humans and rhsus monkys ar ompard' thy ar as gnra' familis, or t
found to diffrent at position 58 (isoluin in humans' thronin in rhsus kinds of information r
monkys), ut idntial at th othr 103 positions. Whn humns ar om- nsis. Figur 5 illustrat
pared with horss, 12 amino aid diffrnSare found, and whn horss ar th rltiv rlationshi
ompard with rhsusmonkys, thrar 11 amino aid differns(Figu4). of th figur, humans al
vn without knowing anything ls aout th volutionary history of mam- to h othr than ith
mals, on would onlud that the linags of humans and rhsus monkys di- shorvs that th last o
vergd from ah othr muh mor rntly than they divrgd fom th hors mot past than the last
linag. orovr, it an onludd that th amino aid diffrn twn volutionary trs m;
humans and rhsus monkys mr-rsthav ourd in th human ling aftr its ah linag' or nagl
sparation fom th rhsrrsmonky linag (sFigur 5). last ommon nsto
h a n g d i n t h l i n a g t
t o r h s u s m o n k y s .T h i
,l that at position 58'
5 ]0 15 20 25 0 5 40 45 50
Figur 4) hav th snr
Uman tHGLtGRKTGQAPGYsYAAN
GDVKGKKItIMKSQHVKGGKHKTGPN
n t o n ( i s o l u i n )w
,h
Monke
aftr it sparatd fom 1
Horse
5 5 6 0 6 5 7 0 1 5 8 0 8 5 9 0 9 5 10 0 1 0 4
umn KNKGIlWGDLMYLENPKKYIP6TKMI
FVGIKKKERADLlAYLKKATN
olular Ronst
Monkey .....7
Hors Svral mthods ist f
opd for intrprtirrg
Figur 4. Amino aid squnof ytohrm-protinsin humans,rhsus
data; som an usd
monkys,and horses.Th squnonsistsof 104 amino aids,ah
rprsntdby a lttr.A dot indiatsth samamino id as in humns.S usd in molula vol
Fisur 3 for th namsof th min aids. maimum liklihood.
D|sAN M|
A ..distan'' is th num
Humn
diffrns ar msur
data) or to rtin maI
--t.,,.n",,, protins or th squn
monke
in Figur 5 was otain
amino aid diffrns
protin. Th amino aid
is rfltdin th num
plamnt of on am
Horse
lotid substitution in
ass it rquirs at last
im -__---.----.->
numbr of nulotid di
Figur 5. Th volutionf ytolrrom-among hurnans,monkys,and horss. ssary to aount fo th(
Th numbsinditth mino :ridrplamnts that hav tkn pl:rin h lutionary tr basd on
ranh of th phylogny. nulotid hngs in z
olculr uolution 19
from atriaand yasts to insts volutionary trs ar modls or hypothss that sk to ronstrut th
Lsistsof 104 amino aids' Whn th volutionaryhistory of taxa (i..,spisor othr groups of organisms, suh
us monkys ar ompard, thy ar as gnra'familis, o ordrs).As pointd out :lrlir' th trs ma two
in in humans,thoninin rhsus kindsof information rlatd to volutionary lrang'ladognsisand ang-
.Wlrn
positions. humns ar om. nsis.Figur .5illustratsboth. Th rarrhigltionshipsof th trs flt
]s ar found, and whn horss ar th rlativrlationshipsof anstry, or ldognsis.Thus in th right sid
1 amino aid diffrns(F-igur4). of th figur,hunrns and rhsus monkys 21sn to mor losly latd
t th volutionary history of mam_ to ah othr than ithr one is to th hors. Statd anothr way' this tr
of humans and rhsus monkys di- showsthat th last ommon anstor to ll thr spislivd in a mor r-
y than thy divrgd from th hors motpast than th last ommon anstor to humans and monkys.
t th amino aid diffrn twn volutionarytrs may also indit th hangs that hav ourrd along
;urrd iir th human linas ftr its ahlinag,or anagnsis.Thus in th volution of ytohrom- si th
l (sFigur.5). lst omnron zlnstor of humans and rlrsus monkys, lr-rarino aid
hangdin th linagthat Wnt to httmns ut none in th ling tha Wnt
to rhsusmonkys' This lnlusionis drwn from th osrvation (Figur 4)
that at positin 58, monkys and horss (as wll s othr animals; s
25 0 5 40 45 50
Figur4) hav th sam amino aid (thronin)'whil humans hav a diffr-
LHGLtGRKGQAPGYsYTAAN
KHKGPN
nton (isoluin),whih thrformust hirvhangd in th human linag
t'.D.. aftrit sparatd frolrl th monky linag.
i 80 85 90 95 100 104
)6TKMItVGlKKKERADLIAYLKKATN olularRonstrutionof vlutionary History
DlsAN oDs
Organisrn r0 1t 12 1-l \4 l5 T6 17 1u t9 20
1.umln - r t3 17 16 1-l 12 t2 17 t6 18 18 19 20 ) i
.) t -)-)
1/
-)o 6 56 66
2. onk1, _12 16 l-5 t2 11 1 16 1.S 17 t7 18 ZT 2 2 5 62 57 6_5
. Dog 10 8 4 6712 12 14 t4 l-l 30 29 24 28 64 61 66
4. Flors -1 5 11 11 t6 16 16 \7 \6 -'t-z- 27 24 3 64 60 68
.5.Donky 4 10 t2 15 15 15 16 15 .tl 26 25 2 64 59 67
6. Pig 6 7 ] 1 t |4 1 30 25 26 31 64 59 /
- .10
7. Rabit 7 8 1r 11 11 25 26 2 29 62 59 t)/
8. Kangarlo -14 t4 15 13 14 30 27 26 31 66 -t8 68
9. Duk -)
11
.) -)/ 2"4 26 2s 29 6\ 62 66
l0. Pigon -4 48 24 27 2.6 30 .59 62 66
11 . h i k n 28 28 26 26 31 61 62 66
12.Pnguin -6 28 27 28 30 62 61 65
1. Turtl 0 27 30 33 65 64 67
14.Rattlsnk 8 40 4| 6t 61 69
1 5 . u n a _44I 72 66 69
16. Srwwormfly _16 .'8 6 65
1 7 .M t h s9 60 6\
|8. Nurlspor <'7
61
19. Sbroms |
20. ndid
B 6 ia ] a ! J-i
i-lo-;1:!QJ v=.-
_151E ] *.: -
- ;
,9
i e. >-;+P+
' a
d * (D :'
0.o.
iq.=.:1 ;J - 5
i' a
^w'^*:v
a.-; = a: o = L:.
r }
rl^'p:-
}3li91;fii
} g i * 1 a i i
. ^ F - I
;;*5n
i <
3 g 1; a .' }!
*
i., - . )
rz a =
- '' l '
,a
-
rN.t
oleculruolutio 141
}* un
i - m - * ! a Sroworm
....ot-+ ||
!N.Od-..)l l rA"^
o.-tc '
9.9
''lN.j |
tl'
o
l
.t;
.6: s :
th powr of th mthod is apparent in that a singl protin yilds a fair| a-
= i
= O > -
t*
<.* uratreonstrutionof th volutionary history of 20 organisms that startd
l . . _ - / * > \
: : = : l R
]ti.,r>zd ,.
to divrgmor than 1 illion yars ago.
Th most ommon produr to transform a distan matrix into a
-j^i-; =t.] .od.
_***it+!
phylognyis alld lustr analysis. Th distan matrix is sannd for th
1 4 2 oleulr uolutirl
smallst distan lmnt,and tlr two taa involvd (say, human and mon- from th linags that .
ky in Tal 1) ar joinlat an intrnal nod, or ranhing point. Th matri vors' fr exampl' dol
is sannd gain fr th nt-smllstdistirn,nd th two nlv ta (sy' gthr aus they pol
hors zrnd donky) ar lustrd. Th produr is ontinud until ll taxa ommon anstor bran(
.!fhn
hav n join1. a distan involvs a taxon tht is alrady }rertof a and oyots.
pviluslustr, th avagdist:inis otiirltwnth n'ton irnl
th prxisting lustr (say, th avrag listantwn humurnand hors
MA|U-LlKL
and monky anl hors).This simpl prtldurssllmstht th rat of vo.
lution is uniforrn alorrg :rll ranl.rs. aximum-liklihood mt
Othr distan mthods (inluding th on usd ttl onstrut tlr tr in availal data. Thy r<
Figur 6) rlax th ondition of r"rniformrt and illlow for rrnqulrats of would mak it possiblt.
volr.rtionallng th brirnhs.or-rof th most tnsivl1,usd ll-rthodslf hang. For ampl, as
this kirrd is alld nighor-joining.Th thod starts' as for,y idntify- mor likly than transv
ing th smallst distan in th mtrix and linking th two ta involvd. Th proaility must assl
nt stp is to mov ths two ta nd alul:lta nw mtri- in whih p o i l i t i sf o r a h i n
thir distans to othr taa ar rplad y th distan twn th nod tr is th on with th I
that links th tw.o taxa and all otlrr taa' Th smallst distn in this nw all possibl trs'
matrix is usd for making th nt onntion' rvhih will twn trvl aximum-liklihood
othr taa or btwn th prvious nod and a nW taon. Th produr is numr of taa is larg
ptduntil all tax hav n onnt]with Oll an()thy intvning whih th proability m
nods. of taa. With 10 taa,
taxa, t numr of pos
A x l M U r . 4 . P A R s
l oNY MoDs vn with powrful o
h i i t i v i f t h n u m ro t
aimum-parsimony mthods sk to ronstrutth tr that rqrrirsth t i m r q u i r d .H u i s t i
fwst(i..,most parsimorrious)numr of hngsstlnrmdltlongll rrhs. sil ts is xmind,
This is rasorralassumption ausit usually will tlr rnost Iikly. But
volution may not nssarilyhav ourrd follorving a minimum path, -
sA|slAL AP|
us th sarn lrang irrstadnray hav <lurrdirrdpndntlr'alorrgdif-
frnt branhs,and som hangs mav hav involvd intrmdiatstps. Th statistial dgr o1
Not all volutionary lrangs,vn thos that invlv a sirrglstp, may and maximum-likliho
quilily proa bl. For ampit lnong tlr t<lurrrr.llltil ssirr DNA, y- strapping. It onsists of
tosin () and thymin (T) ar mmrs of a family of rlatd moluls data point at random a
alld pyrimidirrs;likwis, adr-rin (A) nd guanin (G) llngto a family random sampling pro
of rrrolulsalld pr-rrins.A hang withir-ra DNA sqlln om on ootstrap valu for al
pyrimidin to nothr ( *T) or from on purin to anothr (A *' G)' alld all spis drivd from
transition, is mor liklv to our than a hang fom a purin to pyimi. bootstrap valus aov
d i n o r t h l n v r s ( G o r A . ' o r T ) , a l i la t r n s v s i l nP. r s i m o n r ' thos with ootstrap val
mthods tak into ount diffrnt proailitis of ourrn if thy ar
known.
imum-parsimony mthods ar rlatd to ldistis' a vry formalisti lVlolular Phylog
thory of tonomi lassifiationthat is tnsivlyusd with morphologi.
al nd palontologial dl.rta.Th ritil fatur i ladistis is th idntifi. Th mthds for otai
ation of drivd shard traits,lld syn:1pomorphitraits. A synapomorphi mously improvd in r.
trait is shard y som taa ut nlt othrs l.lsth formr inhritd it gns hav bn squn
from ommon nstor thar quird tlr trait irftr its lirragsperatd has n squendin r
oleulruoltion 14
ids,nodd 1'2nulotids.
iormous volutionary information,
rimation, shown in Figur 6, tl of ytohrome- moluls annot dtrmin volutionary rlationships b-
spisovr a priod longr than twn losly rlatd spis. Fr ampl, th amino aid squn of
volving protin. \/idly diffrnt ytohrom- in humans and himpanzs is idntial, although thy di-
rtion of th amino aids in thir vrgd6_8 million yars ago; twn humans and rhsus monkys, whih
tudy of gntidiffrns twn divgdfrom thir ommon anstor 5_40 million yars ago' it diffrs y
s a m a s o n h. o w v r , o m p a r i s o n only on amino aid rplamnt.
146 oleculr uolutio
Protins that volv mor rapidly than ytohrom- an str'rdidin or. .W.ithin
r|ir. ah
dr to stalish phylognti rlationships twn losly rlatd spis. v o l v a t d i f n tr t
Som protins volv vry fast; th firinopptids-small protins that ar vnts. Sintists n
involvd in th lood-lotting pross-ar suital for ronstruting th th volutionary phy
phylogny of rntly volvd spis' sh as losly rlatd mammals. n u m r o f g n si n v
othr protins volv at intrmdiat rats; th hemogloins, for xampl' tion twn gns o
an usd to ronstrut volutionary history ovr a fairly road rang of g n s o r h i v i n gd l
tim (sFigur 8). volving gns an
On grat advantag of molulr volution is its multipliity, as notd t h a r n o d s l t l w l v .
l u t i o n r yr l a r i o n s h i
rdundant odon su
th nodd mino i
PE
:' ;e ! l sprsd among th s
U }a.
;::>
sgmnts of th gns
G o d i n g p r t i r l n so f g
.9-aE
g . l o t i d st h e t s p i f vr
> nd
.!
s .J s
tt ttt t
vv v v rr}' Th }yIolulrl
0
160 O n l l n s p i u o t ' ts t t r i
h o m < l I o g o um s olul
140 pl. popotionsf nL
I v o l u t i o n a r y h a n g
1?n
I
t o D N A o r p r o i n st h
00
tion. Studis of mll
100 "r"*
maromrlulsmay s
.s,9,
80 It ws first osrv
ns twnhomolt
60 n a r l y p r o p l r t i o n a lt t
r o r . I f r h a t o f rt
smin th voIui
D N A s q u n sW o u
q u n s o u l d r h n h
ing vnts of a phyl
ourrd.
onsidr, for am
gn oding for yto}
o u l d d t r m i n t h t
0O simply y amining
0
O9UUO
o 0
r n h .o n w o u I dn
0 10 2oo 30o 40o 5oo 600 7ao 600 9oo 10oo
E<^ sour' suh as th tt
v
t illins of yrssin divrgne
t i m r h a t l p s di n r
Figur 8. Rats of molular volution of thr diffrnt protins. (Aftr T h m o l u l r v o
Dikrson 1971.\ lok, lik a wath
ollr uolutio 147
ytohrom-an studid in or- alir.Within ah organism af tousands of gns and protins; ths
ps btwnlosly rlatd spis. volvat diffrnt rts, ut vry on of thrrrrfltsth snrvolutionary
ropptids_small protins that ar \.nts.Sintistsan otain gratr and gratr auray in ronstruting
-r suital for ronstruting th th volutionary phylogny of any group of organisms y inrasing th
suh as losly rltd mammals. numrof gnsinvstigatd.Th rang of diffrnsin th rats of volu-
tts;th hmogloins,for ampl, tion twn]I.lsopns up th opportunity of invstigatingdiffrr-rt sts of
histoy ovr a fairly road rang o{ gnsfo ahivil-lgdiffrnt dgsof rsllutiorrin th tr of lit; slowly
volving]nsn b usd to study rmot volutionary vllts. vn gns
olution is its multipliity, as notd thatndslorvly vlving protins an usful for ronstrutingth vo-
lutionaryrlationshipstwnlosly rlatd spisy amintion of th
dundantodon sustitutions (nr.rlotid sustitr'rtions
that do not hang
th nodd mino aids), th intrns (nonoding DNA sgmnts intr-
sprsdamong th sgmrrtstlrirtod fo mino ids), or othr nonoding
a
fE onsidr, for ampl, Figur 6. If th sustitution of nulotids in th
gnoding for ytohrom. ourrd at a onstant rt through tim, n
.' ould dtrmin th tim that lapsd along any ranh of th phylogny
simply y arirring th nrrmr lf nulltid sustitutions alorrg that
ranh.on w<luldnd onl,vto alirat th lok by rfrntl an outsid
50 600 700 80 9oo 10oo
sour'suh s th fossil rord, that would provid th atul gologial
ears sine divrgne
tim that lapsd in at last on spifi linag.
hr diffrnt protins. (Aftr Th molul:r volutionay Iok is not xptd to b nltronomi
lok, lik a wth Or othr titr-rpi that msurs tim tly' ut
148 oleculr uolutio
n a stohastilok th proaility
rt (for ampl, a givn quantity o{
gh day' to rdud y half in
:urs in th atual amount of hang.
hasti lok is quit aurat. Th
lutionary lok lis in th fat that
. ah lok ..tiks'' at a difjrnt
cf a partiular gn or protin-ut s
gnsor protins provids an ind_
y vents.
rnt of vriation osrvd in th vo- z Z3
:han is ptd from a stohasti
rdisprsd''' or somwhat rrati.
ong diffrnt linags ar not s-
in prinipl, to tim phylognti o 2s 50 75 100
-125
d s i r d
, u r m o r g n so r p r o t i n s illions f ersof volutin
xamind than would requird
Th avragrats otaind for sv- Figur 9. Rat lf nuiotid sustitution ovr palontologial tim. ah lf th
rly pris lok' partiularly whn ]5 dots marks th tim t whih a pair f spis divrgd from Ommon
anstor (horizotl scle) and th numr f nulotid sustitutions, or
) n a r y v n t si n v t r l v l o n g t i m p r i -
protin hangs' that hav ourrd sin th divrgn (uertil sl).Th
ngr).
solid lin drwn from th origin to th ()utrmost dot givs th avrag rat of
Lr 9, whih plots th umulativ sustitution.(Aftr Fith 1976.)
rrotins against th dats of divr_
ings) as dtrmindfrom th fossil
rstitutionis fairly uniform. Som
s low th lin at th lowr lft of
owr rat than th avrag for th volutionis slowr, th squnsusd ar shortr, and th tim oms in-
austh mor rnt th diver- rasingiyrmot.
it is that th hangsosrvd will
.A s t h l n g t ho f t i m i n r a s s ,
ny linagar likly to anl on Conlusion
, that molular tim stimats tnd olular iology mrgdas a disiplin 100 yars ftr Darwin' following
i asd on othr mthods and, in- th 1953 disovry of th doul-hli strutur of DNA' th hrditary
ris is a onsqunof th statistial hmial.Molular iology provids th strongst vidn of iologial
ar asymmtrially distriutd. B- volutionand maks it possil to ronstrut volutionary history with as
ar diffrnsbtwn two spis muh dtail and prision as anyon might want.
But ovrstimation rrors ar un. Molular iology provs volution in two ways: first, y showing th
; ar boundd' sin thy annot unity of lif in th natur of DNA and th workings of organisms at th lvl
h of a typial distriution of sti- of nzymsand othr protin moluls;sond,and most importnt for vo-
d, gathrdfrom a numr of dif- lutionists' y making it possil to ronstrut volutionary rlationships
ll shap,with a larg numr of that wr prviously unknown, and to onfirm, rfin, and tim all volu-
hr at on nd and a long tail of tionary rlationships from th univrsal ommon anstor up to all living
th othr nd. Th avrag of th organisms.
)vrstimatth tru dat (Figur As pintd out arlir in this haptr, all organisms ar mad up of th
ratr whn th rat of molular samasi omponnts,whih, morovr' are similarly assmldand usd.
150 oleulr uoltiln
-14
I
o B|LIoGRAPHY
!12
Avis,J. .2004. olu
10
o A B
Sundrland,A: Sin
A y a l ,F . J . , d .1 9 7 6 . o l
*B B a k r ,A . J . ' d .2 0 0 0 . o
o
Lr- arroll, S. B. 206.h
Rlrd of ullutil
h
D i k r s o nR , . . 1 9 7 1 .T h
volution.lourl of 1
Flsnstin, l. 204.lfer
Fnhl,. 2002. Th ri1
UnivrsityPrss.
Fith,.W.x4. 1976.Mol
15 5 55 75 95 1.l5 .15 .155 175 195 215 uolutio,160-.l78. l
Fith, \V. N4.,and . arg
Divergenetime(x100MY) I55 279-284.
Grur, D., nd \7.-. Li. 2
Figur 10. Skwingand ag ias in stimatingmolulardats.h instshows Sundrland,MA: Sin
th tr topology for linagsA, B, and ; t.. and t rprsnt'rsptivly' Hll' B. G.2004. Phlog
alirationand targttims.Th min panl shows a fqunydistriutionof Assiats.
1,000 stimatsof th divrgntim twnlinags and AB in th inst' Hazn, R. . 2005.ge.e
st to hav ourrd3 billion yars ago and taindwith th us of a short (7.5 D: Josphnry Prt
rsidus),slow.vlving(onrplmntpr sit pr 1010yars)protin and Lynh.M. 2007. h orig
with th us of th split twnA and B, st to 300 million yarsago' as a Assoiats.
aliratilnpoint. T and M rprsntatual (3 illion yars)and stimtdman Ni, \4.,and S.Kumar.2
(4'084 illion yars)tims.(odifid from Rdriguz-Trlls t a|.2002.| Oxford UnivrsityPr
Rdriguz_Trlls' F., R. T
toward vrstimatio
Ntir.lldmof
.Wos,
C. R. 1998.Th ur
SisUS 95: 68.5
In all organisms' from atria to humans, th instrutionsthat guid th d-
vlopmnt and funtioning of organisms ar nasd in th sam hrditary
matrial, DNA, whih provids th instrutions for th synthsis of protins.
Th thousands of divrs protins that xist in organisms onsist of th sam
20 amino aids in all organisms, from atria to plants and to animals. Th
gnti od, y whih th information ontaind in th DNA of th ll nu-
lus is passd on to protins, is shard y all sorts of organisms. All organ-
isms us similar mtaoli pathways-squns of iohmial rations-to
produ nrgy and to mak up th ll omponnts.Th unity of lif rvals
th gnti ontinuity and ommon anstry of all organisms.
Th dgr of similarity in th squnof nulotids in th DNA (and
thus th squnof amino aids in th nodd protins) maks it possil
to ronstrtvolutionary history. ah of th thousands of gnsand thou-
sands of protins prsnt in n organism provids an indpndnttst of th
organism's evolutionary history. Th volutinary lationships among all
oIculr uolutio i ),
t BILIoGRAPHY
(A, T' G, and ), whil that of th doul strfur is spifid y th fr. vlus gnoms ar usuz
qunis of the as pairs A and G. RNA is gnrally single strandd, with flunza) pakag svr
a diffrnt sugar rsidu from that in DNA and with urail ass instad of into tlr sam vius pal
thymins' have diffrnt RNAs pa
Although th origins of th gnti od ar unknown' slf-rpliatirrg thus omin diffrnt
RNA moluls may hav n th arlist form of ..lif,'' ut th transition ing multipl inftions,
to gnoms that usd th muh mor stabl DNA ourrd long ago (Joy F{urns and otlrr
1989). Dol-stranddDNA also allows damag to ortdy opying isms long to th uki
from th undamagd omplmntry strand. Sin DNA largly laks RNAs DNA organizd into l
ability to at as a atalyst, this hang must hav gon along with th volu- lls (Figur I). h
tion of a mans of transriing on stand of a gn's DNA into RNA. In dal with th prolm
prsnt-dayorganisms th squnof bass in th ..mssngr''RNA tan- uls, and tromrs
srid from a gn dtrminsth squnof th amino aids in th protin sion to th daugtr
hain nodd y th gn (Figur 1). Thr ar also many RNA molulsof ukaryots, suh as th
dirt funtional signifian, whih r transrid (opid) frm DNA ut gno1,with just on
not translatd into an mino-aid squn. mat' th is rif d
ah mating partrrr.Tl
oRGANlzA|oN oF H GN| MAR|AL
Viruss
Plnt viroids
hromosome Genes messenger spIid Protein Influlzavirus
RNA MRNA Rovirus
Phag tpX174
telomer
.
intron
+l trans|tion
-+
SV4O
Vaitriavirus
Batria
intron
f" . oli
'i',, i o1llsm gitlium
I Nulurrgnomsf trk
enromere Y east (Scchro|l,t))L:s
i ntron
Nmatod (orhbd
Fruit fly (Drosophil m
F Ftsh (Fugu rubrips)
FI
+# Slmandr
t
Mat-lrmal (Homo s|li
telomere ress (Arbido1l sis thlit
Miz (Ze ms)
Fritill11'p ritillri,lss
Ntlr: Arvitions:p
Figur 1' Chromosoms,showingthir major ftursand th andingpttrns kiloss;, mgass
that an snwith stiningmthods.Th figurlso shows a shmatiof a ? mns tht th gnnu
rgion of th hromosom ontaining thr gns'th mssngrRNA produd ? ? t h . r ri t i s b s do l i n .
y thm, and th individuaIprotinsnoddby th mssngrRNA. lr <lrgnisrs.
uolutilof tb Gome ].i.i
l stutur is spifid y th fr- virus gnoms a trsually singl nroluls,som RNA viruss (sulr as in-
,IA is gnrallysirrglstrandd,with
flunza)pakag svrl diffrrrtRNA rrrolulsthat r, diffIltgns
'trAand with urail ass instad of
into th sam virus partil. Otlrr RNA viruss (.g.,alfalfa rr-rosaivirus)
havdiffrntRNAs pakagd into sparatvirus partils.Ths virussan
]od ar unknown, slf-rpliating thus omin diffrnt portions of thir gnom from diffrnt parnts dur-
:st form of ..lif,'' ut th transition ing multipl inftions,a primitiv typ of sxualitY.
l DNA ourrd long ago (Joy Humans and othr nimals, plants, fungi, and man,v singl_lldor!]an-
i damagto orrtdy oying isms long to h ukaryot division of lif' whih is dfind v hving th
nd. Sin DNA largely laks RNA's DNA organizd into linar hromosoms arrid within th rrLrliinsid
Jst hav gon along with th volu- lls (F.igur1). Th hromos<rtnshav spializd rgions (tlomeres)to
nd of a gn's DNA into RNA. In dalwith th prolm of rpliating th nds of dul-stranddDNA mol-
lss in fh ..mssngr''RNA tran- uls,and entromresthat ar quird for ort hromosom ti1nsmis-
rof th amino aids in th protin sion to tlr daughtr lls dr"rringl1divisiorr (Lwin 2008). N,{arrypinririv
r ar also many RNA moluls of karyots'sLthas th grn a|ga hlmdons, hav a prirllarily hploid
ransribd(opied)from DNA ut gnom'with ;ust on opy of h of th diffrrrthrmosorns.Whn lls
. mat' thr is a rif diploid stag, With a hrmosom st ontriutd y
ahmating partnr. This undrgosa spial forrrof ll divisirr (miosis)
l ARIAL
Viruss
Plant viroils ssRNA 0 00p
spIied Protein Influnzavirus ssRNA 12 l 3 . . 5k
MRNA Rovirus dsRNA 22 2kb
Phag<p174 ssDNA 11 5 . 4k
translation sv40 dsDNA 6 ) l]
+# Salamandr
Mammal (Homl spins)
r ess(rb i dopsi s th li)
3 0 , 0 0 0( ? ? )
2 0 , 0 0 0( ? )
27,000
90,000
3,000
1 0 0 b
aiz (Z lttvs) .r0,000(??) .i,000
Fritillary (F ritil lri s sric) 2 7 , 0 0 0( ? ? l L20,000 M
Nofe: Arviations:bp, baspirs;ss, singlestrndd;ds, doul strndd;kb,
lr fatursand th anding pattrns kilbass; ' mgass.
figuralso shows a shmtiof a ? mnstht th gn numbr is lnll' pproximatlyknlvn fror th gnolrrsqlln'
gns'the mssngr RNA produd ?? that it is asdorinompletvidrror anallgrvit squnl glltlt-t-ts
<lfsimi-
d y th mssr-rgrRNA. lr rsanisrns.
1.'6 uolutir of tb Gom
f
\
oI rh rprldutivpross'
Chromosom numrs in th haploid st vary widly among diffrnt
spis:16 in th as of hlmdomos,5 in Drosophil,2 in humans,
but somtims just a singl hromosom (in a spisof ant and an stris
roundworm). harrgs twn spis in homosomal stutur ar om-
mon, involvtng dtlplition.snd dItit.lltsof gneti matrial, inursiosof
parts of hromosoms (Figur 3), and trsloctils(hangsof matrial
twn diffrnt hromosoms). hromosom rarrangmnts our infr-
quntly in volution (roughly on pr gnom pr 10 million yars in mammals,
although som goups, suh s rodnts, hav rts 10 tims highr: ihlr
and Sr-rkoff2003). Ovr volutionary tim) tlr rlrdr of gnswithin a hro-
mosom gradually oms srambld y sussiv invsions. Svrl
human hromosotns sm to hav rmaind unhand in grr ontnt
PrentaI
hromosome
(though not ordr) ov
m a d u p < l fs p a r a r l
oD|NG AND N
or rpeatedgns
U n e q u a| ro s s ingo v r bet wendup|iated Gnom siz is dfin
loid gnom. A gnom
r t r n s r i di n r o m t
quns (Lwin 2008
qes ar'd itros.h
I
transrid but ar rr
Y
latd; in ukryots,ir
as th spli<lsot. |
that ar rsponsil fo
and timing of mssng
Figur 2. op: rossing ovr in a rgion of a homosom ontaining fiv gns,
oding u ftlnrion
A to , shlwing th rsulting rominant ombinations of th gns (indiatd
vn with omplt
y uppr. and lowras lttrs nd lso y lk rrd gry lins for tlr twl
diffrnr parlrtal romosonrs). Bottom: rlssing ovr in a gion rrf
g r l s i r s o d i n g
hromosom ontaining two dupliatd gns that r similar nor.rghthat thy q u n s .T h t a s k i s
may mispair, lading o unqual rossing ovr. portion of oding sq
uolutio of tbe Genom 157
I
s of portions of thir hromosms
r e 2 ) . I n a n i m a l sa n d h i g h r p l a n t s
aploid statis oftn just a rif part
invertd rgion
l-
I
l st vary widly among diffrnt
s, .5in Drosopbil,23 in humans,
(in a spisof ant and an sris
!
n hromosomal strutur ar om-
ls of gnti matril, iursils of
I
lslotios(hangsof matria]
osom farrangmnts our infr-
n pr 10 miiliorr as in mamrnals,
lr non-invertdregion
I
hav ts 10 tirs highr: ihlr
l' th ordr of gnswithin a hro-
y sussivinvrsions. Svral -
raind unhangd in ]enontnt l.
ffi
!
,l
Figur 3. A hrtlnlosom invrsion. Th diagram shorvs th nding pattrn of a
hromosom, and t apparan whn th top prt of th hromosom has
\ lrarrangd so that part Of th gtror in th ight-hand hromosom is th
othr Way around from th orlr in th lft ln.
\
ff,H,i;:il' ParentI
hromosome
(thoughnot ordr) ov th ours of som 70 million yars, ut othrs ar
_/
t:i I mad up of sparatloks drivd from diffrnt atrstralhromosoms.
cphiI gnons (blu.grnalgzr)and proaly volvd around 800 million yrs ago, whil
.-rhronrati!]n()ls mitohondria volvd muh arlir from a group of atria known as -
protoatria(Gay 1'999).
Numbr
,%
Ltr ndosymiont vnts hav also ourrd. Th apiomplxan roto-
of fmilis of te gnom
zoa (malaria parasitsand thir rltivs)and dinflgllatsontain plastids
drivdfrom rd alga;prsumly a protozon ll ngulfd a rd algal ll.
r00 8 mrs of th hromophyt group of alga also ontain a rd-algal-drivd
13 hloroplast, whih has rntly n shown to rprsl.lta whol rd algal
z0 ll.As wll as thir plastid gnom, mmrs of th ryptomond sugroup
60 .)
still hav a vstigial nular gnom drivd from th rd algal anstor (with
ovr 400 gns)arrd thus possssfour gnoms. inluding th mitohondri
49 2.65 and th tru nular gnom (Douglas t l. 2001).
27 0.97
20 0..'
GENE DUPL|AIoN AND GN Loss
umn (llt.t-t
S1uI-ring
nslrti um
(2002).
or frquntly, inrassin gn numrs om fronr gn dupliations.
l l R p t I m n t s ( l r l r t sw i t h r t c d
Nw gns aris from dupliations of parts of th gnom or of whol
. w h i h | i l r t h r r ' u h ( v r \ ( .t r ; l I t s r i -
gnoms(ihlr and Sankoff 2003; Lynh 2007). Singl gn dupliations
s q u n ) ;S I N s ' S h t l r t I t r s p r s l
: h r p l i t t h r l t r g h n R N A i n t r r r l d i t an oftn rognizd aus thy fquntly our as tndm duplic-
th nz1'm ndd for thtr vrs tan- tios |ose to thir prognitor gn (Baumgartn t al. 2003). Th sizs of
l m n t s ( s i r n i I rt L T R l r - n n t sl,l u t suh arrays n thn hang furthr y unqual rossing ovr. An hang
) N A l m n t s ( | l n t st l r a t p l i t twntwo diffrnt rnmrs of an array rts on dsndant with thr
vith short invrtd rpats t ah nd). opisf th squnand on with a singl opy (sFigur 2).
Som dupliatd arrays of gns probly volvd undr sltion that fa-
vord larg amounts of th gn prodttt' for ampl' th larg arrays of
A or (to a lssrtnt)in introns. In gnsthat nod th RNA omponnts of th protin translation mha-
vr,vdnslvpkd, with rnultipl nism (riosoms) and th xpandd aays of sonr instiid-rsistn
rh portion of th D. mlogster gns (Ranson t al' 2002). Oftn, howver, highr grr dosag has no ad-
lmatin is iarglv md up of matrial vantag'so that natural sltionwill not at to prsrvth funtions of du-
1 9 8 8 ) ;t h r g i o n s a r o u n d t h n - pliatdgns.Givn nough volutionary tim, an inativ mutant opy of
\X/rightt al. 2003) and many intr- on of th dupliat gns may om fixd in th spis y gnti drift.
1 9 9 6 ) a r s i m i l . on a dupliatd gn has lost its funtion, fLrrthrmutations may ou ln
it and fixd in th population y drift. vntully th gn may los its
protin-odingaility y mutations rating odons' and it thn oms a
rmnant psedlge;vntually deltions f gnti nltrial my rnrov all
rll, but today's orgnisms hav larg or part of it. Nwly dupliatd gns ar thrfor oftn ptd to
s. An irrrportant'tough infrqttnt, quikly lost fronr gnoms (Walsh 1995; Lynh 207). Loss of gn dupli-
rom oth orgnisms.Batriirrgu- atSmans that' aftr a tim' whol gnom dupliations ar diffiult to dis-
: t r i a sl p i s( o h m n t a l . 2 0 0 0 ) . tinguish from partial dupliations; in ithr sitution only a fration of th
i-rirown gnols'as w]l as th nu- gnomis rognizly dupliatd (IJ7onget a|.2002). \/ithornplt gnom
lgahv hloroplasts,and d lga squndata that provid larg gnom strthsof larly similar gn on.
lls oth ontin mitohlndri (Lvnh tnt,dupliat rgions lr rognizd,vn whn most of th gnswithin
of prokaryortlrigin :rnd aros arly th strths hv returnd to th singl-opy stat. Chromosom arrang-
tril ll was ngulfd into a ukary- mntswill, howvr, vntuallyras all tras of dupliation.
n r-rdosymiont. sqtll1sof gns Somtims' howvr' dupliatd gns may gain nrv funtions y mr-rtation
tosof photosynthtiyanoatria for inativating mutations our. For instn, an nzym that atalyzes a
162 uoltillt tlf th (}llll
for thir own pliation and prtltin synthsisand flr a fw spializdor- th mutation rat pr nul
ganllar funtions. any gns ssntial for organll fr-rntilns hav n atria. For ampl, in }
trnsfrrd to th nulus (Gray 1999). ongoing gn trnsfr hs n d- drashov 2003). Givn th n
ttd frorn plant organllgnoms,s wll as transfrsthi1tratnonfun- man gnom, this translat
tional psudognsof orgnllgnsin th nular gnomsof many spis protin squnof aout
( G r a y 1 9 9 9 ;B n s a s s o n t a l ' 2 0 0 1 ) . tion, and around 80% of tl
Gnome-siz volution involvs svrl prssurs.Thr may advan- nt mutation rat to dlt
tags lf smallr gnom siz, ausir rdud gnom n rpliat fastr, largr numr of dltrro
allowing fastr ll division; this arr oLlr through dltions of gns that th nonoding portions of 1
ar no longr ndd, or of intrgni and introni squns.Th finding th stimat of th human
that ompat gnoms oftn hav ovrlapping gns, in whih th sam H i g h d l t r i o u sm u t t i
strth of DNA is transridin two diffrnt rading frms (Douglas t al. (Crow 199) ma limit p.
2001; l,win 2008), suggststhat sltion fvord siz rdutiorr in ths Thortial modls of th
ass.But loss of som squnsmay sltivlynutral, so that dltions at that mutatinal load
an fixd y gnti drift, as outlind arlir for rdundant gn dupli- 2 0 0 0 ) . H o w v r , t h r i s l
ats. In nonrptitiv squnsthat ar unonstraind y sltion, th mhanisms, whih slow
amount of DNA tnds to dras ovr tim aus rpliation rnistaks sours. Th ourrno
sm to aus small dltions rnor oftn thn insrtions of ases (Grgory rats to zro, ut in sxua
2004; Lynh 2007).If sltivonstraints on gns r rmovd, thy v'.ill twn gns' alllsthat
thn gradually dlin in siz ovr tim s mor dltionsthan insrtions - atd with favoral mtat
om fid y gnti drift. Prtlaly oth sltiv and nonsltiv pr- l i t t l d v a n t a g . u t t i o r
s s sh v no p r a t i n g . ar thus likly to rflt tl
rduing th mutation rat
load (Snigowski t al. 20t
MUA|oN RAES AND H NUMBRS oF GNs
I n o n t r a s t ,i n r g n i s m
ost mutations with ffts on an organism's funtions rdu survival or fftiv gnti rom
frtility (fitnss,for short). Unlss sltilnis vry wak, thr will a al- s P i s . n a l l l t h t i n
an twnth mutational input of nw dltriousvariants into a popul- soiatd with any favora
tion and thir liminatiln y sltion. Th prsn of ths muttions qny as th mutation s1
rdus avragfitnss,lmpardwith a mutation-frpopulatin; this mu- m u t i o n a t st h a n i n s }
ttiol lod depedson th totl numr lf nw dltriousmutations that ig ad is obsrvd in al
aris pr lndividual ah gnration,suh that th n!]ativof th natural log- sltion prssurs(Snig
rithm of th man fitnssof a poprrlatilnis qual to th lad (row 199|.
For a givn mutation rat pr gn, thr will thus a gratr load if th GNoM s|z |N
numr of gns in a gnom is larg. RNA gnoms lak rror-orrting oF NoNoD|NG t
mehnisms and hav trmly high mutatilnrts pr gnom pr rplia-
tion, dspit thir small gnom siz: around 1 nw mutation pr rpliatiln Gnom-siz diffrns
yl for th influnza virus (Drak t al. 1998). Suh gnomsannot' thr. v o l v l a r g d i f f r n si r
for, hav larg numrs of gns, austhir {itnsswoLrld tlm vry DNA, with only minor
low. DNA-asd gnoms hav rror-orrtingmhanisms that giv thm Lynh 2007). In som a
muh lowr, ut not zro' muttion rats; the. oli !]nomis stimtdto in lif history and olo
hav a mutation rat of around 10_r0pr nulotidpr rpliation and a mu- hav mor satllit and ot
tation rt pr gnom of only 0.0025 (Drak t al. 1998). ons' suggstingthat th
ultillular lrganismshav larg gn numrs nd many ll divisions nutral with rsptto s
ah gnration.Bausrrr Orrtionlrn nvr e prft,and th muta- fors (lik th balan
tion rat pr nulotid pr ll division is quit similar to that in atria, lir) provids a plausibl
uoltio of th Gnom 165
ynthsisand for a fw spializd or- th mutation rt pr nulotid sit pr gnration is muh highr than in
ial for organllfuntions hav n batria.For ampl, in humans it is stimatd to e aout 2x10 8 (Kon-
). ongoing gn trrlsfr has n d. draslrov2003)' Givn th numr of sitsthat od for amino aids in th hu-
; rvllirstransfrstlrt at nonfun_ man gnom, this translarsirrto a mutation at p 8nom fo hngs in
l th nLrlar]nollls of mny spis protin squnof aout 0.82 nw ltlttions pr individual h gnra.
tion, and around 80" of ths ar likly to slightly dltrious,yilding a
ral prssurs.Thr may a.lvan- nt mutation rat to dltriusamino aid mutations of aout 0.66. An vn
r rdrr.^d 8nom an rplit tstr, largrnumr of dltriousmuttions is likly to ilusd by muttions in
lur through dltilnsof grrsthat th nonoding portions of tlr human gnom (Asthna et aI.207),ringing
and intror-risquns.Th tinding th stimatof th human dltriousmutation rat to wll aov 1.
rlapping gns, in whih th sam igh dltiousmutation ats and th rsulting low population fitnsss
iffrntradingfranrs(Douglas t al. (row 1993) may limit possil gnom sizs for multillular oganisms.
tiorr fvord siz rdution irr ths Thortilmodls of th volution of grrsthat orrtrol mutatiol-lrats indi-
l sltivlyoeutfell,so that dltions at that mutational load favors a rdud mutation rat (Snigowski t al.
r d a r l i rf o r r d u n d a n tg n d u p l i - 2 0 0 0 ) .H o w v r . t h i s a n o p p o s i n g p r s s u r :t h o s t s o f r r o - o r r t i n g
ar unonstraindy sltitln, th mhanisms,whih slow down DNA rpliation and onsum llular r-
'r tinl bausrpliation sollrs.Th ourn of irdvantagousmutations lso disfv<lrsrduing
ristaks
n thn insrtionsf ss (Grgory rats to zro' ut in sxually rproduing spis with gnti rombination
llnts on gnsr rmovd' thy vyill twngns, allls that inrs th mr'rttionrat will not rmain assoi-
as mor dltionsthan insrtions - td with favoral mutations that thy happn to indu' and so thy gain
otlr sltiv:ttl{ntlnsltir,pro- littl dvarrtag. Mutati<ln rts of sully rproduing lrighr ukaots
ar thus likly t rflt th volutionary lan att:rindwhn th ost of
rduingth mutation rat quals th advantag of rduing th mutational
l o a d ( S n i g o w s ki t a l . 2 0 0 0 ) '
UMBRs oF GNs
In ontrast,in oganismstht produ largly without any volutiorrarily
ganis'sfuntions rdu survivl or fftiv gnti romirrtirn,suh as asual and highly inrding
tion is vry wak, thr will a l- spis,an alll that inrassth mutation rat will rmain prmanntly as-
w dltriousvaril-ltsinto a ptlpr'lla- soiatdwith any favoralutations it auss,and will thus inrasin fr-
. n . h p r s n t l f t h s m u t i l t i o n s queny as th mutation sprads in th population. This may promot highr
1l-ltation-fr population this mu. mutation rats than in sual spis.Tlris is an xanrpl of geti hithhik-
r of nw dltrilusmutations that ig and is osrvd in atrial populations that ar sujtto intns novl
:h tht th rrgativof th naturl log- sletionprssus(Snigowskit al. 2000).
i o n i s q u a lt o t h l o d ( r o w 1 9 9 \ '
r 'ill thus gratr lol if h
GNoM s|z lN RLA|oN o H AoUN
. R N g n o m sl a . . k r r o r - o r r t i n g
oF NoNoD|NG DNA
luttion ratspr gnom pr rplia-
.ound 1 nW muttior-r pr rplition Gnom.siz diffnsamong rlatd taa of higher ukryots oftn in-
t. 1998).Suh grromsannot, thr- volv larg diftrns irr amounts lf highly rpatd, nonoding satliit
aus thir fitnsswould bor vry DNA, with orrly minor diffrns in gn numr (avalir-Smith 1985;
orrtingmhanismsthat giv thm Lynh 2007). In som ass DNA amounts larly orrlt with diffrns
s;th . oli gn()lis stimatd to in lif history arrd ology. For mpl, slowly dvloping spis tnd to
r nr.r[otid pr rplition :rn{ mu- havmor satllitand othr typs of nrrodingDNA tl.rrifastr-dvlping
Drakt al. 1998). ons, suggsting that th siz of th nonoding portion of th gnm is not
n numbrs and nrany ll divisions nutralwith rsptto sltion(avalir-Smith1985). A modl of opposing
)n an nvr prft,nd th mllt- fos (lik th alan twn proofraling and mutation dsrid ar-
n is quit similr to tht irr irtria, lir) provids a plausibl and tstablxplarrationof ths pttrns. Svrl
l66 ullutilttof th ot
prossstlrt invo|v rr()rsin DNA rplition irrrdrtnqual rossin!]ovr tistial orrtions for diff.
n amplitr, rril\rsb multipls lf rvhllrptdunits, s thr r fors this is not ptd if thr.
tht inras opy numr' somtims adding DNA to th gnonrof an in- from salamarrdrs and n
dividrrl;gntidrift an tlrn aus it to sprad to fixation (B. halswlrth Johnston 1992).
t l. ] 994;' Lnc 2007). Sltiln against high opy nurr will oftn op- an w s th footprir
.
pos this pross' ut suh sltion is last fftiv in spiswith small sizs, as assumd rlir?
fftivpopultiln siz (Kimr.rr198). This suggststht thr should onntration in rgions o
rltion twnth fftivsiz f a spis (whilr an stimatlfrol w a n . T h r g u m n ti l r \ '
its lr.l of DNA sqlln .livsity;11,n2007) and th ontiutirln oi ray undrgos unqual ro
rptitiv squnsttl its gnom siz. Brod omparisons aross diffnt pliations). Unequal hz
taa suPport this prdition (l-ynh 2007). Hor'vvr,spissuh as D' uir- opis in th array. ut it i
ilis and D. mllgstr diifer onsidrably in thir mount of rptitiv r is nithr advantagous
DNA u sm to hav similar fftiv opulation sizs (Drrlsophil 72 spis to om fixd for
Gnoms onstlrtium 207I. his suggsts tht othr fators must plav rquirs multipl opis, a r
a ll. aftr rmain singl opy (t
A likli" disadvantagof a larg anlount of stllitDNA is a slolvr dvl- opis an lost again y
opnlllt tim, sin ll ]ir,isionswill slow lf it taks longr to pliat that afft array sizs, th
gn()mswith larg DNA nltunts(Cavalir-Smith1985). \Whnrapid dvl- a tandm arry thus produ
opmnt is important, th alan is tippd towrd lowr opy numr, but i{ ity ovr tim that th pop
slow dvlopmnt is not disfavord y sltion' sltion against th au- ially if unqual xhang
multin of rptitivDNA ,ill wakr, nd sr.rlr squnsan aLlmLl- individuals with high rra
l t ( F i g t r r 4 ) . T h u s l a r g D N A o n t n t d < l sn o t n s s a r i l yi m p h , t l r r rsulrs. Larg rays aris
sltiotlhs favord nroDNA (Pagland Jllrnsttrn1992'B. hirrlsworth (B. Chalsworth t a|. \99,
t al. 1994). low-romination gnomi
Of ours, it might somtilns advantagotlsto hav mor nonoding for that pushs rpat n
DNA. For instan' mor DNA pr nulus might nal organisms with for that ats in th ppos
larg lls to mov RNA mor rapidly into th vtoplasm (vali-Snrith No funtional advanta
1985). l. an on distingtrislrtwn ths possiilitis? pritrrnts DNA squns is tlrus nt
th altr th i.rrountof htrohrolnatin in D' rclogLlstrhardly rrfft thir distributin within g
viailitr, lr dvlopmnt tilr-r'suggstingtlrat larg hangs in opy num- latd spis oftn diffr
r s ( m i I l i o n s o f a s s )h r ' a t m o s t v r y s l i g t a d v a n t a g so r d i s d v a n - quns' whih also strong
tags (John and Miklos 1988). Vry slight fitnss diffrnsan, howvr, ]v{iklos 1988; Drosophil
importnt in vllution, so this is not onlusiv' In omparativ tsts dos not man that satlli
tht xnrin dvlopmnt tils and gnom sizs irross diffrnt spis, Drosopbil, portions of t}
th bln rrrodlprdits thilt rlationship will flund vn ftr st..t- mosomal hvior at lnlo
volvd in ntromr [un
probaly did not aus t}
insrtion5,
5ma|| in|uding -..--->
ratdditions
st|lit
mor likly, ths arrays v
insrtis,
|rge i|uding
s lir and wr latr o-opt
small lrge
genome genome RANsPosAL
<- prssur
undrmUttiOn
dltions highostsof rpling
DNA ost typs of transposa
<- fvord
s|tivIy deletins slowgrowtfl t h m s l v si n t o n w p o s i t
<--UPU3| rossing
ovrwithgeneti
drifl nal opy. Ths transpos
m n t n u m r so v r t i m u
Figur 4. Opplsing fors atir.rgOn gnom.siz vrlIution. For ampl e, tn D. mI
uoltio of the Geom 167
liationand unqual rosslng ovf tistialorrtions for diffrns in nular volum and ll siz, whrs
rptdunits, so thr ar fors this is not exptd if thr is an advntag of more DNA. orrtd data
tding DNA to th gnom of an in- from salarnandrs and nWts still show a signifiant rlation (Pagl and
sprlto fition (B. hrlswoth
Johnston 1992).
lst high op numr wil| oftn op- an w s th footprint lf a mutational prssur that inrass rray
'ast fftivin spiswitlr
small sizs,as assumd arli? Th distriution of stllit DNA, with a strong
This suggststht thr should onntration in rgions of infrqunt gnti romination, suggsts that
lis(rvhihn estimatd fom w an. Th rgumnt involvs a simpl modl in whih a tandm rPat ar-
nh 2007) and th ontriution of ray undrg<lsunqual rossing ovr (plaind arlir for tandm gn du-
Brod ompisons ilross diffnt pliations).Unqrral xhang dos not hang avrag nurrs of unit
7).owvr, spissulr
as D. uir- pisin th array, t it inrassth rpat numr rang. If rpt nunr-
aly i thi amount of rptitiv r is nithradvantagousno disadvantagous'gnti drift may irusth
population sizs (l)ros-lphil 12 spisto bom fid for just a singl opy. Baus unqual rossing ov
.sts that otlrr fators must pla rquismultipl opis, a rgion that rahsth singl-opystatwill thr-
aftr remain singl opy (unlssa nw dupliation ours). Howvr, xtra
t of stllitDNA is a slowr dvl- opisan lst agin y unqual hangs.In th asnof othr fors
siow if it t:rkslongr rl repliat that afft array sizs' th omind ffrs of drift and unqual xhang on
i r - S m i t h1 9 t l 5 ) W
. hn rpid dvl- a tandm aray thus produ an asymmtry:thr is an inrasing proail.
l towird lou,r opy nun-rr,lrut if ity ovr tim that th population finds itslf with just a singl opy' sp_
ltion,sltionagainst th au- ially if unqual hang is frqunt. If thr is som sltion gainst
r, and sulrsqunsI1aulnu- individuals with high aray sizs, an quilirium disrriution of array sizs
lt dos not nssarily imply that rsults. Lag arrays aris only if rossing ovr is xtrmly infrqunt
rdJlhrrston1992:B. hlswoth (B. harlsworth t aI. 7994). Th lsrvdssoiation of satllit rrys with
low-rominationgnomi rgions thrfor suppots th hypothsizd wak
ltltgousto hl, nror nonoding for that pushs rpat numrs upwd; w s its ffts only whn th
:lusrnight nal organisms with for that ats in th opposit dirtion (unqual rossing ovr) is rmovd.
n t o t y t o p l a s m( a v l i r - S m i t h No funtionl dvantg to hving larg loks of tandrnly patd
ln ths possiilitis? priments DNA squns is thus nssary in ordr to plain th main faturs of
n in D. mllgstarhrdly afft thir distriutionwithin gnomsand among diffntspis.Morovr, -
g tht larg hangs in opy num- latd spis oftn diffr gratly in th typs and amounts of satllit s-
' r y s l i g h t a d v a n t g so r
disadvn- quns'whih also strongly suggststhat no sltionis involvd (John and
lt fitnssdiffrnsan, howvr, Miklos 1988; Drosophil 12 Gnoms onsortium 2007). Howvr, this
t o n l u s i v .l n o l n p r t i v t s t s dosnot man that satllitsnvr hv nssaryfntions in organisms' In
t o m s i z s r o s sd i f f r l r rs p i s , DrosophiI, portions of th htrohromatin ar ssntialfor orrt hro-
nship will found vn ftr sta- mosomal havior at miosis, and human o-satllitsqunssm to in-
volvd in ntromr funtin (Sullivan et al.2001). But ths funtions
dditions---------> proaly did not as th vlution of ths squnsin th first pla;
mor likly, thsarrays wr first stalishdy th prosssmodld ar_
lir and wr latr o-ontd for ll funtions.
+ lrge
- genome RANsPosAL LEMNs
highosts
ofrpling
DNA Most typs of transposal lmnts (Ts) (Tal 2) can insrt opis of
slowgrwth thmslvsinto nW positions ir-rth grrom' oftn without lss of th origi.
nal opy. Ths transpositions also rat a prssur toward inrasd l-
mnt nums ovr tim unlss som othr fors kep T numrs in hk.
-siz volutin.
For ampl, in D. mlogstr it is stimatd tht on avrag t last on
168 urlltilnof th Gom
ltl gnomvry 10 gr-rrations in rarrangmnts hav fw or no harmful ffts and n sprad in populations
rlad(asid t al. 2000). h
bausof gnti drifl, thry orrrriuting to volutionary h:rr-rg in lrro-
gnom(Karninkrt al. 2002), so
mosomstrutur. But many rarrangmnts disrupt gn funtions nd r
Llly tiv lmnt is alut 10_apr
highlydltrious:nrany humn gntidisssr ausdy hromosom r-
rrangmnts(glmi disclss),inluding mny ns (Diningr t l.
in th gnom? Thr i1two on.
2003).If this is n importalrt disdvurntg t having high T aundan, it
94). First, Ts ould maintaind
. host orgarrisrs; prditstht Ts sould altmulat mainly in gntrmi gions rvith low -
for instrr. thy
ominrtionrts' sin tlry ar tlr lss likly tl us harnrful rarrang-
rtiorrsy insrtinginto gnsOr rg-
mntsnd will thrfor not rmovd y sltion against th produts o{
ida is tat insrtions f]lostly atls
topixhangs'As mntiltldarli, this is oftn tru' For anrpl, a larg
srllting oding or gulato s_
pportion of tlr Y hromlsms of many spis onsists of -drivd s_
triousmutations' inluding som
quns(B. harlsworth t a|. |994|. Howvr, thr ar othr possil x-
srtionsinto oding squns (os-
planatinsfor this pattrn' srrhas th dud ffiiny of sltirrin rgitlns
|y dltioLrs muttions r usuall1,
of th grromwitlr rdud rorintion(sltr disr"rssiorr).
d as mntiondarlir, it is inded
Ts miry somtimsindu favoralmutations or hromosom rarrang-
f highr organisms Ts ar largly
mnts'and sltionwill thrl sprad thm throughout th plpulation.Son-r
g tt irrsrtionsther r harmful
-drivedsqunspp21rto rgultgn tivity (Diningt l. 200)
ing onsortium 2001; Kamirrkr
or onfr othr nfitson thir hosts (Brookfild 2003). As w argud for
:vidnthat T movmnt is disad-
satllitsquns'howvr, a prsnt-daynfitdos not impl,v that sl-
lis, suh as st' rh nrold Nzl-
tion prlmotdtlr initial sprd of tlrssqtlns.lr is possil tht rgul_
lhanisms th:lt rstrit h rat of
tory or r,nodin8 funtions hav n aquid by suh Ts long aftr tlry
r aL.2007). orovr, t viw tat
wrinsrtdinto thir Prsnt loations and fid in th population.
hosts' futr:rl.o]utionsts on th
is is nhandy th prsn of
:om stalishedin th Iirst pla. MAssIV AUULA|oN AND PoPULA|oN
ir ailit1'to sprerd y slf-rpliation sIz DIFFRNs
ssion of nw opis to th offspring
Although th rat of inrs of Ts y trnsposition in spis suh as
rsuallyhav harmful ff:tson thir
D. melnogsle/smslow (on nw lmnt in th haploid gnom lmnt
h i s v i r , , T a u n d a n w i l l d p n d
oughlvvry 10 gnations)'opl,numbr lvill inasrapidl,vovr volu-
t attssinrasdopy rrumrvr-
tionarytimsalsif no opplsingfo oprts:for xlmpl,with th rat of
g opy numrs (B. lrarlsworth
transpositilnstimatd arlir, a tnfold inras woLlld tak only 2,000
mplis that th rdution in fitnss
Drosopbil gnrirtiotls,lss tlran 2,000 yars. Lrg diffrnsin T Lrrr-
vrgqual tl-rhan tht an l-
dansamong rltd spis,s in th maiz and ri ampls dsrid
r a r o u n d l 0 - a p r g n r a t i o ni n
arlir,a thus radily plial if sltion prssus against thm r
:ssurmust wak. Thls has r
smtimesrlad. Arrothr ampl is provild Droslphil mirnd, itl
:mininglmntpositions on th gi.
whih art autos()nlm attahd to th Y hromosom out 1 million
ls. Th frqunisof T insrtions
yrars ago; this hs sin n transmittd only from fthr to son, ausing
oW at a givn pla olr t hrt>mo-
this gnom gion to stop romining usrrmirrtiondos not o-
lisadvantagos. Th strngthof s-
ur in Drr,,sophil ma|s.h rgion has sin thn aumulatd a high dn-
's,and it agrswith th stimatd
sityof T insrtionsinto nonoding squns(Bahtrog 2003)' Gnoms of
19941.
somspis(.g.,lrumns arrd miz) lrva vry hig ovrIlT aundarr
V lrv alrady disussd sltion
(abl2), and at many hromosomal sits, Ts may b prsllt at high f-
[ftsof transpositions on th host.
qunis(or vn in all individuals) (Ostrtag and Kazazian 2001)' ontrast-
g ovr btwnlmnts irr diffrent
ing lvirh th low instilnfrqunisin Drossphil.
h i s u s s h r < l m s o n tin v r s i o l r s ,
\fhy ar thr suh larg diffrnstwn spis?Prhaps th ruh
(B. harlswortht al. 1994). Som
largr nrounts of intrgni DNA and largr introns in humans than in
1 7 0 uolutio of tb Geom
Drosophil 1nthat insrtions hv lss risk of ausing harmful mutations Gns irr gnonrr.gion
(although this would not tu if intrgnirgions' squnsoftn ontrol similar volutionry pros
gen ativity). A mor intrstingpossiility dpnds on th waknss of s- ls of adaptation. his ma
ltion against ah nW T insrtion. If Ts rarly is from a sit, gnti omirring hrotnosomal rt
drift an somtims lad t a sit oming irvrsilyfid for an lmnt. (ph:rpslso in pIt Lls
In spis with low population sizs, suh as humans, sltion against l- strikingly illustrr th
mnts is lss fftivin opposing gnti drift' and this possily allows th ( D . h r l s w o r t h t a i . 2 0
prssrrrfor lmnt uildup to prdominat (Lynh 2007I. Nw insrtions s o m g n s ' t r ' i n gr h
into lady_upidsits wold flot prvntdy sltion(thrwould pai, ut tlr Y laks n)()st(
e no diret hrmful ftts,just a slightly gratr risk of topi xhang), th gnti informtion on
and so lmnts ould snowall ovr tim. This may aount for th fixd l- nonr()mininghromoso
mnts in mnrmalian gnoms' ut rrot for th vry high T aundan in p d t h i r i l i t yr o m a i n
maize, a spiswhos high DNA squnvalaility indiats a larg pop- fastr than tir -linkd
ulation siz (Tenaillon t al. 2002I. Romination in maiz may largly d i s d v e n g o u sr . e r i e r l t si
onfind to gns' so that Ts in intgni squns ould not rmovd gns thn lik normal gn
y topi hang. Thus lmnts would uild up twn gns' as o. s s o f s . l i n k dg n si n
srvd (Sanigul t a|. 1.996)'owvr, th austion ould th othr ( F i d o l i s s o n n d l l g n.
way round: Ts may simply romi lss than l.lostsquns. gns addd to th Y hro
onfirm hetthis is du tll i
mutations (Bartollrnand
H VoLUIoNARY oNsoUNs oF GNI
Similar ut mildrfft
RoMINA|oN AND PoPULAloN s|zE
gions with diffntroml
SltionagainstTs is only on Way in whih small population siz and lak odons that nd irr G or
of romination an importnt for gnom volution. Romination ontnt of third oding pos
rts play important ols in numous voluionryprosss'spiallyth o th djant r-rontrdi
volutionary advantags and disadvantagsof sual rprodution. Svral gs, and ' thlitl, ter
diffrns among gnom rgions r onntd with romination rat flts th ation of sl
diffrns' nd th ffts of low ronrination rts ar similar to thos of fastr rrd/or mor aurt
small fftivpopultion siz (Grdo arrd Chalsworth 2001). tin squir(Akasi t al
Sx and rombintiorr llow sltion to at indpndntly on diffrnt sits is' horvvr, vr1' small o
within th gnom. This inrassth ffiiny of sltion.For ample,if fa- protin squns.so that
vorbl mutaions aris indpndntly, but vry rrly' at two sits, th popu- prfrd ovr A. In an o
ltion will rarly trntainany individualsthat arry doul mutants'vn if this omPonnt of tlr gllomr
is th fittst omination (unlssth popultion siz is so hug that th sond whr romination is in1
mutation vnt happns for sltion fis the first favoral mutation). G- mal odon usag ight
nti rombination allows th doul-mutant omination to e formd. would lss isdthn
Whn th is littl or no enti romination' or whn rprodution is asx- o f t h o s r r ' r ]r l l t i o n s
ual, sltio on on gn thus impds sltion ating on othr gns' making ontnt of oding squn
fvoral mutations lss likly to sprad and dltrious muttions (suh as and for th wakning of
amino aid hngs in prtins) mo likly to fid y gnti drift, pati- ttd for th gns addd
ularly whn population sizs ar low (Gordo and harlsworth 2001). This and harlswlrth 2006).
Pross proaly ontriuts to gnon-lrdution in symioti atria that Horvvr, th G ()nt
liv within inset lls (romination btwen tria of ths spis is pr- . lgs also <lrrltsv
vntd by isoltion);thsrdudgnomsalso hav fastramino id squn ing sqttnsand rrontr
volution' as prditd givrrth lowrd aility of sltionto liminat muta- m n t i ( ) l ] d l r l i r 1 \ { r l r
tions with wakly disdv:rntagous ffts(oran 2003). rats i1nd dirtiolls of mut
uolutkl rlf the (}ut 171
rats. rtainly, not all muttions oLlr qually frquntlv' As lrady mn- latiol.lsizs drsdas
tiond, th rlativly high A ontnt of nonoding DNA is ausd by muta- trons xp()ss gns t() ]
tional bis toward AT. A pross that may oppos this is bisd ge hang th prltin irnd
onursion; aus of th molular mhnism of gnti romination' spliir-rgmust Onsv
whn on gnom has G at a sit and th sam sit in th oth has AT, r. y s l t i o nt l n l y i n s i
omination an podu a vry slight ssof G in th daughtr gnom prssur inrss intron
(arais 2003). This will inasG ontnt in oth oding and nonoding pokllry<ltsand prs.ll
squns' ausing Gs to fid mo frquntly than ATs' just as though 2003; Lynh 2007). Sl
Gs wr favord by sltion.Baus it is pat of th romination pro- o r r h r e s o n s .h u i n
ss, isd gn onvrsion is xptd to at mainly in rgions with high Again, lr<lrr.vr, it s
rats of romination, lading to a highr G frquny in suh rgions. Prsn o l l i n t o n sl l l
Thid positions of odons will hav an dditionl nhanmntof G from aus intron siz and r-
th fftsof sltion for odorr ias' Statistialanalyssof th G ontnt h l p p r l . i , f t l i t l s t l n .i n
.Whatvr
of th gnom of D' mlnogstel"supPort this hypothsis, as do data from t l. 2002). tlr
th highl romining parts of sx hromosoms of nrammls (arais to lm domstiatd
2 0 0 ). suggstdfor tltlrr gn<l
ltiv onstritints on intr
kryot gnoms (Hallig
nlusions F.urtlrr progrss in unl
volution is to ptd
Gnom volution, lik othr volutionry hangs, is opportunisti. hangs trm possil. Th Tw
in gnom faturs hzrvonsqunsof two kinds. First, hang may lad to gnoms <lf 12 rnmrs.
futhr hangs. For instan, hr{rmosomrarangmntsafft romina- stP in this ]irtion(Drt
tion. A gn an mov into a rgion of lowr or highr romination, afft. th rtnsntlrrlgsin p.
ing th ln twn iasd gn onvrsion that pushs G ontnt up partitllar vnts in gnorl
vsus mutation Prssur in th othr dirtion, So that GC ontrrt will start ing spissoirld allorv s
hanging. If a rarrngmntrdus romintion, as happns whn a hro- auss Of many hangs.-
mosom arr is joind to a Y hromosom, th drasdaility of natrrrl s- possil to onrpar indp
ltion to liminat wakl disadvntagoushangs may ultinratlylad to mon frors, sirrrihr to th
th loss of funtionality' or vn th physial prsn'of gns from this hro- volr.ltionayhngs(.g
mosom' as indd has hppnd in D. tird (D. harlsworth t al. 2005). ous tsts of hypothssthl
Sond, vn whn a gnonl fatur r,olvswitlrout ing favord y in-
dividual sltion,th an oasionlly ptntially advantagouseffts BILIoGRAPY
for organismal funtions. For ampl, Ts that ontain squnsthat on-
trol th prssion of thir own gns n lso fft th ativity of gns A k s h i 'H . , R . l \ { .K l i m n ,a
s | t i l r rl r n d h r ' t > | t
los to thir insrtion sits.Initially parasiti slfishDNA (spradingpurly
49-60.
as a onsqunof sltion for th squns'own aility t rpliat)an Th Aridopsis Cnlm In
..domstiatd''
bom as a part of th gnom of a spis,prsntin all in- flllvring pltt rbidl,
dividuals nd funtioning in its host, rathr lik th organlls'whih wr Arvin, A. A.' G. J. Hantln
on indpndntorganisms (Brokfild 2003). This possiility is often ovr- rovids an alirtivd
lookd, ut it is important to avoid th naiv attitud that prsnt.davn. Asthl-r,S., \. s. Nol, G.
St:rnll-ttovnnlgl<rul2
fits an plain th origin of irll gnom fatus.
tl.rhuma gt.ltlm.Prr
Genom vlution is far mor ompl than straightforward adaptiv vo- 12410-1241 5.
lution y natural sltion,and many fatursmay hav hd a long history in Bahtrg' D. 2003. AunrL
gnoms for ny nfis t<l th organism volvd. It has vn n lIrtlr
trtrl.lsp()s<rns,
Bitll,lgl, llt] L u,lltt!i,'tt
suggstdthat introns may initially hav sprad within gnomswhn popu-
uolutio lf th col 17.1
Barns, T. ., Y. ohara, A. oulson, nd S. kim|. 1995. ilti romination, Gray, . W. 1999. volutilrro
nonoding DNA and gnmi organiztion tn norhbditis lgs' Getis d Dulopmt 9: 67B_(
147: I59-179. Grglry, T. R. 2004. lnsrtton
Bartolom, ., and B. harlsworth. 2006. volution of minl.idse1unsand G 24: 1-4.
ldon usg on th DrOsOpil mird no-sx hromosoms . (]tis 174: Hallign' D. L., and P. D. Kig
20-2044. D ro sop b il gt.lomrvaI
Baumgartn,A., S. annon, R. Spnglr' and G. ay. 2003. Gnm-lvl Rsrh 16: 875_884.
volution of rsistan gns in Arbidopsis thli. Getics 165 9_319. a r v v ' P . ' , a n d M . P a g l .1 1
Bnsasson,D., D. . Zang, D. L. Hartl, ar-rdG. . witt. 2001. itohondrial Bioklg'. oxford: oford l
psudogns:volution's misplad witnsss. Trds i llog d uoluthn Intrnatilnluman Gnom l
16:314-321. analysis of th human gn
Brookfild, J. F. Y. 2003. oil DNAs: Th poahr turnd gmk epr.urrt John, 8., ar.rd(i. L. G. Miklos.
Biolog 13: R846-R847. uolutio, Lndon: Allrr l
Bruggmann' R., A. K. Bharti, H. Gundlah, J. S. Li, S. Young, A. . ntaroli, F..S. Joy, G. F. 1989. RNA vlut
\Wi,G. arr, G. Fuks, . G. Du, . Rymond, . . stp, R. Y. Liu,.|' aminkr, J. S.' . . Brgma
L. .!7hlr,
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ihlr, . ., and D. Sankoff. 2003. Strutural dynrnisof ukaryoti hromosom symionts. urrt Opii
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gnson the Z and.W sx hrmosoms. Gtis 15.5:1903-1912. 1 12-24.
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uolutio tf th Gom 17-\
]ekimi' 199.5.ioti ombination' Grv,. !. 1999. volutiorr f organllar gnoms.urrat piio i tis
l in (,orhbditiselgs. Gtis d Dukl1lmt9: 678_687,
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llution of amino-id squnsand Cut 32.4:|-31.
o-shrl-lsot-t-ts.
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is thlin, Getis 165 9-]-9. Harv,P. H., nd . Pagl. |991 , h tlmprtiu thod i F'ullutior
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nlvsisof th hum !.n()m.Ntur 409: 861-921 .
ohrturnd gamkpr.urrt John' B., and G. L. G. iklos. |988. h ukrot Glm i Duklpttd
uolutirm,Lonrin: Alln rrd UIlwin.
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.!(hlr,
J. Quknush,!. B. Barzuk, D. A. S. . Lwis, G. M. Ruin, . Ashurnr, and S. . lnikr.
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1 7 6 uolutiln of th Gome
2000.The evolution of
BioEssays22: Speciation,the evolutionary processby which new and distinct lineagesarise
and mirintain their independenttraiectories,has been a perplexingphenome-
'terminingcentfomere
non ever since Dirrwin (18-59)first proposed that all living organisms have
uiewsGenetics2:
diver:sified frorn sharedancestors.There are severalreasonswhy unclerstand-
ck, J. F. Doebley,and i n g h o w s p e c i a t i o no c e u r si s c h e l l c n g i n g .F i r s t . m o s t s p e c i r r i o nt v c n t s h r v c
ion alongchromosome 1 occurredin the past, leaving only the end products, species,:rs a signattrreof
- 14 1 3 . their occurrence.Species,l.r<tth exrant and extinct tornts, [re,rrthe cher,rctcris-
rt. 2002. Evolution of tics rlssociatedwith their divergencefrom conlrlon ancestors'but in addi-
ing invaded by
tion, they also carry unique phenotypic and genotypic changes that have
ion 19: 1154-1161.
occurreclsince speciation.Disentanglingthe characteristicsassociateclrvith
ew fnnctions? Genetics
the speciationprocessfrclm those thzlt have evolved since divergellcecan be
,olution and particularlydif6cult.
4sof tbe National Second,new speciesarise through r'rvariety of evolutionary mechrrnisms
(e.g.,natural selecticln,sexual selection,genetic drift, anci mutatioll), and
of recombination rateand is
therefore,finding a unifying definition that describesand clefinesall sf-'ecies
Arabidopsisthaliana.
an impossiblechallenge.Thus much clebatehas occurredal'routthe nature of
a n d p a t t e r n so f m o l e c u l a r speciescharacteristicsand how scientistscan distinguish speciesfor taxo-
lar Biologyand Euolution nomic,systelratic,and conservatiorrpurposes.
Finaily, speciationis a seriesof processes, with :r beginning stageof initial
nultiple vertebrate a middle stagewherein species-specific characteristicsare refined
divergence,
tns. Proceedingsof the
bv vari<,rus forces of evolutiotr,and an end point at which ir new speciesbe-
)L.
slrion.gnti drrft. a
Speies Speies
gnotypi diffrnsbt
how nd why nw spi
in whih nw spis r
primary mans y whih n
dnt voIr-rtionarypathwa
hv gul.rto rsllv th
xllo
Th Natur of Spi
Spis onpts, :r st ot
a o u n d i n t h l i t r a t u r ( }
llng-standingdat enro
lm is ntrd arlund ir
llll. pttrn-orintd <lrPr()
Dlvr frnsof opinion tln wht
plial (Figur 2). P.rtt
spis. For ampl, th
Common y Thodositrs Dozhrs
( | 9 4 2 , \ 9 6 ) , d f i n ss p
Anestor
nisms. Rprodutiv islla
arrirs to sussfulrp
Figur 1. Th prossof speiationis a ntinuum.InitiaIdivrgntwn
Poss-orintd dfinit
populatinsis followd y priod of angnihangwithin ah nw linag
m h a n i s m s p, h n t y p i
whrin spis.spifihartristisar rfind y various fors of volution.
In som spislong pridswith littl volutionaryhang(stasis)may ()ur. ion of som lvl of div
one formd,spismay furtherdivrsifyinto nw linags(ladognsis) or s p e i s i n t t l d i s t i n t n t i t
may om tint. Th hallngin th study of spiation is unovring ( v a n V a l n 1 9 7 6 ) d f i n ss
whre along this ontinuuma spisatuallyxists. l q u i l n t s n d l d n '
from othr spis thr<ru
llogial hartristlst
volutionary iologists ovr th past 15 yars has ld to mny iting ad- grphi xhangaility'' 1
vans. Ths inlud th disovry of svral spitio gs that onfr r- Al:rn Tmplton (1989)
produtiv isolation, a ttr undrstandingof th importan of sympatri uniqu llogiirlprr)prt
spiation (spiationwithut gographiisolation);and a surrtionof in- volutior-rarySpis <
trstin how natural sltionan lad to th formatilnof nw speisas y- s p ar e t v o lu t i o n 1 r' l j
produts of adaptiv diffrntiation. on pross' that is, voluti
In this ssayw outlin som of th major hallngsin undrstandingsp. l u r i o n a r y l i n a g d f i n i t i
iation and showasxampls that illustratth major advansin th study sin n formlizd with
of th origin and divrsifiation of spis. w start with th prolm of and ollagus in th Phy
dfining spis, and how an undrstanding of th undrlying mehanisms Spis ar d{inds sP
that lad to th formation of nw spis may the st approah in at- that rsLlltrorn divrgn
tmpting to dfin what a spisrally is. Nt, w outlin th various ryps s p i sd o n o t o m d i n
o f i s o I a t i n g r r i st h a n n i n i t i t 'p r o m o t .a n d o m p l t t h s p i a i o n ompltly sd (FigLrr
pross.Finally' w disussth volutionary mhanismstht driv th spi- Wht do w do with all
ation pross. Outlining th rlativ rols tht natural sltion, sxual dfin spis irnywy? T
hc Pttr d Prlssof Spitil 1 7 9
Th Natur of Spis
Spis
onpt Dfinition Rfrns
Rfrns Lr'
intrbrling D o z h a n s k y1 9 3 7 ; BS
lulationsthat ar 1942,1963
ltivl1,
isolatd
srrhgroups.
-rIusivgroup T m p l t o n1 9 8 9
ls havingth
lr gnti
rgrphi
rility.
;anismstht Sokal and rovllo
r to hoth 1970
tint fom oth
pultinsrhat Patrson-l985
nmon
l systm
rt-rogniion
nlusivgroup Tmplton1989
ns havingth
)I gntl Figur2. An illustr:rtionof lrorv diffr:ntspisonpts dfi.rspisat
rographi diffrntpoints in th spiationprossaftr th splittingof a sil ling
rility. into two. lfhrr divrgntlinagsdisprsinto diffrntaitats'ih tgial
adptd van Vln 1976 Spislnpt(S) will dfin tlrrns sprtspis.Th volutin of
;arrisms
ologilnih, rprodutiv isolation(ISM1 or ISM2) twnth two lincswill lfin
paratlyfrom sarat spisundr th BiologialSpisorrpt(BS). iinags must
tsidits rang. rrsnt monlphyltiiads(all p<rpultilr.rs irdt-snd.i frlm singl
o m m o na n s t rl)r f r l
i n gr t l g r l i z ds s p i su n t l t h P h y I o g n r i
tag Simpsn 196 1;
Sis onpt (PS).
l-lsnd nt Wily 197tl
f populations)
paratlyfrom
with r"rniqr-r sintistsall spissm to know thir own oundaris' vn if billgists
.y tndnis
nnotsm to agr.Thr ar, howvr' som vry imprtant sons ftrr
r lristorialfat.
h a v i n gs o m f o m o f d f i n i t i o nt h a t s i n t i s t s n u s o o g n i z s p i s ,
sr digr.rosai a r f t1 9 8 ,
ven if diffrnt speis rquir different ritria for dfining thir ound-
rdividual 1989
aris.First, dfining iologial ntitis s spisprovids a svstmatiman-
withinwhih
arntalPattrn nr in whih to lssify divrsity and is th sis of modn taonomy.
and dsnt. Sond,dfining diagnosal groups as spis rrsponds to th disrt
ntitisthat ist in natur. Third' on annot study th pross of how nw
l r s f s i l l i n
spisaris without having som dfinition rf what a spis is in th fist
pla. Forrth, spis, s th last inlusiv tgory of livrsity, rprsnt
thvolutionaryhistory of organisnrs'Finlly, iodivrsity is gnrally ma-
surdy th numr nd kinds of living organisrs,and ths stimtsar
usuallymad at th spis lvl. Thus for a varit of asons, inluding
atgtlrizingand onsrving divrsity, as wlI s undrstanding its oigitls'
182 h Pttr d Pr-lss
of S1litk
olli
Th Natur of IsolatingBarrirs
tI
. r t i l i z n t i ll n d o n r r i l ] gr p r ( ) .
taa ar mor likly to xhiit aldan's rul; lldr taa oftn show svm-
omplt th spiatron
mtri hrid prolrns that fft orh ss.
ovrlap in gographi rar
Although Haldn's rul has primarilv n osrvd to rsult in hybrid
srv 5 strong ai
strility or inviaility of th htrogamtisx, sol vidn suggststhat
whos ranges ovrlap (s
htrogamtihyrids ray suff draslfitnsstlrrough a varity of othr
or whn prviou.slyisoll
mhanismsas wll. For arpl,Nal Davis anl lllagusshowd tlrt in
o n d r y o n t a t ) 'i n t r i n s
th Notropil uttrflies Arti ftim and . mtb strong assrrtlrtiv
rding om ns
mating urrd within ah spis'ur whn thy wr rossd artifiilly,
importan of rprodut
fmal hyrids showd dud trtiiity,onsistrrtwith ldan's rr-rl(t-
of spiation' and muh
mal uttrflis ar htrognlti;Dvis et aI' 1997). In addition to this
n t r do n r h m h a n i
mor typil rsult,howvr,hyrid fmalsfrom th . mtb (fmal)x.
twn spis'
ftit (mal) ross showd rdr'rdtnlnyto milt (Dvis et a|. 1997),
lassi modls of th s
his study provids vidnthat Haldarr'srul mzryxtnd yond th typ-
bution of populations un
ial osrvation of hyrid strility or inviaility, and that hyrid {itnssmay
gographi isolation to t}
drasdthrough th prmating and postmting arrirs'
d g o f g r t g r p i s
A omplt gnti undrstndingof postzygoti islating isis n.rh
an important ontriutol
mr omplx than aldan's ul aln implis. Of partiular intrstis th
of spition hav fous
onpt of speitir gns. Speiation gns ar gns that diffr tr-l,n
(natural sltion,sual
ta and us rprodutiv isolation trvnthn-r.A gn lld d1,55g'',
d i v r s if i r i o n e n d s p i
for ampl, shows vidnof rapid volution in srmfruit flis,dspitrl-
ativly slow volution in other organisms. This gn appars tl rsponsi-
l for strility in hyid mals trvenD. simllusnd D. mouritia (Ting H RoLE oF NA
t al. 1998). Lik olyssesJ]nsthat prinrpid voltion(t lilst in
Th imortan of adapt
som taa) may likly to us gnti inompatiilitistwntaa and
llgicl spcitio) Was fil
thus good andidats for spiation gns.u of th wrrk ln sPla-
in th 1940s in dsriin
tion gns rmains thortiurl; howvr, volutirr:ryilogists r addrss-
livd that spititlnin I
ing iting qustiorrssuh as whthr partiula gns ar gnrally rnor
divrglrt physiologia1r
likly to lad to rprodutiv isolation, or whthr gnsassoiatdwith r-
m n t s . n s M y r \ | 9 4 2
tain funtions ar ttrndidatsfor spi:rtingnsthn tltlrrs.
latd gnti diffrns
a n d | o g i e la d e p t a i o
rprodutiv isolating mt
Th Prossof spiation
dutiv isoltion is th dri
allopatri spiation (FigL
volutionary mharrismssuh as natural sltion,sual sltion,and g-
Nl[or rntly th imp
nti drift an promot diffrrrstwn populations in phnotyps and
d u t i v i s o l i o n i t h ri n
thir undlying gnotyps'vl1to th trrttht diffrnt ;lopuiatiorrs-
n important rl fo natr
om fixd for ontpltly diffrnt ltrnativs. Why, thn, is not vry
om lar. Laoator1
larly diffrntiatdpopulatiln nsidrd a diffnt spis?Th ru of
hav ofirmd tlrat o
th prolm sts on two issus.First' htl.,vmuh gn flow lurs tr-l,n
thr rzrisdfruit flis (D. l
diffrntiingpopultions? Gn flow :rtsto honrgnizpopulatiorrs both
vironmnts (varying tml
phnotypially and gntially;hn it works against slti<lnand drift in
s o m l v l o f p r n t a t i n gl
promoting daptiv or randrt diffrntiation' rsptivly.Sond, lrow
in diffrr-rtnvironmnts
muh diffrntiation is noLrgh?This issu is thlrIrirand points tO th im-
dpndnt lins raisd in
portan f rprodutiv isolation in th spitionpross.Although divrg-
s r t h o w t h p r r l s sm i
ing populations do not hv to volv intrinsi arrirsto rprodution, som
In th simplst modl
brrir to rprdutiv harrg mllst prsent to sto gIr florv and
volvs twrr popuiati
h Pttrn d roess of Spcitio 187
ul;oldr taa oftn show sym- ompltth spiation proess. For allopatri populations' whih do not
ovrlapin gographi rang' th physial distan imposd y gography an
)nosrvdto rslllt in hyrid srvas strong arrir to gn flow. owevr, for divrging populations
s' som vidn suggststhat whosrangsovrlap (sympatry)or ar adjnt to on anothr (parapatry),
l fitnssthrough a varity of othr or whn prviously isolated allopatri ppulations om into ontat (s-
lis and ollagusshowd that in ondaryontat),intrinsi or trinsi arrirsto gn flow that prvnt intr-
n d . m t h s t r o n g a s s o r t t i v rding om nssary to omplt th spiation pross. Thus th
hn thy wr rossd artifiiurlly, importanof rprodutiv isolating arrirs has n paramount in studis
nsistntwith Haldan's rul (f- o{ spiation,and muh of our undrstandingof th pross of spiation is
t a|. 1997). ln addition to this ntrdon th mhanisms that favor th volution of rprodutiv isolation
from th . mth(fmal)x . btwnspis.
ny to mat (Davis et a|. 1997). lassi modls of th spiation pross fousd on th gographi distri-
s rul may tnd ylnd th typ- utionof ppulations undrgoing divrgnnd th rlativ ontriution of
ility,and that hyrid fitnssmy gographiisolation to th formation of nw spis(Figur 4). Although the
ostmtingbarrirs. dgreof geographi sparation twn divrging popultions is rtainly
tzygotiisolatingarrirs is muh an important ontriutor to th disruption of gn flow, mor rnt studis
nplis.of partiular intrstis th of spiationhav fousd on th importan of volutionary mhanisms
ns ar gns that diffr twn (naturalsltion,sxual sltion,gntidrift, and mutation) in promoting
nthm.A gnalld odysses, divrsifiationand spiationithr in allopatry or in symptry.
tion in sm fruit flis, dspit rl-
h i s g n a p p r st o r s p o n s i -
H RoL oF NAURAL sLIoN lN sPE|AloN
simulns and D. muriti (tng
rinrapid volution (at last in Th imprtan of adaptiv divrgn in th spiation pross (trmd eo-
nompatiilitistwntaa and logilspitio)was fist notd y th gntiistThodosius Dozhansky
rs.Muh of th work on spi- in th 1940s in dsriing spiation in Drosophil. Dozansky (1946) -
rolutionaryillogistsar ddrss- livdthat spiation ln Drosophila ourrd minly through th volution of
rrtiulargns ar gnrally mor divrgntphysiologial ompls that wr sussful in diffrnt nviron-
whthrgnsassoiatdwith r- mnts.rnst ar (1942) Was th first to point out that many of th umu-
:iationgnsthan othrs. latd gnti diffrns twn populations' partiularly in physiologial
and ologialadaptations to partiular nvironmnts' Wr also potntially
rprodutivisolating mhanisms. Suh ..y-produt'' volution of rpo-
dutivisolation is th driving for hind ayr's lassi mod f viarian
allopatrispiation (Figur 4).
sltion,sxual sltion,and g- Mor rntlyth importan of ologial divrgnin promoting rpro-
n populations in phnotyps and dutivisolation ithr in allopatry or in sympatry has n rognizd, and
tnt that diffrntpopultions - an importantrol for natural sltionin a varity of routs to spiationhas
nativs.Why, thn, is not vry omlar. Laoratory primnts (rviwdin Ri nd Hostrt 1993)
: d d i f f r n ts p i s ? h r u o f hav onfirmd that ologial spiation an our; primntal studis
w muh gn flo,ours twn that raisd fruit flis (D. melnogstr or D. psudoobsur) in diffrnt n-
:s to homognizpopulations oth vironmnts(varying tmpratursor food sours) hav dmonstratd that
lorks against sltion and drift in som lvl of prmating rprodutiv isolation volvd twn lins raisd
;iation, rsptivly.Sond' how in diffrntnvironmnts,whil no prmating isolation volvd twn in-
is thornir and points to th im- dpndntlins raisd in th sam nvironmnt. Ths primnts dmon-
iatiopross.Although divrg- strathow th pross might also work in natur.
nsi barrirsto rprodution, som In th simplst modls of ologial spiation, rprodutiv isolation
prsnt to stop gn flow and volvstwn populations inidntally as a y-produt of adaptation to
188 h ttr tld Prlssrlf Spitkl
-*/B/
\
finhs on Santa uz Is
aks adaptd fo rushi
((\/-
\7 I o w a t s o f s y l l l r
sis with smallr ak
aus song plays a ntra
this linkag twn ak
its onomitant influ
Sympatri pls of y-produt volut
portane in spition of
PARALLL sP|l
oF oLoG|AL l
ParallIvolution of sim
ronmnts strongly impli
s p i a t i o n n r s u I tf o n
produtiv isolation that
4. lassi modls (lr rods) tlf spiatitln r asd on th dgr of
I-.igr-rr rlatd populations as a t
gographi spration s t driving fo of spiation. In viarian alloptry,
nstrl ppulations of a on-ontinuous, larg rang of a spis r
sparatd y virirr vnt (.g.,mountains, riv, oan). Diffrns in
nvironmntal onditions on ithr sid of th arrir to disprsl pronrot loal
adaptation and y-prldut rprtldutiv istlliltion, 1adingto th formtion lf would rapid in suh fou
nw spis (ayr 1942). Th prihral-isolats or foundr-fft modl of ould rsult in gntiror
spiatiln p()stultsgnti drift as th prirry mhnism that prrmots i s o l a t i o na n d r s u l ri n r a p i
divrgn n1spition. Priphral isolats or foundr vnts rlur whrt prrpulationsthat ovrlprn
pirrrd Outward from th dgs of thir distriutions to OuPy
lpr.rlatilr.rs spiationwhr rprodu
nw l-raitats,rr lvhn ir s.ral]numr of ir-rdividLrlsdisprs to nW loatior-r not geographiallysparat
nd stlish a nw population. ayr (1954) prtlplsithat gnti hang aompanid y assortativ
Th ttrld Prlcssof Spition 189
a->\ 9
o
(Shlutr and Nagl 1995
from symptri spiation
t\*.***
4- te^ ulr nvironmnts have l
2l
.**,*-..'
gn in daptiv traits ;
uildup f assortativ m
ns in thos adaptivtra
h tlrrspin stiklbt
systm for th study of p;
9
o ak is a marin fish fol
4- Alaska and British Columl
2r_ **#& #, This spis is anadromot
t h j u v n i l S t g i n m a r i
6a adults tl brd. Indpnd
=---t
'-
4|_
2L-
*
4' ** &** to multipl stram-rsid
th oan. In addition,in
nizations y marin stikl
ir ttt&L b th rrt of th glairs
-t
2L pndnt volution of lak
r I a k s .S p i a t i o ni n s t i
LLL
nw envionmnts and th
;F
-a\
L ody-siz diffnsarrd
+
prfrns for mal ody
8
%-*.
6l-
4-
kHz 2l
0.5s
t"
.Tfl
oN IN Hos
'toN
divrgnwhr omplt gogrphi isolation is not lways rquird to ini- prfrnsfor onspi
tiat spiation. of D rosolt h iltt psendolts
Although phytophgous insts offr svral wll-doumntd ampls of lutlne to mat with ht
sympatri spiation though host shifts, a rnt study provids at last on popultions (Noor 199.5
mpl of th potntial for sympati spiation via a host shift in a vrt. frtil, allowing flr gn
rat' Afrian indigoirds (gnus idu) ar host-spifirood parasits, on of th st-dotr
laying thir ggs in th nsts of firfinhs (gnus Lgonostit)' and diffrnt t h i a n d l i m n t i o t y p
indigobird spis spializ in pzrrasitizingdiffrnt spis of firfinhs.In- ak. oward Rr"rrrdl:
digoird nstlings ar rard along with th host young and mimi th mouth lor21toryrosss anl ir
markings of thir rsptiv hosts, thus lowring th proaility of ing r- rids suffr a foraging d
jtdy thir host fostr parnts.In addition to this uniqu mimiry adapt- prntal nvironmnts.It
tion y young indigobids, as adults, mal indigoirds mimi th song that is vironnrnt with rmpl
sung y thir host spissir, and fmlsrard by a spili host spisof parrrtal spis.Horvv
firfinh prfrntially mat with mal indigoirds that sing thir natl host- parr-rtlnvironmnt, lit
spifi song. This failitats host-assortativ mating and lads to rprodu. rats :rnd wr smallr at
tiv isolation among diffrnt spis of indigoirds that parasitiz diffrnt Wr poor at thing
host spis of firfirrhs.A rnt phylognti study y ihal Sornson fild nvironmnts iru
and ollagus(Sorensont al. 2003) that ompard n-litohondrialDNA s- In adlitiol-l' nthi f
quns for a numr of diffrnt spisof indigoirds and thir rsptiv likly to mat witlr irrtrr
firfinh hosts dnronstrtd that diffrnt indigoird spis that parasitiz mals from al|opatri p
diffrnt firfinh hosts ar gntially diffrntiatd from on anothr, a sults support a modl of
finding onsistntwith assortativmating of indigoirds rard y a partiu- t h r o u g l l n t r l r ls | t i o
lar firfinh host spis.Thse rsults suggstthat divrgnthrough loal fitnss in ithr parntlll t
adaptations in nstling mouth markings, adult nrl song opying, and f- How impotant h.rs r
mal prfrns for host spis song typ has ld to positiv assortativ h p t l | mi s t h l t r i n
mating twn diffrnt host-spifi spis of indigoirds. This pattrn of pattrns of strongr div
host-spifi adaptations linkd with positiv assortativ mating within a tions. Imagin that two s]
host typ is onsistnt with sympatri spiation irr Afrian indigoirds gr of prldutivisrll
through host swiths in th firfinh spieswhos nsts thy parsitiz. I i k l v o r l t i n u t ( ) ( ) i s
ls of produtiv isolat
or th tintion oi th
RPRoDU|V lsolAloN MAY B RE|NFoRD
spis pairs) that w ol
Y NAURAL sL|oN
volvd a high dgrof
Natural sltionan at to dirtly inasth mount of prmating isola- following sondrv o
tion btwn two divrging populations by slting against hyrids formd v o l r ' di n m a t i l l g l i s i o
twn thm. This pross' first proposd Thodosius Dobzhansky drrrrinsth prssof rt
(197) and trmd riforcemelby Frank Blair (1955 ), an our in arasof romot divrgnirr rn
sondary ontat whn prviously allopatri populations om into ontat must tightlv linkd wi
and intrrd. Whn hyrids twn formrl allopatri populations ar ity. Tlrs quirmnts
partially infrtil, rinformnt n omplt th spiation pross. Rin- Ths argumnts do n
formnt was popular onpt ftr th modrn synthsis aus it allows plting th spiatilnpr
natural sltion to driv spiatin dirtly, ut its mpirial dmonstration that th vidn for tn '
has n diffiult, and its istn as an important omponnt of ologial siv. of th two pross
spiationhas n ontrovrsil(.g',Noor 1999; arshall t al. 2002). rr'ith ologiel s1rgi1
Most of th vidn for rinformntoms flm omparisons twn rinfo:tnt-th first i
allopatri and sympatri popultions in th strngth of fnral mating spiation; th irnpotn
h Pttrn d ross o| Spcitit 1 9 . t
lation is not always rquird to ini_ prfrnsfor olspifis.For irmpl, fmals drivd from populations
of Droslphil 1lseudotlbst,lrasvrrrpatriwith D . persimilis lisplay grat r-
',rlwll-doumntdxampls of lutanto nlat with htrospifimls than do fmals 1rorn aIlopati
rnt study provids at last on populations(Nolr 1995)' F1 hyrid mals ar stil,ut hyrid fmals ar
iation via a host shift in a vft- frtil,allowing flr gn hang and sltionagainst hyrids.
r host-spifirood parasits, on of th st-doumntdirmplsof rinformntours in th n-
(genusL gostit), and diffrnt thi and limnti OtYpsfound in lak populations of th thspinstikl_
3 diffrntspis of firfinhs. In. b a k . H o w r d R u n d l a n d D o l p h S h l u t r ( 1 9 9 8 ) d m o n s t r t d o t h i n
rhost young and nrimi th mouth laoratoryrosss nd in F1 lrr.bridsdttd in wild populations that hy-
,wringth proaility of ing r- rids suffr a foraging disadvntagrlativ to thir parntal spis in th
:ion to this uniq mimiry adapta- parntalnvirlnmnts.In th laoratory F1 hybrids rard in a ommon n-
indigoirdsmimi th song that is vionmntwith ar-rrpl assto food showd growth rats similar to thos of
; rard by a spifi host spies of pantalspis.Howvr, in fild studis F1 h,vidsrard in itlrtyp of
ligobirds that sing thir natal host. parntalnvirlnmnt,littoral :rnd limnti zons of laks' had lowr growth
tiv mating and lads to rprodu- ratsand wr smller at maturity than ithr pur parntal spies.Hyrids
ndigobirds that parasitiz diffrnt Wrpoorr at athing nthi invrtratsand plankton in th diffrnt
gntistudy y ihal Sornson fild nvironmtsausof intmdiatgill-rakr numr and ody siz.
.onrpardmitohondril DNA s- In dditiln, rrtlrifmls from svmpatri poptrltions wr rrruh lss
o f i n d i g o b i r d sa n d t h i r s p t i v likly to mt rvith intrmdi:rt-sizd F1 hyrid mls than rvr nthi f-
t i n d i g o i r ds p i st a t p a r a s i t i z mals from allopatri populatilns whr limntis did not our. Ths r-
liffrntiatd from on anothr, a sultssupport a modl of rinformntof siz-assortativmating prfrns
o f i n d i g i r d sr a r d y p t i u - through natural sltilnagainst intr-rdiat-sizd hyrids that hav lower
ggstthat divrgn through loal fitnssin ithr prntl environmnt.
adult mal song opying, and f- How irrrpotanthas rinf<lrtnnt n in ss of ologil spiation?
yp hs ld t positiv assortativ Th prolm is that rinforrnntis not th orrly plnation for osrvd
:isof indigoirds.This pattrn of pattrnsof strongr divrgn in mting prfrnsin sympatri popula-
'sitiv assortativmating within a tions.Imaginthat two sympatri populations had alrady volvd a high d-
spiation in Afrian indigoirds gof rprodutivisolation wlril still in allopatrv. Ths populations will
:is.hsnstsrhy prsiiz. likl,vontinu tO Oist,whil formrlv alloptri populations with low lv-
ls of rprodutiv isolation n121y lost ithr tlrr<rugh gn Ilow and fusion
or th tinti<lnof th rarr population. Tlrus th only poprrlatins (or
r R|NFoRED
spispairs) that w lsrvliving in sympatry in natur ar thos.that
voivda high lgrof isolation in allopatry, not usof rinformnt
as th amount of prmating isola- follwing sondv ontat. In addition, ronrir-rationtwn loi in-
ly sltingagainst hybrids formd volvdin mrtingdisions rrd th hratr(s)ir.rfluning hyrid fitnssurr-
'osd y Thodosis Dobzhansky drminsth prss of rinformnt.In ordr f rinformntto proprly
k Blair (1955),an our in aras of promotdivrgnin rntpr[rns,gnsthat od for mating disions
rtri popultionsom into ontat must tightlv linkd with gns that od for hyrid infrtility or inviail.
rmrly allopatri populations ar ity. Ths rquimntsmay not oftn mt in ntural populatiolls.
rpltth spiatin pross. Rin- Thsargumntsdo not shw that rinformntl.rsnot oprtd irr om_
: modrn synthsisbaus it allows pltingth spiti<-ln pross during ologial spiation.Tlry only suggst
tly, ut its mpirial dmonstration that th vidrrfor th widsglradimportan <lfrinformnr is inonlu-
important omponnt of ologial siv.of th twl prosssthat an promot rprodutivisolation assoiatd
> o r1 9 9 9 ; a r s h a l l e t a | . 2 0 2 ) . with logial spiation-divgn with isoltion as a y_produt nd
t oms from omparisons twn informnt-thfirst is wll doumntdnd almost rtainly irnportant in
n th strngth of fmal mating spiatiorr; tlr importan lf th sond still rrdsfurthr strst:rrrtiation.
196 The Pttern nd Prrl.,sslf SDeitil
akgrlund radian i
sxUAL sLE|oN AND oLoG|AL sPlA|oN
ovrall rightnss of tJ
Sxual sltion' a for that ats on traits that inrasmting suss'an m n t h i r v v t l l v d r i
inflund y th nvironmnt and' as suh' miry ontriut to ologial nd tallsmissiun t |
spiation.Sxul sltionthat lads to spitionirs a rsult of divrgnts- hitats'whr th a
ltion twnnvironmnts iln rsuit fronr sptial viation in sonday i n d w l . r ps i g n a l sw l l u
sxual traits, suh as linal variation in mal plumag or oloration in r- wirlr tlrWrongdw|.r
spons to a vaying nvironmntal gradint. For ampl, linal variation in and hrrlss likly t
th availaility of foods high in arotnoids that ld to right rd and or-
ang oloration in mal irds has n suggstdas th primary aus for di-
H RoL oF s
vrgn in plumag among popultions in mal lrous finhs, rpodus
NoNoLoG|
mius (Hill 1994). A rr-ro{lthat dmonstratsth importan of praill
linal variation in fml mting prfrnsas a sult of nvironmntal o d l s t r f s u a Is |
variation in mal signling traits was first proposd y Russll Land (1982), mating prfrnsfor
ut good ampls of linal variation that lads to spiation via sxual s- u [ r i o n s n d i v g l l
ltion ar laking. and spiation. Suprf
Natural sltion through variation in fatursof th nvionmnt whr s p i e t i l l l . p t t h i t
mating signals ar propagtdn sult in divrgntsxual sltionof mal m l n l t i n g ,s i g n l r l sr h
mating signls' suh as ourtship displays or nuptial oloration' lading to p r o p r i st o t h s i g n l I
prmating isolation twn populations from divrgnt signaling nviron- p r o m < l d h r t ' u g hd i v t
mnts (Ryan and Rand 199; Boughman 22).Natural sltion an also on modl of fma
at upon th snsory systm of fralsas wll, lding to snsory iassfor runWa pross (or ..
rtain proprtisof ml signals (.g',brightr oloration, largr siz, loudr F i s h r i n p o s s sa r
and longr durtion of volizations). Partiular faturs of lal nviron- tlr tttlstxrm rtr.tl
mntal onditions ould altr sensory ias or mk som mal traits ttrat p r J i s o r t h i g h r
ploiting biass in rtain nvironmnts than in othrs. In addition, fmal tion of ths traits ris
mating prfrns for mal traits that ar indiators of suprior quality an that nod th attrat
also inflund y th signaling nvironmnt' For xampl, if partiular fo that teit. Fisr.s
ml traits ar mor rlil indiatos of quality in som nvironmntsthan a n d t h i s u n d a n tt
in othrs (Shlutrand Pri I99), divrgnirr prfrnsbtwn nvi- promoting th volr.rti
ronmnts for thsmal traits ould lad to prmating rprodutive isolation 1994); howvr, studi
twn thm. Thus diffrnt ovolutionary paths of signals nd prfrns ult to pinpoint. On
in diffrnt nvironmnts ould ]nratspiatiorl. span length in stalk-
Th st xampls of an importirnt rol for sual sltionin prolnoting Grald Wilkinson and
spiation as rsult of varying signaling nvironrIrtsom from spis wild flis' a slrort-y-
that liv in environmnts that vary in light irrtnsiryor transmission propr- o s l i o n .t h y f o L r n
tis that influnth sptral quality of signals (i..'whih olors irr st r- a I l y s I t dm a | t r e i t
fltd y partiular nvironmnts).For xampl, allopatri populations of from th lin sltdf
Aolis cristtlls lizards our in two diffrnt nvironmnts (ri and with short y spans' .
msi) that diffr with rspt to light intnsity and th sptral quality of prfrr1mals with l
olors tht r st fltd.nul Ll and Lo Flishman (2004) hav r n s i o r - s p a nl t
shown tht mal anols frorrr th two nvironrnts diffr in th olor and s t a l k - y df l i s t h e t d i
pattrn of thir dwlaps (folds of skin undr th throat that ar distnddin S x u l s l r i nm l
mal displays usd aS tfritorial nd mating signals)in th dirtion that in- g s \ .U n d t h i s m o d
rasssignl dttailityin ah haitat. als fom th xri nvironmnt fmal lvith informatil
hav darkr and slightly rddr dwlaps that miriz ontrst whr th mk mating dision
Th ttr d Prrlssrlf S1litiltt 1 9 7
suprior quality to hr offspring. Although this mhanism may potntially Sxual sltion has t
lad to spiation if th indiator traits diffr twnpopulations, lik run- part aus of its dir
away sltion,mpirial xampls ar rar. Som vidn indiats that a 2 0 0 1 ) . S p i so f i r d s r
good.gns mhanism may promot diffrns in ml gnitalia that lad thought to xprin st
to divrgn in som inst taa, although this hypothsis is still undr d- monogamous linags;tJ
t (oskn and Stokle 2004).It should notd that for oth indiatr spios (Mitra et a|. 199
and Fishrian traits' th lak of onlusiv mpirial vidndos not mn of high sPitioI1 r a t si
that divrgnis unlikly to a produt of ths mhanisms;rathr, it is prisonsof dihromri
oftn diffiult to distinguish whih mhanism may hv oprtd in th past. with fmals dull) lads
An additional rol of sual sltionin spiationan trad to undr- usually right) lads (l
standing th mhanisms assoiatd with sual onflit. Sual onflit amountS of fthrornar
ariss whn th st stratgy for ln sx is diffrnt (or vn dtrimntal) hav n osrvd in an
ompard with th st strtgy in th othr s. This antagonism is pr- and othr invrtrats
ditd to lad to a ovolutionary arms ra whr diffring sts of traits - t a l . 2 0 0 1 ) . A r n t x
twn th ss intrat to dtrmin th natur of mal-fmal intrations rsult f sual sltio
(Arnqvist and Row 2005). An ampl of within-spissual onflit is Lupl. Tamr Mndl
th ffts of sminl fluid in Drosopbil melnogstr. Sminal fluid has spis at molular mar
n shown to nfiial to mal fitnss' rduing fmal rmating and in- tiv ags of th diffrnt
rasing ovulation, ut it is toi to fmals; th mor sminal fluid thy r- i n o n l d o f t h s r i
iv, th fastr thy di. Sual onflit apparsto our aross a wid rang spis pr million yars
of taxa (Arnqvist and Row 2005), ut prditaly,som of th st vidn iation in this group app
for a o1of sual onflit in spiation oms from insts (Arnqvist t al. song trill rat, whih f
2000). Baus sual onflit may lad to rapid volutin in th rprodu- appar to any olog
tiv trat, it follows that it may also inrasth pa of rprodutiv isola- rikt spis.Ths a
tion twnpopulations. tion an an importn
Prmating isolating bar
sXUAL sL|oN' lsoLA|oN'
AND RAs oF sPlAloN H RoL oF Ml
Sual sltion,ausof its rol oth in promoting th volution of pr. Muttional hangsass
mating rprodutiv isolating arrirs and in ontinud divrgn twn thought to a primar
spis,may hav onsidra1impat on th rat of spiation (rviwd rWhit (7978, 336) onl
by Panhuis t al. 2001). For xampl, taa with a wid rang of snsory pr- th primary rol in th
ption apailitis may b partiularly pron to spiation through sxual widsprad rol of kay
sltion, sin thy hav n lvatd aility to produ or dtt a road aus th aumulation
rang f signals, thy visul' aousti, or phromonal (Andrsson 1994). t h t t u g h t t h l a r g , l i r
Spis-rih groups f frogs' for instan, an dtt a widr rang of problm with th viw
advrtismnt-all frqunis than thos in lss spios linags, potntially tion ntrs on th diffi
aounting for why suh linags of frogs ar mor spios ompard with lations rhing fiatto
thos with narrowr frquny prption rangs (Ryan 1986). In addition, ltrnativ rarrangm
rinformnt,as dsridarlir, an omplt th spiationpross upon rids, thry rduing
sondary ontat y nhaning divrgn of fmal mating prfrns. mosoml rarrangm
Sxual sltionplays an important rol in sympatri spiation as wll. As- trm hromosomal r
sortativ matin8' th propnsity to mat with lik individuals, hs n pro- in hangs in lrromos
posd s on way to quikly volv rprodutiv isolation (lyn and orr hromosrmal rarrang
2004) and is a nssaryomponnt of th sympatri spiation pross. ar infftiv rrirs1
Tll ttrn d Process of Spitio 199
gh this mhanisnrmay potntially Sul sltiorrhas n linkd tl rlativly higlr rts of spiation, in
.ffrtwnpopulations, lik run- part aus of its dirt fft on mat rognition traits (Panlruis t al.
rr. Som vidn indiats that a 200l). Spis of irds with highr lvls of promisuity, for ampl, ar
frnsin mal gnitlia that lad thouglrt to prin strongr sual s.ltionthan losly rlatd, mor
gh this hypothsisis still undr d- moogamous linags;ths promisuolls linags gnrally ar also mor
rld notd that for oth indiator spios(itra et al'.1'996).othr studis of irds oform to th prdition
mpirialvidndos not man of high spiation rats in taa with strong sxual sltion,inluding om-
;t of thsmhanisms:rathr. it is parisons of dilrromati (ss of diffrnt olors; nrals gnrally right,
nisrrrmay hav opratd in tlr past. with fmalsdLrll)lads vrsus monohromati (otlr ss th sam olor'
r spitionan trad to undr- usually right) lads (Brralough t l. 1995) and lads with vaying
th sr'rlonflit. Sual onflit amountsof ftrornamntation (ollr and urvo 1'998).Similar trnds
is diffrnt(or vn dtrimntLrl) hav n osrr,din angiosprnrs(nd thir assoiatdpollinatos),insts
o r h r s . T h i s . r n t g o n i s mi s p r - nd othr invrtrats,arrd fishs (rviwd in Andrsson |9941 Pnhuis
l wlrr diffring sts of traits . t a l . 2 0 0 1 ) . A r n t x a m p l h i g h l i g h t i n gt h r a p i d r a t o f s p i a t i o na s
n a t u r o f m l - f m l i n t r a t i o n s rsrrltof sual sltion hs bn shown in Hawaiin rikts in th gnus
of within.spissxual onflit is Lupl. Tamra Jv{ndlsonand Krry Shaw (200.5)usd divrgnamong
l mllgsler.Sminal fluid has spisat rnlularmarkrs liratd with gologil stimtsof th rla-
s, rduingfmal rmating and in- tiv agsof th diffrnt Hawaiian islands to show tht th rat of spiation
rls;th mor sminal fluid thy r- in on ld of thsrikts from singl islnd was alulatd at ovr four
p p r st o O u ra r o s sa w i d r a n g spispr million yars,th highst rat vr alultd for arthropods. Sp-
'ditaly, som of th st vidn
iatio i tlris group appars t<lb promotd y diffrnsin ml outship
orsfrom insts (Arnqvist t al. songtrill rt, whih fmals us in spisrognitin, austhr do not
to rapid volution in th rprodu- apparto arry ologially distinguislringfaturs that spart diffrnt
astI]pa of rpodutiv isola- rikt spis.Ths and many othr ampls highlight how sual sl-
tion rr n irrrportntvolutionary flrtht lads to raid divrgnin
prmatingisllirtingbarrirs' rsulting irr high rats of spiation.
{,
HE RoL oF MUA|oN lN sP|A|oN
Lorn Risrg and ollagus hav studid rominational spiation in ontirrs diploid sp
sunflowrs (Helitbus spp.) and th ffts of hromosomal rarrangmnts spis Hl uersiolor
in ontiuting to th spiation pross.Rominational spiation ours lopilty (H. hrsoselis
whn two diploid spis intrbrd and giv ris to a nw linag that is northl.l lralf), ut thy
both frtil and tru rding br.rt is rprodutivly isolatd from both tlrir rarrgsovrlap. Di1
parntal spis.Risrg (2001) argus that hromosomal rarrangmnts produtivly isoltd frt
that ar htrozygous in hyrids hav majr ffts on supprssing romi. nl diffrns) and p
nation twn hromosoms that ay th rarrangd sgmntsand gns intrrding. als pl
linkd to ths sgmntson th hromosoms. Gns unlinkd ro th ho- rats, with th dil<lidh
mosomal rarrangmnt sgmnts,spially those that ar nutral' frly in- r i l t 5t h a n t h o s t l f t h t
trogrss twn the hyridizing parnt,al spis. Rarrangmnts that l Grhardt arrd oll
supprss rombination ut lak a ausal fft on hyrid fitnss ould at ttraploids ar highly s
synrgistially with linkd isolation gns to xtnd thir ffts ovr a largr valr'latingall hart
gnomi rgion. Thus hromosoml rarrallgmntsproaly do play an im. n a t i n g g a i n s t a l l s r v i t h
portant rol in rduing gn flow aross spis rrirs, ut not Irssarily A n ts u l vl l y A I i
through th mhanisms of rdud hybrid fitnss traditionally nvisiond. dmonstratd an unusu
Th hyrid sunflower spis H' omlus, whih rsults from hyridization spis H. ursir>kt.Us
twn H. us and I.I' petiolris, is thought to hav arisn in suh mitohondrial gn rgr
mannr. In ddition, H. omlus has highr fitnss in novl nvironmnts this study showd thirt
whr oth of its parntal spis pognitors do poorly. Suh n adaptiv r i u dt o m t r l r i p l o r i g ,
rol for hyridization in fostring spiation has bn suggstdfor a numr 8). Two of th thr dip
of plant and som animal spis (Arnold 19971Dow|tng and Secot 1997). suggstd y th ourr
not in th iiving dipl
sPlAloN Y PoLYPLolDY t t r l l l i d sa r o \ t h r ( ) u
diffrnt dipllid nsto
Polyploidization ours through th dupliation of ntir sts of hromo- rls from diffrnt dip
soms and is on of th fw mhanismsthat an produ instantanoussp- symptry. Busth dl
iation (polyploid spisar rprodutivly inompatil with thir diploid latd from on antlthr,,
prognitors). Polyploid spis ar formd ithr trough gnom duplia- u r s i tl l t , r . P l p l t ' i di z . r t
tion of a singl diploid spis(autopolyploidy)or through th fusion of two sisntharrgin th dv
or mor diploid gnoms as a rsult of intrspifi hyridization (allopoly- ll siz, whih hs n
ploidy). Both mhanisms of plyploidy ar ommon in frns' mosss,alga, ially prodd utotrip
and virtually all groups of vasula plants, partiularly angiosprms (St- hngs in trill rat s a r
ins 1950;Lvin 2002). til.ltlit twn lirrgs
How polyploid spisonrstlishdis not fully undrstood.Svral rinftlrd tri Il-reltdiffl
onditions, partiulaly ommon in plants, might nal a nw polyploid to gry tr frogs illustrat
inras and form a vial population. Ths inlud slf.frtilization' vg. ais nd furth shows
tativ propagation' highr fitnss than th diploid prognitor spis, or mnt lls, an lirt
nih sparation from th diploid prognitor spis'Indd,many polyploid
tax rprodu y slfing or vgtultivpropagation, nd most diffr from
H RoL oF G
thir diploid prognitors in haitat and distriution. lnrassin ploidy altr
ell siz, watr ontnt rat' rat of dvlopmnt, and many othr physiolog- Irr thory, gnri drift
ial proprtis' so many polyploids may hav sltivadvantagSin nw outward from th dg
ologil nihs imnrdiatlyupon thir origin. whn a small nulrlr tl]
An nusual as of polyploid spiation in animals ours in th gay rr nw popr-rizrtilr-r.
Irrpr:
frog spisomplx found throughout th astrn Unitd Stats.Th omplx fomatirn rf rrw spt
Th Pttr nd Procss of Spition 201
diploidnstorsillustratsmLrltipl
|lursiolor.Th rlationshipsof nsqunsof Spiation
H. uiuo,ar dpitdat th
' sp.A and H. sp. B' wr infrrd Th ulk of this ssayhas for"rsd on amplsof living spisand th veioLts
ay trfrlgs,H. uersiolor (4N), mlranismsy lvlrih th ar thought tl lrav spit.l.Yt spiati<lrr has r.
loid arrstors(A x B, A x , arrd sultdin tlr normous divrsityof norv-tintanimirls,plants, irrld riros
ntlings(.g.,N, NW) ar unovred in tl-rfossil rrd as wll. Dtrmining pross frorrr pattrn is
spisthat sharsa divrsgn
hallnging;hn most studis tlf th spiation prtlss fous ln living m-
. (FromHollowy t al. 2006.)
pls.Although n-ruhdathas ntrdOn th mod llnd tmpo t>fspition
s illustratd y th fossil rord (phlri gduirlisnl, tlr sllrv trr-rsitiorr
of
on spisinto anothr' vrsuspuntuatdqr'riliriurrr' rapid volutionarydi-
vrsifiationand spiationfollowd y long priods of stasis),volutionar,vi-
' spcition(Figur4) was first ro- ologistsgnrallvagr that th sam volr'rtionarylhanismsthat prorot
ed priptri spitio) and was divrgr-retod1,nrong popultitlnsof living spis(nrirovt>lutiolr) r r_
rds and mmmals, populations p- sponsilflr siationvnts(mrovtllution),otlr lngoingar-rdin th past.
..parnt''
spis wr oftn highly Th tmpo of spiation vais dramiltially among diffrnt taa. Som
rs diffntspis.N4ayr proposd groups'suh as hrsshoras (gnus l-imulus), hav rmind unhangd
suhlolizdfoundir-rg populations for ovr l 50 million vars, w,hil othrs' sr'ls Hr'r.iianrikts (gnus
Te Pttrn d Pross of Spitio
Lupl), hav spiatd rapidly, with up to 4.17 spis pr million yars )iz, A., and M. R. nrr'
(ndlson and Shaw 2005). Thus spiation an our quikly or slowly, p r z r . g tilt r p r o d u r i v i
undr varity of ologil onditions, and with or without th volution of p . l g r l i t o r . g u t t t u s .i
Dozhnsk,v,T, 1937. Gt
rproiutiv isolation.
LIr-rivsitPrss.
Unravling th mystris of how nw spis aris is on of th nrost xit- l 946. (io.rpltt'pr
ing ras of volutionar iology' and although w hav mad muh progrss spis lf Drostphil.
in undrstanding spiation sin William Batson's arly onrns, futur Dowlir'rg,. ., d . l,. S
studis promis to yild vn mor nw and fasinating disovris. Th hop in th divrsifi:rtiol.ttlf :
28 59_61'9.
of th futur, in prsrving and protting our plat's iodivrsity, lis in an
F d , . | .L , | 9 9 8 . T h a p p l el
r.rndrstanding of th pross of speiation and ommitmnt to saving th Onvntionalwisdom a
produts of this pross: th spis that ist on rth today and thos that ds., lld/ess Flrnts:S
hv yt to voiv in th ftrtr-rr. Urrivrsit,vPrss.
Fishr,R. A. l930. Th ,t
Prss'
lBL|oGRAPHY F u t r t v t ,D . J . , J . S . W l s h T
G n t i v l r i l r t r l litr l l t p l
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Arnold, . L. 1997. Nturl Hbridiztil d uoltil. Nw York: Oford F l ' l u Ii , l D , B i , ' l ' ' 1 ' :
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promots siation in insts. Prdigs of th Ntttilttlcdm lf Sis volution of ormutrt
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zski.and R. . Silrglied.1997. 1'dn,R. L. 1997. hirary of spisOnpts:Th dnoumntin th sag of
lxrnding H:rldn's ru|e. I'rldigs th spisprlll.In M. F. laidg, A. H. Dalvh, nd . R. Wilson. ds.'
164:845-8-51. Spis:Th Uits lf Birldiursit-, 3.s1_424.Noru.ll, : hapman and Hll.
206 h Pttr ttd Pross lf Spitio
ayr, . 1942. Sstmtis d the rigit of Spis.Nw York: olurra Sits' J. W.' J.., and . Nlrl
Univrsity Prss. rvisitd. S stmti Zt-
19.5.1. lrarrgot gnti nr,ironnalld volution. IrrJ. Hulv, Sokal, R. R., and . J. rov,
A. . Hrdy, nd . B. Ftrrd, ds.' ullutils Plss' 157-1B0. [-ondn: valuation. Alri
G o r g A l | l l. l l l d I j n w i n . Sorson, J' D., K. . Sf,
1'96.ill S1lis d t'olutil.arnridg,A: Blknp Pssof brood p rsiti indig'b
avrd Unir,rsiPrss. S p i r i t t - l. . 1 9 9 8 .T h t l l f
innon' J. s.' s. lri, B. K. Blkn.ran,L. Dvid, D. . ingsl1',L..|anris.ln, D. J. Howard nd S. H'
J' 1rou,arrd D. Shlutr' 2004' vidn ftl llt>g's rol in spii]tion. 32o-3z9.Oxford:Of.
Ntttr 429 294_298. Stins, G. L. 195. rit
ad, L. S., nd S. J. Anold. 2t)04. Quntitti\,gntirlls<lfstll sltin. Ul-rivsitPrss.
reds i rlllg1, d t'tllutitl 1'9:164_2-71. Sturtvant' A. H. 1920. Gr
.r.' Hyrids wtth Drosophl
N{ndlson, nd K. L. Shaw. 200.5.Rapil spiition in n irrthrood. Ntur
4:375-76. Tmplton, A. R. 1980. Tht
N{itr, S., . I-ndl'nd S. Prrrtt-Jorrs. 1996. Spisrihnssvlris rvith 94 1|1-1038.
mating systnl it ir.ds./,,uk 11:544_.5.'1. l989. h maning
N4ollr,A. P.' nd J..|. trrvo. 1998. Spiation nd flrthr<lrnrnnti.ttilrr irr irds. n d J . A . n d l r , d s . ,
uolutitlll 52: 859_t]69. A : S i n u rA s s o i t
Noor, M. A. F. 199.'. Spitilndrivn y nturl sltiot-t in Droslphil. Nfutr ing, .-T.' S. . Tsaur, .
3 7 5 : 6 7 4 - 6 75 . homoox at th sit O
1999. Rinformnt and Othr onsqunsf symatry. Hrdit U s h i r n a ,R . . n d . A a m i .
83: 503-508. Ntur 425 679.
Orr, . A. 199.'. Th populti<)ngntisf spition:Th vllutionof hyrid van Batnurg, F. H. D., n
impatiilities.Gctis 139: l 80.5_18 l 3' oiling: omputr pt
1996. Dozhnsky, Batsln,and tlr gntisof siatior-r. tis 114 van valn' L. 1976, ologi
1 . ] .1] _ l 3 5 . Whit, .J.D' 1978'od'
Panhuis, T. M., R. K. Butlin, NI. Zuk, nd T. Trgnza.2001' Srtalsltiorrnd ompany.
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spiation.rltds i oklgl d t,ltil 16 64_37 |. Wily, . o. 978. Th vo
Patrson' H. . H. 1985. h gnitionOnptof sis.ln . S. V:r' d., Zoolog 27:17-26.
.Wilkirrson,
Spis.rd Sci|tti,21-29.Transv:rIMusut onlgrph rlo. 4. Ptlria, G. S., nd P. R.
South fria: ransvaal -Nlusutlt. slion on irn g
odos, i. 2001. lrrltlvluti<lnof nlorphology nl voirl signill struturin Rol Soit' of Lodt
Drwi's finlrs.Ntur 409: lti.5_]88'
Ri, W. R. 1998. Intrgnornionflit' intrlous ntgonistior,olutiot"t, and th
l.llutionof rpodr-rtiv isoltitln.In D..J' Hlward nd S. H. Brlhr,os.,
dlss Flrms: S1lisd Spitio,261_270. Nw Ytlrk: ofrd Urrivsity
Prss.
Ri'.!(. R., and . . Hlstt.1993. I,aoratoryprints n spietion:!(/l-rat
hv rv lrlrdin 40 yrs? F|lltklrt47: 167_1653.
Risrg,L. H' 2001' hrrnrtlsmaIrrngmntsand spiatiOl1.Trds i
tfulgd tllutiltz16: .51_j.58.
Rundl. . D.' and D. Shlutr. 1998. Rinflrlentof stiklirkr.nat
prfrns:Syrnptrr.rlsnrmpt. t l l tttio .'2,:200_208.
Ryan, M. J. l986. Nufoanrtomyir.rf1uns spilrtinrilts :]nlOngxnurJns.
Prldigslf th Ntiol rlm tlf SiltsL/S B3: 179_182.
Ryan, M..|., nd A' S. Rnd. 1993. SpisrOgllitiond sul s|tinas
unitary prolrn in ar-rimalmm unitio. lltttitt 47 : 647 _6 57 .
Shlut,D.' nd l,. M. Nagl. l995. I)arlllspiritlrly natural slti<ln.
lritt Nturlist 146: 292_()|.
Shlutr' D.' and T. Pri. l99.j. Holrst-y,PrtiOn,anl p1lLrlirtiorr divrgnin
sxually sltd traits, Prot:digsrlf th Rln,l Sit lf Lldo, Sris B
2 - 5 3 :1 1 7 - I Z Z .
.I'lttlll1'.
Simpson, G. G. 1961 , Pritlipls f il Nw Yrk: lLtlnbi
Univrsitv Prss.
h ttt d Pr.>css
;f Spitiotl 207
.|ousntagolisti
ovllutill.t,
nd th
' J. wrd .rndS. . BrlIr,ds.,
61-270. Nr,vYlrk: Oford Univrsity
rmntof stiklakmat
tlrllutio 52: 200_208.
ipirionatsmon8 l1urns.
S i c sU S 8 : \ 7 9 _ | 8 2 .
ognitionnlsxual sltlns a
'n.L.uolution17: 647-657.
siation1'natural sltton.
ption'and oultilndivrgllin
Rl Sllit'tlf I'lttd, Sris lJ
|o||orIJ'.
Nrv York: olLtmia
i
I
Y
208
uolutio d DeueloDmt 209
ev0li"rtr0nary
mehisms
g
)vlopmnt t
i
is at th girrningalra1ya vrt-
se).
rt fr th nltitln of rapitulation
tiorr thory rahd its zrrith in tlr
,hosstatmntthat ..ontognyra-
tr nihts ist als in kurz Rpitu-
'dayrrapsulats fo rany iologists
Jvlopmnt.kl, strong su-
s an hiit of phylogntihistory,
qunduring tlr dvlopnrntof an
ogrssir. dir,grr of initilly simi_
lh, turtl,ird. and humn mryos
ury volutionardvlopmntal iol.
; H:rkl'svi'uvof rapitrtlation as
ltionaryhrrgrathr thn simply
rd this mharrismto onsist of two
sonto th n1of dvlopnrnt and
.mor rvitdarlir stags.
ptions to strit rapitulation
y Bar (1828). H alld thm
Figur2. Rapitrrltitln as nvisind1.Hakl. Th gratomparativ
in muh th sm light tat most
anatlt-t-tistrnst kl was tln of t strongstProPonntsof t notion that
s tht hav onvrgd in fom Ltt
thvoIutionr,v historyof 'r spisis rplavddrringits -rrylogial
nr-roying ofr.rstion of volution- -l
dvlomnt(Hakl tl66).His famos illustratinsof vrtratmryls,
nds of nognti features.. ht- onof whih is shown hr,lpitprgssivdivrgnin ovralisimilaritis
lopr-nntvnts' and htrltop' or amngspisas lvlopnllrtprods.Th mryos ar (from lft tl right)
t strutur.Both trms ilr now usd thosf a 6slr,arrrphiian. tr-rrtl, ird, pig, ow, rzrllit,and hurt-t. Not that
any hangin th timing o loation all tlr mryos lrvgill slits t an rl Stag(top ru) and tht th umn
Ly (Raff 1996; Klingnrg 199s). m r v o h s t i |( m i d d l r o t u , r i g h tt)l - r a t i s l t r \ o s t ( b o t t o m r l , r i g h t ) . I t
Wasth prslltlf thsplt>gntiaIlv nintftursin rrrlrryos, dspit
Llrnt'mor gnrl us:rgof th
tlrirsnirl dults,tha strgl}.i.rrssd Hkl.(Frlmkl 1866.)
rpitultr. tmporal hangs, as
ory ons'
ak rapitulation thoryt nor Ws
, dtsaor-rtapitulirtiorrlvr
modrn synthsiswas undr way; amollg ta' al1d of th lristory of spifi harirtrs.hs thr prtial
l mr'ltualapathv that aros twn appliationswr all first proposd during th nintnth ntury, ut for
aout this tim irnd lastd trntil th somrvhatdiffrrrtrasons thy rmairrimportant tody.
Although tlr influnof rapitr.r. N()t mn radrs of tlris olk lvilI nd to prsuadd that th pross
rrvologyontinud to play an im- of volution taks pla. Antiipatirrg a rathr diffrnt audin, Drwin
mrshaldall th vidn h ould fid to support th onpt of trnsmu-
tationi]mong,spiswhn lr wrot oll the rigi of Spis(1859). \Whr
.,spifi''
Bar,who rjtdth noti<lnof volr"rtion, spok of ..gnrl''and
ln, Phylogny, dvlopmntaltrits, Drwin intprtdsimilaritisamong mryos as om-
plling vidn of ommon dsnt' ..mrology is to m y far th
strongstsingl Iss of fats irr fvor of lrang of forms,'' lr l,vrot in a lt-
vlutionarylristory, thy nonth- tr to Asil Gra (Sptmr10, 1860) shortl afterth publirion of tl-rori-
llution,of phylgntiltionships gl (Burkhardtt al. 199, 350). mryologial similaritisrmin prsuasiv:
rt
21 2 uolutiolt nd Dculrlmttt I
i
traits not manifst in adult anatomy' suh s th prsnof gill slits and a and sns or$ans prsr
tail in hurrratlmr1'<ls, ar tangil vidni dsntw,ithmodifitin. anstor or aros indr
A solld pratial appliation of omparativ mryology is irl phylogr- simila mryologial
ti analysis.Darwin pointd out in th rigi tht ..ommunity in mryoni sin orrspondn
strutur rr,lstltnmtrnitvof dsrrt''(18.i9,449). Sysrmtisrshav n 1994). nrrologial
applying this onpt nthtrsiastiallyvr sin. Sval signifiant insights mologis' suh as th
into phylognti rltionslrips am aout through us of this approah' origin of vrtrtja'
suh as th dis<lvr},thatrnals r arthopods tlrr thrr rnllr"rsks as vind y th rr
(Darwin 1854) and th ralization tat asidians ar losly rltd to vrt- (Burk and Fduia
brats (Grstang 1928). In oth ass'adults r highly lrivdanatomiallY, an our (d Br 1:
ut larva provid lr phlognti links. Sirrrilaitisirr mrylsor lrr. rains on of th nros,
providd th first vidnfor phylogntirlationshipsamong mny Othr. In rnt yas th
lvis anatomilly distint taa and oftn rm:rindth only dirt vidn to th prssion of
r-rrrtilth :ldvnt of rr-rolulr data sts.Th nrrlsf mrry lads r sd sd <ln th osrv
on ths mryologial ritria, sul] as Spirlia (for th spirl gomtry of during th dvlopm
mryoni ll divisions found in som anirals)and rnonoots (for th singl highly divrgnt anat
.s7ith
otyldon tlrat norishs th mryo of som flowring plants). tlr olland 1999). For
widsprad ppliation of DNA squnsir-rphylogntis,lrrryologial xprss not onl th
traits play a lss important rol in idntifying lds today, ut thy an po- funtion (myosin, at
vid an iortant rlity hk whrr rrw rlatilnshipsar proptlsd. trol th transriptiln
h squnof mrytllt>gial vnts has lso n invokd s asis for srvation that dvl
polarizing haratr-stattransftlrmtion in fornral phylognti analyss pattrns forms th z
(Nlson \978). his ltrlg|ttiL. rit('ri(,tllis zrsdon ir litrl :rdingof I994)' lgs (Pnganib
Br's laws: that rlir vnts duing dvloprrrnti1r mor gnral ;rnd ar homologous in ar
thrforaros arlir during phylogntihistory than latr ons. Th utility viningly rquirs o
o i t h i s i t i o n h s n q u s t i o n do n f i r s t p r i n i p l s( d Q u i r t l z 1 9 8 4 ) , rs of th sam gn
n d i t l a r l y f a i l s i n r n a n y r a l - w o r l d a s s( M a 1 9 8 9 ; R a f f l 9 9 6 ) . o u t - suh as spurious simi
group omparistln is far mor widly usd today, nd th ontogntirit- son 2001). Rigorous
riln is intrstingpinrarily as :rn ampl of th psistnf raglirr-rlationist homollgous strlltur
thinking a nturY aftr Hkl. among dutrosom
Th third pratial appliation of omparativ mrytllogyis s ritrion and of ody sgmnt
lf homollgy alon ] spifihartrs.of th tlrrpplitions rrrntiond tl 1'997: Azhnov
lrr,this rmains y th far th most impotant tod (Hll 1994; Wilkins
2002). Rihard own originally proposd th trms homolog and log
in 184 with rfnto his lloti()not th arl-rtyp' gnrlizlldy or- FItohrony
ganizatill-lshard among spis rvithin a larg gottp (Palrhn 1994).
Own, who was ostnsily strongly opposd to th idir of voltltion, intro- N{any vnts during
dud th trms t() ontrst twtl two kirrds tlf rrtomil sinrilitv partiular pa a
among spis:homology for orrsponlingprts of atl arhtypand nlrl- vnts; vn dvlop
ogy for nonhomologous traits with sirilrfuntions (Own 1848). Btlth trms in a onsistnt squ
wr soon propri:rtdy r,olutitlnarl'illogists(PnhrrI994) ltndar still vn largly indpn
trsdto distinguistwtwo funlrmntallv diffrntlristorialhypothss: most ilny hang in
ommon vrsusindpndntvtllutionaryrigins of a trait (Hall 1994). of othr aspts (Ra
Distingishing twn homllogr.rsand rralogous struturs is <lftn hang in th tinring
straightforward,ut in mll1,ssit is dinglydifliLrlt.A psistnthal- ld to th viw that
lng' for instan,is distinguishing,hthrth appndags,odv sgmnts' htrohroni (Goul
uoltitl td D uellpmet 2 1
l s th prsnof gill slits and and snsorgans prsnt in diffrnt animal phyla ar drivd 1rom ommon
t tlf dsntwith rodifiation. anstoror aros indpndntly (lor and lr)ilmr1997; Nilso22\. t
rItivmryologyis in pylogn- similar mryologial origin has long n onsidrd ritrion of lromology'
igi tat..ommunity in mryoni sin orrspondns in antomial rganization ar oftn larr (all
1859' 449). Systmatistsl.rav 1994).mryologial omprisons hav idrrtifidsom initially surprising ho-
r sitr.Svral signifiant insights mologis,suh as th oigin of nrmrrralianar ossils from jawbons nd th
lut through us of tl-risapproah, origin of vrtratjaws from gill arhs.This approah ontinus to usful'
arthropods rathr than mollusks as vind by th rnt dat aout digit homologis among arhosaurs
idiansr losly rltd to vrt_ (Burkand Fdui 1997;Wagner arrd Gauthier 1999). Although xptions
Lltsr highl drivd anatomially, an our (d Br 1971; Roth 1984), similarity of mbyologial origin r.
s. Similaritisin mryos o lrva mains on of th rnost usful ritria of homology in anatomy'
rltionslripsmong many othr_ In rntyars th mryologial rition of lrorologyhas n tndd
rmaindth onlv dirt vidn to th prssion of dvlopmntl rgulatory gns. This appliaion is
h namsof many lads ar asd basdon th osrvtion that th sam rgulatory protin is oftn prsnt
ipiaiia (for th spirl g<lmtylf duringth dvlopmntof horrrologousstrutursvn whn thy give ris to
imals)and monoOts(for th singl highly divrgnt anatomis and vn in distntly rl:rtdtaa (Holland and
som flowring plants). With th Hollnd \999). For instan, musl lls throughout th animl kingdom
s in phylogntis,mryologial xprss rrot only th strutural protins that ndow thm with tlrir spifi
,ingldsttldy, ut th alr pro- funtion (myosin,tin' nd r-tinin)ut also rgultory protins that on-
l rlationshipsar proposd. trol th transription of ths strutural gns (ylD and MF.2). Th o-
las also rrinvrlklas a asis ftlr srvation that dvlopmntal rgulat<lrygns ar prssd in similar
t in flrnll phr,logrrtianalyss pattnsforms th asis of ontrovrsial prposals that ys (Quiring t al.
lr is asd tlrr a lital rading of 1994),lgs (Panganianet a|. 1997\, and sgmntation(Holla nd et a|. 1997)
ivloplrrntar mor gnral and ar homologous in arthropods and hordats.Framing suh argumnts on-
history thrrlatr tlns.lr utility viningly rquirs nrpring orthologus gns (i.., orrsponding mm-
first prinipls (1Quiroz \984), brs of th sm gn family) and ritially assssingaltrnativ hypothss,
; s( M a l 9 8 9 ; R a f f 1 9 9 6 ) . o u t - suh as spurious similarity in gn prssion (Aouhif t,a|' 1997; David-
. d t d . r r . r l d t I t o n t ( ) g ( ' n t ir i - son 2001). Rigorous appliations of gn-prssionritri:r to idntifyirrg
lf tlrpsistn of rapitulationist homologous struturs inlud ompaisons of th ntal nrvolls systm
among dutrostomgroups (Williams and Holland 1998; Low t al. 200.3)
' r t i v m l l r v o l . l g iys s . t r i r r i r l n and of ody sgmntsamong divrgntgroups of artlrropods (Avrof nd Pa-
)f tlr thr rppliationsn-rntiond te|1997i Azhanov et a|. 1999\'
lportanttoday (Hll l994; \/ilkins
d th trms homolog and Iog
aht1,p, a grrralizdod1,lr- Htrohrony
n larg group (Panen 1994).
l s dt o t h i J . r< l f v t l l t l t i o n ,i n t r o - any vnts during dvllpnrnt happn in partiular squn and at a
,o kinds tlf antomil sirnilarity partiular p aus th a triggrd by, or dpndnt upon, arlir
ing prts of i]n arhtvp and anal- vnts;vn dvlopmntalvnts that ar not ausally rlatd oftn unfold
funtions(Own 1848). Btlth trrs in a onsistnt squn aus of gloal oordination or simply aus
ologists(Panlrn1994) and ar still vnlargly indprrdntprosss r h tighrly orrtrolid. As a rsult, al_
rtallydiffrnthistorial hypotlrss. most any hang in on asptof dvlopmentwill altr th rat or squn
originsof a trait (Hall 1994). of othr aspts (Raff 1996). Th rsult is a bterlhro'an volutionay
lnd arrllglusstruturs is lttn hangin tlr tinring of dvlopmntalprosss.This simpl onntion hs
:dinglydiffiult. A prsistnt hal- ld to th viw tht all volutionary hang in dvlopmntis fundamntally
-rrth ilppndags,ody
sgmnts, htrohroni(Gould 1977; Kinny and Namara 1991|'
21 1 d Dulopmt
F'ullutil
Mldrn idas aout volutionary hangs in th timing of dvlopmnt prsptiv that was soli
grW Out lf rapitr.rlationthory. Th aumulating wight of ptions and N4Namara 1991')'
stadily rodd th gnrality of akl's trminal addition and ondnsa- H t o h r o n yi n r l y
tion as volutionary mhanisms.Howvr, thr wr two additional fators fiation of dvlopmnt
that brought aout th dowrrfall of rapitulationthory: a lar undrstand- 1980s and 1990s. Dv
ing of th mhanismsthat undrli hrdity and th rognition that natural ryos and in omplx l
sltionn at troughout th lif yl (Gold 1977;kaff 1996; \Wilkins rohrony in th fossil rr
2002). By th 190s it was widly aptdthat th iogntilaw is nithr 1991)' Dvlopmntal
a mhanism of volutionary hang nor an aurat dsription of it. vn to suh pfosss as mor
th most voifrous ritis of rapitulatitlnthory rognizd,howvr,that p r i s o n s o f g n x p r s s
par:.rlllsfrquntly do ist twnvolutionry history and mryologial p o r l h a n g s .s o m i n
vnts. out of ths dats am nw and ultimatly mlr rwarding pr- Dikinson 1988; avn
s p tVl s. formations (.g., Wray l
On important insight Was that hangs in th timing of dvlopmntal lar that htrohrony
.!7l-
vnts lftn rprsnt adaptati()nsrathr than y-prldutsof hang. ould aris through a va
tr Garstng, a pugnaious and witty riti of rapitulation, assrtdthat A third important rns
..ontogny d<lsnlt rapitr-rlat phyitlgny, it rats it'' (Garstng 1922, rathr than pross (Raf
82). Garstang natly invrtd th aklian viw lf ontogny as a rord tims onsidrd to al
of phylti history ar-rdargud instad that ntural sltion ats on dvl- hang (Gould 1'9773
opmntal timing, thry produing hangsin anatomy. intrprtd original onption. To t
m a n y s g l i f i v o l u t i o n a r y h a n g si n a r l y d v l o p m n ta s a d a p t a t i o n s ,i n - traits, howvr, htro
l u d i n g i n t r p o l a t i o n s a n d l o s s s t l f l a r v a l s t a g s , h n g s i n m t h o d s o f annot aus hangs l
mtmorphosis, and altrations in th timing of rprodutin (Garstang nompasss a rih arr:
1 9 2 2 , 1 9 2 9 ) . S t p h nJ a y G o u l d t o o k u p t h i s t h m i n o t r l g n d h . mbros d Astors
loge 0977), rvhih tid htrohrony to th lassil ologial frm. sifiation of mods of h
work of r-sltion and K-sltion (r-sltionprdus adaptatilns'suh 1977; |eret a|.197
:rs rpid growth and arly rprodution' for suss in unprdital nvi- th varity of tmporal
ronmnts; K-sltionprodus th onvrsadaptationsin mor stal vidd a pris voaula
nivtlnmnts). any authlrs hav mphasizd th links twn ht- Htrohrony has fad
rohrony nd suh lif.history traits as fundity, timing of rprodution, opmntal volution, in
n d a d u l t o d y s i z ( . g . , K i r r n y a n d N m a r 1 9 9 1 ; R y a n a n d S m - pervasiv that doumnt
l i t s h 1 9 9 8 ) . T h i n p o r t a n tl s s o nt h a t m r g df r o m t h i s W O r k i s t h a t s i m - I/ilkins 2002). Th mp
pl hangs in dvllpmntal timing an prldu hirngs in organismal for volutionary hang
p h n o t y p t h a t a r o l o g i a l l yr l v a n t . hangs aris, nd how t
A sond, rlatd insight Was that volutionary hangs in dvlopmntal of phnotyp.
timing an ()urthroughout th lif yl. As arly as 189.5Frank Lilli sug-
gstd that adptatitlttsn aris in th arlist vnts of mryognsis.
(iarstang (1922) argud that nw traits ould appar at ny point in dvl- Dvlopmntal or
opmnt (not just th trminal addition advlatdy Hakl), and that su-
squnt modifiations ould shift thir tin-ringithr arlir r latr (not just A ntral gal of volutr
Hkl's ondnsation, involving progrssivshifts to an arlir position in ryp produ hngs in
ontogny).Dvlopmntal iologistsw losing intrstin volutionary pr- tntion on hrrri
sptivst th tim' howvr, and th fw volutionary iologists whtl wr idntifying whih ompl
studying htrorlnyfousd on lat lif-historystagsrathr than mbryos stalishing th numr
and larva. Most studis of htrohrony thrflr mphasizd hangs ifiations in traits. Ths
in dvlopmntal tiring rlativ to adult nat()myor to sxual maturity, a rn to th dvlopm
uolutio d Dulomet 215
ngSin th timing of dvloprrrnt prsptivthat was solidifid y influntial ooks (Gould 1977;cKinne
rumrrltingrvight tlf ptions nd Namara 1991).
's trminal addition and ondnsa- trohrony in arly dvlopmnt ttratd rnwd intst as th runi-
'r, thrWrtwo additional fators fiation of dvlopmntal and volutionary iology gathrd pa during th
trrltilnthory:a lirr undrstand- 1980s and 1990s' Dvlopmntal iologists amind htrohrony in m-
lty and th rognition that natural bryos and in ompl lif yls, whil palorrtologistsdoumntd ht-
(Gould1977;Raff 1996;Wilkins rohrony in th fossil rord (Raff and Wray 1989; Kinny and Namara
d tht th iogntilaw is nithr 1991).Dvlopmntal biologists garrto apply th onpt of htrohrony
an aLlrtdsription of it. vrr to srrhprossss mrrphognsis,ll linag,rrd gn prssion. om-
)n thoryrognizd,hwvr' that parisonsof gn prssionamong losly rlated spisrvaldmany tm-
utionar1'history nd mryologial poral hangs' som in th asn of ovious anatomil orrlats (.g.,
nd Llltinratlymrrrwardirglr_ Dikinson 1988; avner 1992) and otlrrsorrlatdwith phnotypi trans-
formtions (.g.,Wray and Bly 19941 tverot and Patl |997).It am
gsirr th timing of dvlopmntal iar that htrohrony, lik othr kinds of phnotypi transformations'
.Wal-
r thn v-;lrodutsof hang. ould aisthrough a varity of volutionary mhanisms.
i t i o t r a p i t u l a t i o n a, s s r t dt h a t A third important insight was shift to viwing hrrohrony s pattrn
n y , i t r a t si t , ' ( G a r s t a n g 1 9 2 2 , rathr than pross (kaff 1996). Until quit rntly htrohrony Was som-
lian virv of ontogny as a ord timsonsidrdto an volutionary mhanism that produs phnotypi
l1tnilturl sltioll ats on dvl- hang(Gould 1977; Namra 19B8), a urious prsistnof akl
angs irr anatom\.. intrprtd original onption. To th tnt that dvlopmntal prosss ar hrital
r l y d v I o p m nats a d a p r a i < l n isn' - traits, howvr, htrohrony is an aspt of phnotype lik any othr and
'val stags,hangs in mthot]s of annot aus hangs in phnotyp (Raff 1996). Htrohrony nonthlss
timing <lfrprodution (Garstang nompasssa rih arra of phnotypi transformations. Gavin d Br's
p this thm tn togen d Ph- mbros d ncestors (1940\ was th first attmpt at a lmprhnsiv las-
r to th lassial ologial fram- sifitionof mods of htrohrony, arrd svral othrs hav followd (Gould
.Wray
:ltiorrprtldusdaptations, suh 7977;|rch t al. 79791Raff and 1,989I.hs analyssmphsizd
, for sussin unprdital nvi- th varity of tmporal hngs that ar possil in dvlopmnt and pro-
nvs adaptatins in mor stabl vidda prisvoabulary for dsriingtheslrangs.
r p h s i z dt h l i n k s l l t w l r l r t - Htrohrony has fadd somwht as a ntral thm in studis of dvl_
fundity,timing of rprodution, opmntalvolution, in part austmporal lrngsin dvlpmnt ar so
l MNamar 1991;Ryan and Sm- prvasivthat doumnting anothr as is no longr surprising (Raff 1996
mrgdfrom this rvtlrk is that sim- Wilkins 2002i' Th mphasis has shiftd to undrstandingth gnti asis
an pro{uharrgsin organismal for volutionaryhangs in phnotyp, th ologial ontt in wlrih suh
hangsaris,and how th organization of dvlopmntiassth volution
lutionar1.hangsir-rdvllprnntal of phnotyp.
. A s l ya s l 8 9 . 5 F a n k L i l l i s u g -
arlist vnts ()f mrygnsis.
ould appar at ay point irr dvl. Dvlopmntalonstaint and Dissoiaility
dvoat] ,vHakl)' and tht sub-
ming ithr arlir or latr (not just A ntralgoal of volutionary iology is undrstndinghow hangs in gno-
s s i v s h i f t s o a r l r l i r p t l s i t i o n i n typprodu hangs in phnotyp. volutionary gntiists hav fousd at-
. l o s i n gi l l t r s itn r ' t l l u t i n a yp r - t n t i o n o n h a r t r i z i n gg n t i v a r i t i o n w i t h p n o t y p i i m p a t ,
:w volutionaryillogistswh<lwr idntifyingwhih omponnts of this vaiation sltionoprats upon, and
-historystagsrathr than mryos stalishingth numr and rlativ impat of muttions that undrli mod-
ony thrfor mphsizd hangs ifiationsin traits' Ths qustions hav n addrssdlargly without rf-
lt anrtlnr1. lr to sul maturity' a rnto th dvlopmntal rhnisrsthat link gnotyp to phnotyp
216 uolutio d Dulopmet
evolutionary redL
sin th shism twnvolutionary and dvlopmntaliology during th
1930s and 1940s (ayr |9821Wilkins 2002), volutionary and population
gntiistshav gnrallytratd dvlopnrntas a lak ox whos ontents
hav littl to ontriut toward undrstndingvolutionary prosss.
Th validity of this approalr has n hallngd rpatdly. harls
.Whitman, Frogs
an arly volutionary 8ntiist'wondrd whthr ..th laws of
dvlopmnt lrrdsom lins of [phnotypi]variation nd favor othrs''
( 1 9 | 9 , 1 1 ) . B y t h 1 9 8 0 s s v r a l v o l u t i o n a r y i o l o g i s t sW r p r p a r dt o n -
swr in th ffirmativ (ayr 1982; Holdr 1983; aynard Srith t al.
1985; Athur 1988; Alerh 1989I. Th notion that th orgnization of dvl-
opmnt imposs iasson th prrrdutionof phnotyps smsrasonal-
to liv othrwis is to assum that vry possll phnotypi outom is
qually likly, whih is manifstly not th s (Alrh 1989; \/ilkins2002). Salamanders
Th tru hallnghas n idntifying th undrlying soursand lng-trm
onsqunsof suh dvlopmntalonstaints.
vidn of dvlopmntalonstraint has om from two sours.on is
forbidde morphologies, phnotyps that on an imagin and tht sm
funtionally rasonal ut ar nvr osrvd.Th sond is th rstritd
rang of phnotypsthat mrgfollowing natural or primntalprtura-
tion of devlopmnt. In oth ass som of th st ampls om from
t t r a p o dd i g i t p a t t r n s( F i g u r 3 ) ( H o l d r 1 9 8 ; A l r h a n d G a l 1 9 8 5 ) . B experimental redL
Although thes approahs onviningly dmonstratthat th prodution
of phnotypsan iasd,th aussof thsiasshv rmaind lusiv.
A phnotyp ould ..missing'' baus it is diffiult to produ dvlop- mitotiinhib
mntally, or ausof a physial or gomtri limitation, a lak of gnti
variation, or an unrognizdfitnssost. Distinguishing among thspossi-
ilitis has provn quit diffiult. vn whn it is possil to infr th xis-
tn of dvlopmntal onstrints y poss of limination' idntifying
what aspts of dvlopmnt ias phnotypi vition reiss furthr hal'
lngs,and fw lar ampls ist.
A rltd qustion is why suh iassist. on planation is that dvl-
opmnt is ativly ..uffrd''to produ onsistntphnotypi outomsin
spit of gntior nvironmntaIprturations.This ida was arfully arti- developinglim
ulatd y onrad \/addington(7957), who nmd it liztitl.Dvlop-
mntal biologists hav long rognizd a similr phnomnon thy ll
regultio:th aility of an mry()to onvrg on a normal phnotypfol-
Figur 3. Bisd produtio
lowing primntal manipulation. \7addingtonblivd nliztion to b
may rstitth rang lf pl
th produt of gnti assimilation' a pross whry a phnotyp that was fro-rprimntalprtur
initially invokd y th nvironmnt is susquntlygntiallyfid through r a n g o i p h n o t y p sr h i s
sltilnto maintain that phnotyp. Ivan Shmalhausn (1949) proposd i n d p n d r lt a s st l f d i g i t r
that stailizing sltion produd muh th sam rsult. othr possil amphiiirns.AIthough four
sours of dvlopmntalonstraint ar a progrssivloss of fliility s th thssult from th loss ol
prosssof dvlopmnt om urdnd y historial lgais (.Wimsatt tratingth dvlopingliml
A l ha n d G i r l ( 1 9 8 . 5w )
1986) and th liklihood that modifitions in arly dvlopmntwill hva
frogs and salmandrs.ln
roadr phnotypi impat thn latr modifiations (Arthur 1988; Gould
random ut instirdnfor
1989\. spis.(Bsdon Alrh l
evolutionaryredutionin digit number
l dvlpmntaliologv dr-rringtlr
)02). volutionary and populatitln IV
nnts lak o whos ()ntnts
ding volutionaryprossS.
^fl/
\l
n lrllngdrpardly. harls
..th laws of Frogs
;t' wondrdwhthr
typil variirtionnd favor ohs''
raryiologistsWr prpard to an-
lldr 198; aynard Srnith t al.
tion tl"rtth organizlrtionof dvl-
Lof phnotypssmsasona1- ill
,.''/lil'
. . r yp o s i h l p h r l o p i o u o r i s
Salamanders
a s ( A l r h1 9 8 9 ;! i l k i n s 2 0 0 2 ) .
Lundrlyingsoursnd long-trm
\l
;ti]ints.
has lmfrom two sours. on is
lt on n irlagin nd tht s-r
srvd.Th sond is th rstritd
g natr-rral or primrrtalprturl"la-
of th st ampls om from
. 1 9 8 3 ;A l r h a n d G l 1 9 8 . 5 ) . B eperimentalreductionin digit number
;l1,dmonstrtthat th prodution
f thsiasshav rmaind lusiv.
' it is diffiult to prolu dvlop- mitotiinhibitor
)mtri limitation' a lak of gnti
:. Distinguishingamorrgths possi.
hn it is possilto infr th xis-
pross of limir"rtion,idntifying
otypi variation raiss furthr hal-
nstraint on volutionay hang' within a population y minimizing its immdiat phnoypi impat On ny
) J o s p hN d h a m ( 1 9 3 3 )p o i n t d givnindividr-ra[;rlr inrasdgnti variatiotl rnight latr ot-t-t
gist for
s s s ,s u h a s l l d i v i s i o n , m o r - th mill of sltiorrunder altrd nvironmntal iumstansor irr diffr-
ontinu indpndntly aftr - ntgntiakgrounds (Gison and Dworkin2004). On th othr hnd, g-
phnomnon dissoibilit and nti variants that might othrwis produ phnotypi onsqunsand
rlyingbasisfor dissoiility may thrbybonrvisil to sltionmight b maskd and thrforfftivly
:nmongnralftur is proaly drift s nutrl llls.W still hav only ir limitd undrstanding of th d-
pltfuntionalindpndn- grto whih sltionstalislrdrnhanismsrhat uffr phnotvpi veria-
rd Kirshnr 19971Hansen 2003). tion and th dgro whih suh mhanisms limit lr ias th vrlutionay
lity is that it allows modifiations fatof lads (\ist-rhard 2003).
ut ftirllyompromising th ntr-
LH2 Ho-t
Anirals in diffrnt phyla ar so Dl
nologous strLltursnd dtrmin.
tntiotls(Nilsn 2002)' This ph. td
'lr ntai thnls of omptiv |nS
BMP4
.d (Hall 1992 kaff |996). Th no. Ap t:holin
D
a n i m a l sh a s a t r n i q u o m i n t i o n D
rhittural(suh as ody symm. hh Nkr2.2
' of ll divisitln)o Ap Ms
, r oth (suh s lG-H
lody-pIanol1ptlras dply inflr.r-
animals up ttl th prsnt (Gould
Figur5. Ho prado. Homllogous gns pi]ttrn t mrvos of fruit
97). Dvloprrrntaliolgists, fo
flisnd mi dspit th plrylogrrtidistan nd phrrotypi disprity lf
rofourddiffnsin th way in
thesanimls. Th mlst fmous of ths r gns lf th Ho lmplx, whih
rrorphgnsislur ir-rmryos of stalishposition of l.od rgions |ong th ntoPostrior is. Mnv othr
7).No on prditlthat rnany of honr<llogous gns pttrn th rins, hrts, ys, and lims of arthrolds and
t to rgulatlvlopmntthrough- hordts.Tlrs htlnrlogol.lsglls ar prssd in roughly similar pattrns
duringdvlrlpmnt ut odu r,ry diffnt phnor1'ps(Grhart ar-rd
l dvlopmntlor rllutilnaryi- Kirshnr 1997; Wilkins22). Tlris asi rsr.rltsuggststnsiv ..rwiring''
of dv,lopmrltalgn ntworks drrring th ours of volution (s txt). (From
s h o w ' t h r i r r l i I l rg t l < l t n i() r g n i -
lfray 2003.)
osl1lhil and mous: this imag has
224 uolutio d Dulopmt
What is lar at this point is that th sam asi omplmnt of dvlop- and rgulatoy squn
mntal rgulatory gns' som of whih ar prssd in roughly th sam r- a o l i s h i n gs u h i n t r r
gions of th mryo' an produ a wid divrsity of nimal ody plans. of muttions, singl as
Figuring out how this lurs is on of th grand hallngs of volutionary t l. 200J). Mny prot
dvlopmntal iology (Ghart and Kirshner 1997; arroll t al. 2001; nuli aids, and small
Davidson 2001). A dtaild rsolution of this Ho parado lis many yars d r a m t i f f t s. hngi
in th futur, ut hangs in th organization of dvlopmntl gn nt- tion or altr affinity dr:
works ar likly to fom an importnt part of th solution (arroll t al. an rat or modify a i
.Wry
2001; Wilkins 2002; t l. 200). rgion.
Two asi kinds of rwiring within gn ntworks hav n doumentd, hangs in dvlopm
roughly orrsponding to quantitativ and qualitativ hangs. Th first, or pat on th history of lif
quantittiv' atgory involvs a hang in th loation or timing of prs- and why gn ntwlrks
sion of a rgulatory gn. Suh hangs an aris whn a ntwork is altrd
upstram of th gn of intst.Th loation or timing of th dvlopmntal
prosssth gn ontrols will hang as a rsult, potntially altring organ- Frm ly{arovoluti
ismal phnotyp. For xampl, th spatial xtnt of Ho gn xprssion dif-
frs in ways that orrspond to sgmnt idntity in rustaans (Avrof and Phnotypi hangs,inlt
Pate| 1997) and to vrtrl anatomy in ttrapods (Burk t al. 1995). Th variation within populati
sond, or qualitativ, kind of ntwork rwiring ntails a hang in whih d- mntal volution at thrs l
vlopmntal prosss ar rgulatd. Ths hangs an our whn a nt. aout th gnti asis ft
work is altrd dwnstrm of th gn of intrst.For instan,signalingvia wild populations (as opp
th rptor Noth gults spifiation of nurnal ll fat in insts and th la) or aout what k
spifiation of whit lood ll fat in vrtrats (Hitzlr and Simpson tion in ntwork funtlon
1991; Guidos 2002), involving two vry diffrnt sts of fftor gns. Sim. happn ovr llng tims
ilarly, th tansription fator rahyury rgulatsnotohord dvlopmntin This pauity of inform
urhordats and hordats ut plays no suh rol in othr phyla, whih lak D v | l p m n t al i o l o g i s
this strutur; instad, it rgulats dvlopmnt of th gut and othr stru- influns of gnti an
turs (Thnau and Sholtz 2003). hv workd largly witl
Dozns of ass r now known whr an volutionary hang in th x- gnt that omprisons 1
prssion prfil or downstrm targts f a rgulatory protin has altrd nisms. For thir part' po
anatomy. Som ass nompass hangs at th sal of ntir ody plans, a n d h v f t l u s dp r i m r
suh as th transition from rdial to ilatral symmtry during arly mt- protins, protin strutL
zoan volution (Martindl t al. 2002) and th latr transition ak to radial rathr than gn intrt
.Wray
symmtry in hinodrms (Low and 7997). Othrs involv modifia- hr ar, of ours'
tions in organs' suh as th tansition from fin to lim during th origin of variation in th oding :
ttrapods (Shuin et a|.7997), or volutinary innovations, suh as arthro. u l a t o r y g n s ( . g . ,A y a
pod wings (Avrof and ohn 1997). vn subtl diffrns in anatomy ar T h s o p u l a t i o n - l v ll
oftn th rsult of modifiations in gn ntworks: ampls inlud diffr- o f s g r g t i n gv a r i i o n
ns in ristl pattrns in flis (Strn 1998; Skar and Simpson 2000) and opmntl mhanisrns
olor pattrns on uttrflywings (Brakfildet a|. 1996i Bruntti t al. 2001; toris (.g.' Wray and
Gompl t al. 2005). 1998), or among popu
How do hangs in dvlopmntal gn ntwoks aris? Thr ar many t a l . 2 0 0 4 ) , a n r v a ll
ways in whih this an happn-any spifi intration etwn maromol. hangs in sltion. (
uls is a link in a gn ntwork' and any diffrn in intrations altrs th out population gn
ntwork. Th two most ommon and important kinds of ntwork hangs opmnt rmlins a gleri
ar proaly intrations twn protins and intrations twn protins vlopmntal iology.
uolutil dd D eU|opmt 22.t
am bsi omplmnt of dvlop- and gulilt()rysqttnsin DNA (Davidson 200 l). Crating, rnodifvirrg'or
' prssdin roughly th sam r- aolishingsuh intrations an <lurwith th sirnplstand most ommon
l divsit1,of niml ody plans. of mutations' singl as sustitr-rti<lns
(Grhrt nd Kirsirn 1997 wra
grand hallngsof volutionary t al. 200). Many protins ssoiatspifiallywith othr protins or witlr
'shnr19971 arroll t al. 273 nuliails,and sr.n:rll
hangsin inding affinity or spifiit,van produ
this Hox pradox lis many yars dranratiffts.hngirrg singl nrirroilid n rt a spifi intra-
zation of dvlopmntal gn nt- tion or altr affinity dramatially; similarly' hanging singl nulotid
lart ot th solution (Carroll t al. enrtlr nrodify inding sit fr tr.lnsripti()nfator ilr a rgulato1'
rglon.
e ntworkshav n doumntd, hrngs i dvlopmntalgn trtwokslaly hv had a profound im_
d qualitativhangs.Th first, or pat On th histlry of lif. As disussd nxt, hwvr, undrstandirrghow
n th loation or timing of xprs- nd ,hy gn ntwrks r ..rwird'' rrnainsa ntrl hallng.
:an aris whn a ntwork is altrd
tion or timing of th dvlopmntal
a s u l tp
' o r n t i e l l ya l t e r i n go r g a n - Frm Marovolution to irovolution
xtnt of Hox gn xprssion dif-
idntityin Llstaans(Avof and hnotypiangs,inluding vt>lutionay hangsin lvloprrrnt, gin s
ttrpods(Burk t al. 1995). Th varii.ttion within populatilns.Rltivly fw str.rdis hv addrssddvlop-
wiring ntailsa hang in whih d- mntalvolution at this mirovolutionary sal. As sult, w know littl
]s hangsan our whn a nt- aboLrtth gnti sis flr nrodi6atir-rs in dvloprntalgn ntrvorks irr
: intrst.For instan,signaling via wild ppulations (as opposd to What indud mutations an aOmplish in
Lof nuronlll fat in insts nd thla) or aotlt wt kinls of volutilrraryml-r:rnisls s()rtgnti \,ri-
vrtrats(Hitzlr and Simpson tion in ntwork funtion (s opposd tl simply osrving tlrt hn{:saIr
jiffrnt sts of fftor gns. Sim- happnovr long tirnsals)(Purugganar2000; Wry t al. 200).
:gulatsntohord dvlopmnt in This pauity of informtion drivs from tw<lhistoril trrrdsin rsarlr.
,uhol in othr phyla, whih lak Dvlopmntalilogists, in th first instan, lrav stivn to liminat th
)pmnt of tlr gut and oth stru- influnsof grrtird rrvironmrrtlvri:rti<lfrorrr thir rral'ssan.l
havworkd largly with modl systmsthat ar so phylogrrtiallydivr-
n volutionryhang in th - gnt that omparisons ild littl inflrrrrtionbotrt volrrtilnaryr-nh.
f a gulatoryprotin has altrd nisms.Fo thir part' popr-rlation gntiistshav largl1,ignord dvl<lpmnt
s at th sl of ntir body plans, andhavfor"rsd pimarily on nzymsralthrthirn dvlopmntalrgtllatory
ttrl symmtry during aly mta- protins,rotin strutur rtlrr tharr gn prssilln, alrd singl gngs
nd th latr transition ak to radial rathrthan gn intratilns.
a 7997). othrs ir.rvoivmodifia- Thr ar' of <lurs,prions. Svral stlldis hitv xirnlind gni
om fin to lim during th origin of vritionin th tldirrg nd ls-rgulatlrysqunsoI dvlopmntal rg-
ionar innovations,suh as rthro- u l a t o r yg n s ( . g . ,A y a l a a n d H a r t l 1 9 9 ; I - u 1 w i ga n d K r i t m n 1 9 9 . 5 ) .
:n sutl diflrnsin anatomy ar Thspopulation-lvl arralyssprovid a glirr-rpslf th n.1turand lvl
ntworks: ampls inlud diff- of sgrgatingvariirtion ftlr fw rgul:ltory gns. lmparisons of dvl_
)98; Skar and Simpson 2000) and o r n n t anl t h a n i s m sa o n ] l < l s l yr l a t d s p i sw i t h l i s t i n tl i [ h i s -
ld t al. 1'9961'Brunettit al. 2001; t o r i s( . g . ,l / r a y a n d R a f f 1 9 9 0 ; S w a l l a n d J f f r v 1 9 9 6 ; G r i t a l .
1 9 9 8 ) ,o r a ] o n g p o p r - r l a t i l nrsv i t h d i s t i n t r o r p h o l l g l s( . g . ' S l r a p i r o
r ntwoks aris? Thr ar many t a l . 2 0 0 4 ) ' n r v a l h o w s p i f i d v l l p m n t arlr r h n i s m rs s p o n d t o
:ifi itrationtwnmaromol- h a n g si n s l t i r n .O v r a 1 l , t l r o u g h , t h g n r a l l a k t l f i n f o r l a t i n
l diffrnin intrirtions altrs th a o u tp o p u l t i o n g n t i sa n d s r a l l . s a l v l u t i o r r r y h : l n g si n d v l -
:lportntkinds of ntwork hangs o p m e n tr m a i n s a g l a i n g g p i n t h r r t n i f i a t i o no f v o l u t i o n a r y r r n d d _
r s n d i n t r . t r i o nbs t w np o t i n s vloptr-rti.rl iology.
226 uolutild Duelopmet
l lrmnand th physiologial Shpiro,. D., . ' Marks' . L. Piel,B. K' Blakmn, K. S. Nng, B. Jnsson,
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lations. Ntur Getis 32,: Shuin'N., . Tain, nd S. B. arroll. l997. Fossils' gnsand th volution of
animl lims. Ntur 388: 639_648.
aptation:Th adaptation of DNA. Skr'N., and P. Simpson. 2000. Gnti analysis of ristl loss in hyrids twn
Drosophil mlogster and D. simIs provids vidn for divrg of
ion ptlstzvgltiisolation: Fors, ls-rgultory squns in the ht-sut gne ompl. I)t,elopmtl
1094. Biolog 221: 148_|67,
lgisof th Vrtbrt ShIton' Strn,D. L-. 1998. A ol f Ultrbithor in roplrologial difftrn twn
Drosrlphil spis.Ntur 396: 46-466.
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o|pdr.1tiu B io l o g, 2I-62. San variation. uolutio 54: 179-1091,
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.Wray,
/alkr'G' A. B. J. Swalla, 9-361.
'.
Th oigin nd volution lf animl Striklr,A. G., Y. Yallmoto,and W. R. Jffy.2001. arly and lt hangs in
\dml, of Sin:sUS 94: Pa6 prssiOnolnpany y dgnrtiorrduring vfisdvlopmnt.
Duelopnrctt Gs d uoluti.lll2|1: 138-144'
4. Pirir-rulprssionpattrns of Swalla,B. J., and W. R. Jffrv. 1996. Rquirmnt of the gn f xprssion of
long.grmtls.Ntur 67: hordtfatursirr a tillssasidin |ara' Sie274: 125_128.
Thnu,U., and . B. Shlltz.2003. origin and volution of ndodrm and
lation gtisof rgultory gns. msdrm. lntcrtkll Jourl of Dulopmtl Biolog 47: 53_59.
Tru'J. R., nd . S' ag.200]. Dvloprnntalsystmsdrift and flxibility in
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ony: Dvlopmntalmhanisms and .!agnr'
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B i l o g 4 5 : . l 0 8 _ 2 2 '
26 uoltil nd DuklDlnet
Foundations in Natu
U n d r s a n d i n gw h y s p i
q u i r s d r w i n g i d sf r o m
half ntury ths idas, t
bn wovn into a ohr
iorists had to larify hw t
th mundn to th fasrn
to ask' ..Why ar thy
(196) proposd that thr
tions. Thr wr standard
] volutiOnf devlopmnt.
bg1'47:675-684.
r o i h i n o d r md v l o p m n ti s
t |20, Supplmnt:97_106.
l h o f f ,M . P i z r .M . V . R o k m n '
Soial Bhaviorand
transription1 rgultion rn
20 177-I4I9.
ralgulatorygns and hinodrm
Most nimals liv solitary livs' Apart from oming togthr to mat' most
individuls forag, mov aout, anl void prdatos on thir own. Yt as
highly soial animals ourslvs, W a fasinatd by spis that dvlop
.W
long.trm riationships and liv soial livs. want to know why suh
.!e
spisliv in groups and why th groups tak th form that thy do.
want to know why som groups prsist, whras Othrs ar mor transitory.
W want to know why ooprtion oms in many typs and why oopra-
tiv and omptitiv tndrrisshp most rlationshipsnd thus oist in
mny soitis.\Walso want to know why partiular rols twn individ-
uals and th ss dvlop and why som appar univrsl, whil othrs do
not. Answrs to ths qustions ontinu to m]'ut sin w oftn look
to animalsto lrn aout ourslvs'th ility to fous th lens of sinon
answingthsand other qustionsabout human soiality has n loudd
y prolms.Bhavior, lik all oth fatursof th phnotyp,volvs, and
thfar that what nimals do, humans must also do has oftn ausd popl
to shy away from asking' or at tims vn bing allowd to ask' qustions
aboutth adaptiv valu of soial bhavior.
27
28 Soil Behuiord Soiobioklg
ontognti dvlopmnt' nd volutionary history. What h addd was a baus saf nst sits.
fourth that foused on unravling a havior's funtion y idntifying how it safst trs. \With opp
augmntd an ator's survival or rprodutiv valu. This mphasis on x- polygyny and right m
plaining havio in trms of sltivnfitsto individuals hlpd mov - tis of many othr gro
plnations of soial funtions away from what is good for groups to what is parativ lns (Lak 196
good for th individuals that ompos ths groups (\/illiams1966). Tinr- mthods of atgorizin
gn's insights idntifida onfusion that oftn prvadsth study of havior, among spis hav
namly, that ther ar roadly two typs of planations:proimat and ul. omparativ mthod t
timat. Idntifying triggrs,or rlasrs,of a havior or th timing of its ap. among havioral out
paran during ontogny yilds proximat planations austhy fous road-rush undrsta
on th mahinry within animls tht gnrats th havior. Idntifying yond what ritis (.g
how a bhavior altrs an individual's fitnss or th way in whih historial gain a mhanisti un
antdntsdtrmin what matrial onstituts th rprtoir of a spis why partiular soial r
yilds ultimat xplanations;thy fous on th volutionary roots and ons- irumstans was n
quns of havior. of ours' this dihotomy is lurry sin mhanisti x p l a i n i n d i v i d u [d i
pattrns hav funtional onsquns,and as a rsult thy too volv. But
for th most part' kping th lvl of planationslar whn aounting for
why animals havth way thy do was a major rkthrough.If this pra- mrging Thorti
ti had n aptdwidly, many of th argumntsthat hav plagud th
study of soial havir, spially as it rlats to humans, ould hav n Two diffrnt thoris
avoidd. ogists. Why animals sh
Sond, sarhingfor a havior'sfuntion, or adaptiv valu, nssitats thmslvs,is a qustio
assssingits rlationship to th nvironmnt.Long for Darwin, iologists was puzzld aout wh;
.W.hat
intrprtd morphologial traits in rlation to th nvironmnt. Dar. through ourrd wh
win did Was dmonstratthat suh adaptationsould aris without a rator' Until th mid-1960s th
thus omplling arly naturalists to sarh for nvironmntal prssurs tht dwrds (7962) am
ould serv as sltiv,or shaping' gnts.on of th arlistand most om- mans of rgulating po
plling studis that dmonstratdthat the adaptiv valu of ntir soial sys- opratd. Williams (19
tms ould xplaind y diffrns in ology Was rook's (1964a, ial intrations augm
1964) study of wavrirds.By xamining th foraging, nsting' grouping, Hn sltion atd
and mating havior of ovr 100 spis of Afrian Wavrs that oupid individuals, thn aou
haitats ranging from tropial forststo grasslandsand svannas,rook was straly hlp ripints
al to dmonstratthat ologial fators assoiatdwith th distriution of prolm. Hamilton's in
food and prdators Wr rsponsilfor th road soial diffrnsthat th only dirtly through
spisxhiitd. Thos spisthat livd in forststndd to solitary' in- rlativs. Th ominat
stivorous,trritorial, draly olord' and monogamous. Crook argud that clusiue fitess and provl
ths traits ovarid baus insts wr hard to find. Sin fding young vn stril asts of ins
rquird frqunt visits to th nst y fathrs and mothrs, monogamy and prodution of rlativs
pairwise dfnse of aras whr insts would rppar ld to monogamy fiint,'' and Hamilton
and trritoriality. rook also suggstdthat uilding rypti nstswithin tr- ration of te kiship
ritoris and th volution of dra oloration would rdu prdation. on- drivd y a rlativ m
vrsely, thos spis that livd in savannas and othr opn landsaps rlativ dvalued y th
tndd to soial, trsof sds'olonial, and polygynous' with mals ov- for undrstanding th
rd in rightly olord fathrs.Sin sds oftn ar lumpd, Crook - and has n supportd
livd that nfitsssoiatdwith groups ing al to sarh fo sdsovr That nfits and ot
wid aras ld to th volution of soiality and that llniality mrgd for th dvlopmnt of .
Soil Beuior d Soio|iolog 29
.!7hat aussaf nst sits wr rar, induing mny pairs to nst togthr in th
r history. h addd was a
lr's funtion y idntifying how it safsttrs. !ith opportunities for ukoldry prvalnt, rook argued that
tiv valu. This mphasls on x- polygynyand right mal oloration wr also favord. Over th yars soi-
its to individulshlpd lnov - tisof many othr groups of animals hav bn analzd by us of this om.
hat is good for grouPs to what is p a r t i v l n s ( L a k | 9 6 8 i l a r n a n 1 9 7 4 i K r u u k 1 9 7 5 ; F r i k 1 9 7 5 ) .A l t h o u g h
r groups(Williams 1966). Tin- mthods of atgorizing havior and lassifying volutionary rlationships
n prv.rdsth study of havio' among spis hav n rfind y ovrlaying havio on phylognis, th
[ plrrtitlns: proimat and ul- omparativmthod y itslf, with its rlian on statistial orrlations
hvioror th timing of its zlp_ among havioral outoms nd eologial fors, ould only provid a
planationsausthy fous road-rushundrstandingof th volution of animal soitis.To mov -
nratsth havior. Idntifying yond what ritis (.g.,Lwontin 1979) |aimd wr ..just.sostoris'' and
ss or th way in whih historial gain a mhanisti undrstanding of ologial ausation, ttr gsp of
titutsth rprtoir of a spis why partiular soil rlationships among individuals dvlopd in spifi
t h r ' o l u t i o n e r ryo o t s a n d o n s _ irumstanswas ndd, and this rquird th dvlopmnt of thoris to
ltom is lurry sin mhanisti plainindividual dision mking.
d as a rsult th too volv. But
. n t i o n s l rw h n o u n t i n gf o r
Lmajor rakthror'rgh. If this pra- mrgingThortial Framworks
argumntsthat hav plagud th
:latsto humans, ould hav n Two diffrnt thoris mrgd, on from soial thorists and on from ol-
ogists.Why animls should hlp others, oftn t th xpns of rproduing
i o n ' o r a d a p t i v v l u ,n s s i t t s thmslvs' is a qustion that ppld biologists for nturis.vn Darwin
nt. Long forDrwin, iologists was puzzld aout why so mny soial insts wr stril. A major rak-
ln to th nvironmnt. What Dar- throughourrd whn Hamilton (1,964)proposd th ida of hi sltio.
rionsould aiswithout a rator' Until th mid-1960s thr was onfusion out th units of sltion.Wynn-
r for nvironmntlprssurs that dwards (1962) hampiond th viw that soial havior volvd as a
. on of th arlistand most om- mansof rgulating populations; hn goup havior ws whre sltion
adaptivvalu of lltir soial sys- optd.\(illiams (1966) ritiizd this viw y dmonstratingthat most so-
in ologwas Crook's (1964a, ial intrationsaugmntd th fitnss of individuls that ngagd in thm.
n g t h f o r g i n ,l 1 s t i n gg, r o u p i n g , Hnsltionatd on individuals. Br'rtif bnfitsof bing soial aru to
of AIrin wavrs that oupid individuals' thn aounting for th volution of altruisti ats that dmon-
asslandsand savannas,rook was straly hlp ripints ut aru no dirt nfits for altruists boms a
i sslitd with th distribution of prolm.amilton's insight was that individuals an pass on thir gnsno
h road soial diffrnsth.1tth only dirtly thr<lughoffsprirrg ut als ir"rdirtlyvia th offspring of los
in forststndd to b solitary, in- rltivs.Th ornination of ths two nefitsprodus an individual's ln-
d monogamous.rook argud tht Iusiufitess and provids a fitnss aounting for th volution of altruism.
: hard to find. Sin fding young vn stril asts of insts an volv if thy an suffiintly nhan th r-
:hrsilnd mothrs, monogamy nd produtionof rltivs.But th dvil is in th dtails of what onstituts,.suf-
would rappar ld to monogamy fiint,'' and Hamilton providd a alulus for dtrmining this with th
Latulldingrypti nsts within tr- rationof the kiship oeffiit. His rul that th nt rprodutiv nfit
tion woLlld rdu pdation. on- drivdby a rlativ must gratr than th ost inurd y th altruisti
anns and othr opn larrdsaps rlativdvalud by th riprol of rh kinship offiintprovidd the ky
al, and polygynous,with mals v. for undrstanding th volution of th rnost fundamntal soial haviors
ds oftn ar lumpd, rook b- andhs n supportd y many studis on insts and vrtrats.
s ingabl to sarlrfor sds ovr That nfitsaIrd osts ar fftd y kinship also ratsth onditions
ality and that <llor.rility mrgd forth dvloDmntof onflit within familis and twnth ss (Trivrs
240 Soil Bbuior ]Sociobiolog
1972, 19741Trivrs and Har I976). Sin parnts ar rlatd qually to all
thir offspring, ut offsping ar mor losly rlatd to tlrmslvs than to providd th lns for
thir silings, onflit ovr ptdinvstmnty parnts will oftn mrg. nting options. And v
In gnral, offspring will attmpt to prvnt parnts from qually sprading mining whih s th.
rsoursamong all thir offspring. Howvr, th rltionship bven parntal volutinary history a
invstmnt and sual omptition nray hav n th most profound tn. Whil th soial th
sion of kinship thory for shaping th study of soial volution. Armd with part of th polygyny-t
th rsults of Batman,s(1948) study of fruit flis that showd that rprodu- rang of ll1aring svstl
tion y mals and fmals was limitd by diffrnt fators, Trivrs proposd widr rang of o[
that whn on s invsts mor in th raring of ffspring than th othr, potntial for polygyn
mmbrs of th lattr will ompt for mmrs of th formr. In ssn,th tion of th limiting s
sx that invstsmor in raring omsa limiting rsour'and th s with find it diffiult to ontl
..ss'' in groups, or rsour
rsours should invst hvily in mating tivitis, inludirrgfind.
ing and guarding mmrs of th panting sx. grgat without inurr
Th sond thory mrgd 1rom ology. Pionring work y Orians, as in th N4aynard Sm
.Willson mals; if fmals wr
Vrnr' and showd that th mating systmof rd-wingdlakirds
dpendd on how ologial onditions afftd th ss in diffrnt ways for polygyny ould nol
(Orians 1969). Thy slrowd that within on population oth polgyny and
monogamy ould oist, and that th lravior of fmals indud this
mating-systm variation. Thy argud that on landsaps with rsour mrgnof a S
paths that varid in quality, mals on those of th highst quality ould d-
fnd nough rsoursto suPport th luths of m:rny fmals. on thos of Th lmntsof a synt
lowr quality, howvr, trritorial mals would luky if thy w :rlto h a m l 9 8 6 ) . F m a l sr v
.Wht aus thir soial rl
sunough rsoursto support th offspring rfon tmal. mad
thir modl uniqu was that it dpndd on assssmnts y frrrals-th dmands of th physi
polygyny thrshold-to dtrmin whthr mating polygynously or monoga- sminal rviw, ths
mously was in a fmal'sst volutionary intrst' and prdation prssu
h undrlying prmis that fitnss dpndd orr what othrs wr doing pndd mostly on solv
st th stag for aynad Smith's (|977) insight tlrat th volution of th saf sites for thmslv
fou major mating systms is a gm in whih what is st for on sx d- mts. oniy aftr ss
pnds on what is st for th othr. Maynard Smith proposd tlrat whthr lndsps wr dtr
On s should invst in aring urrnt offspring should dpnd on how fo mting opportunit
likly th othr s Was to ontinu raring thos offspring. Sin parnts' in- ations y rdingma
trsts ar similar ut not always idntial, Maynirrd Smith showd that hirvior ultimatly ld
thr was a pair of strtgis' on for th ml and on tr th frrral,tht, y suadults and dv
whn prformd togthr, would lad to an volutionarily stl stratgy g n a t i o n s l s o a r i
(SS). Suh a stratgywould our only if it did not pay ithr parnt to d- itis (Rubnstin 198
part from his or hr stratgy.Thus it was possil for iparntal ar, uni- Th synthti mod]
parntal ar proffrd y ith s' or no parntal ar at all to an SS wr bundnt, sp
undr spifi ologil onditions. IWhatth nvionmnt dtrminsis th would low nough t
tnt to whih pantalar is nddto rr offspring. If only on parnt is aging or antiprdator
ndd,thn the liklihood of itlrpartnr dsrtingdpndson th asof and th groups tht fo
finding an additinal mat. For fmals in partir'rlar' this liklihood is also dfndd y singl ma
afftd y th xtnt to whih prnting rdus hr frtility in futur rpro- sult. Flowvr, if rsou
dutiv pisods.Fo th first tim soial asptsof mating havior ould tion among fmales rr
haatrizd nd tid to srlifi sltiv fors. Suzrl asvmrtis would vary, nd fmal
than dfnding unst
Socil Behuior d Sociobiolog 241
partsar rrlatd qually to all
providd th lns for idntifying how rsours and prdation afftd par-
sly rlatd t thmslvs tharr to
nting options. And whn only on prnt ws rquird for raring, dtr-
nnt parntswill oftn mrg.
mining whih s tha should dpndd on trad-offs shapd y oth
1t parntsfrom qually sprading
volutionary history and rrnt ologial irumstans.
:, th rlationshiptwnparntal
!7hil th soial thorists Wr panding th havioral dision-making
v n th most profound xtn-
part of th polygyny-thrshold modl to aount for th volution of th full
y of soial volution. Armd with
rangof mating systms' ologists wr panding th modl to aount for
rit flisthat showd that rprodu-
widr rang of ologial dtminnts.For mlen nd Orling (1977), the
Jiffrntfators,Trivrs proposd
potnti:rlfr polygyny dpndd on oth th spatial and tmporal distriu-
aring of offspring than th othr'
tion of th limiting sx. If fmals wr widly sattrd' thn mals would
rsof th fornrr.In ssn,th
find it diffiult to ontrol mor than on fmal. Altrnativly, if tmalslivd
limitingrsur,and th sx with
in groups, or rsours wr lumpd so that fmals ould tmporaril ag-
:l mating ativitis'inluding find.
ggatwithout inurring muh omptition, thn polygyny ould rsult. But
g s.
as in th aynd Smith modl' muh would dpnd on th havior of f-
ogy. Pioning work y orians'
mals; if fmals wr rptiv mostly at th sam tim, thn th potntial
lg systlof rd-wingdlakids
for polygyny ould not ralizd.
fftdth ss in diffrnt ways
ln population oth polygyny and
havior f fmls indud this
hat on landsaps with rsour
mrgnof a SynthtiThory
os of th highstquality ould d- .!Vrang_
Th lmntsof synthti thory wr rystallizing (Rubnstin nd
hsof many fmals.on thos of
ham 1986). Fmals wr viwd as th driving for of soial volution -
vould luky if thy w al to
:fspringof on fmal. What mad austhir soial rltionships rfltd thir bst rsPons to mting th
dmarrdsof th physial nvironmnt,and, as Aland r (|974) argud in his
l on assssmntsby fmals-th
sminal rviw, ths largly fll into two atgoris: rsour distriution
r matingpolygynously or monoga-
and prdation Prssur. Maimizing rprodutiv suess y fmals d-
r intrst.
pnddmostly on solving prolms posd y aquiring rsours or finding
lndd on wht othrs wr doing
saf sits for thmslvs or thir young rathr than on finding or suring
) insight that th volution of th
mats. only ftr assoiatiorrsamong fmls and thir distribuions on
whih what is bst for on sx d-
landsapswr detrnrinddid mal stratgisassoiatd with ompting
nard Smith proposd that whthr
for mating opportunitis om into pIay' Susquntrsponssy fmals to
offspring shold dpnd on how
tionsby brdingmls and, in turn, rsponssy rdingmals to fmal
g thos offsprirrg.Sin parnts' in.
bhavioultimatly ld to an volutionarily stal mating systm.But ations
ial, aynard Smith sl.lowd that
by suadultsand dvlpmnt of rlationships among mmrs of diffrnt
mal and on for th fmal. that.
gnratinsalso am into play as soial strutursdvlopd in omplx so-
an volutinarily stl stratgy
itis(Rubnstin1986, 1994).
f it did not pay ithr pant to d-
Th synthti modl prditd that in nvironmnts in whih rsolJrs
s possilfor biparntal ar, uni-
wr abundant, spially whn thy wr vnly distriutd, omptition
ro prntal areat all to an SS
would low nough to prmir fmlsto aggrgat.If suffiintlylrge for-
t th nvironmntdtrmins is th
agingr antiprdator nfits ould drivd y ths aggrgating fmals,
r a ro f f s p r i n g I. f o n l y o n p a r n t i s
and th ]roupsthat formd we not too larg, thn ths groups ould
rr dstingdpndson th s of
dfnddy singl nrals, and so aild harrn_dfns polygvny would r-
n partiular' this liklihood is also
sult.Howvr, if rsursWr mor pathily distriutd, so that ompti-
r d u sh r f r t i l i t yi n f u t u r p r o -
tion among fmals was priodially intnsifid, thn fmal group sizs
lsptsf mtirrghavior ould
.tiv fos. Sxual asymmtis would vry, arrd fmal assoiationswould om mor tratrsitry.Rathr
than dfnding unstal groups of fmls, mals instad would attmpt to
242 Socil Behuior nd Soiobiolog
ls. In ths systms of rsour- am an aility to susiston mor aundant, lowr-quality vgtation..i7hn
ld dfndth st paths and thus it was pathily distriutd' as Was th s for impala (Apcrlsmelmpus),
:s for th longst priods of tim. If duk(Redu spp.), and sm gzlls (Gzll spp.), mals dfndd
.ily ut i paths that wr lrg, th st paths tht fmls prfrrd. !hn th vgtation was mor vnly
sonallyin abundan' thn omp- distriutd,whih oftn rsultdsimply from th fat that largr spissuh
; would failitat th formation of as land (Turotrgus orv) and ap uffal o (Scros cffer) ould us
angwidly and follow th shifting Vn th lowst-quality items, largr groups formd. Baus th largst
als would thus ford to ithr spisviwd lrg ontinuous swards of a landsap as aptal, r-
'r, and thn tnd, on rprodutiv souomptition was virtually liminatd,nd many mals ould assoiat
position thmslvs at th intrs- rvith many fnrals.With suh high lvls of rnale_nralomptition' dfns
- for fmals to visit tem (lk.king of a smal] sugr'rpof frals anrimpossil, and dominan dfns
ral omptition would gnrat a sstmsdvlopd.
polygyny, nd in th lattr as f- Th sam sorts of onntions btwn hanging ologial irumstans
ppotunityof simultanously om- and havioral disitln making shad th typs of soiality xhiitd in
.s7hnvr
whom to mat. rsous oth taa. A rif survy of mammals illustrats som of th mor gnral
riutd,high lvls of omptition pattrns.For th quids, th los assoiation twn food and watr
n-
ups. As a rsult, individual fmals als hors (quus bllus) and plains zera (quus burhelli) fmals of
nsuringa rgular supply of a r* diffrntrprodutivstatsto assoiatprmanntly.Thus mals ar al to
ls who wr srhing for mmrs dfnd suh groups, nd so-alld harms form. .Whn ths two rsours
d prdationrisk, pairs would oftn ar widly disprsd, as fo Gvy's zra (quus grui) and th Asiati wild
sult. ass (quus hmioous), fmals of diffrnt stats ar prludd by mta.
"nan 1974) and ts (Bradury and oli onstraintsfrom foraging togthr.As a sult,mls ompt for trri_
ologyatully shapd fmal dis- torisalong trvling routs that tak fmalsfrom fding aras to watring
ptions of this modl wr supportd points (Runstin1986, 1994\'
lassistudy of Afian ntlops il- In flids, fmls rmain sparatdwhn food is sar and haitats ar
atdwith physiologial onstraints dnslywoodd ausin suh irumstansindividul pry an b aught
By showing how ody siz afftd and nsumd for omptitos an intrvn.In mor opn haitats and
:ivd and thn rspondd to th dis- whr oth pry and omptitors ar lrg arrd mulr mor numrous, oali-
J prdators or-rgrsslands, Jarman tions of fmals frm to help hold on to kills until thy ar ompltly d-
nrasd survival and rprodutiv vourd (Pakr 19s6). If ths fmal olitions ar thmslvs larg, thn
rdtht th smallst-odidspis, thris prssurfor mals to ggrgtto ontrol rprodutiv assto f-
<rs(phllpbus spp., Slvipr mals.Thus th only higly soial flid is th svanna-living|ton (Pnthr
klipspringrs (reotrgus oreotr- Ieo)' In this plntiful landsap th lopari (Pnthr prdus) rmains soli-
-quality vgtation. But baus of tary auseit an safly ah its lag pry in trs.
'padto widly sattrd. High For anids, monogamy is th rul. But variations do our: small-odied
;ks of prdation mad trritoriality foes (Vulps spp.) somtims hiit polygyny, and larg_odidhunting
s. airs gnrallylivd in wodd or dogs (Lrl pitus) arrd tirnr wlvs (is lupus) dvlop polyanlry
vgtationto grow and rnw itslf (ohlman 19s6). Typially anids nd hlp from norrrding
foagrs
:rritorial ownrship via snt rthr to hlp nurish latating mothrs. Fr th midsizd iakals. suh as th sil-
', thssmall-bodidspisdud vrak 1aka|(is mesomeld, th hlprs ar young from prvious lit.
f rnivorswould pry upon thm. trs tht annot thmslvs find sussful rding loations. For th
r physiologiallydtrmind ditary smallst spis, howvr, prpartum invstmnt in young is rlativly
amdistriutd on th landsap small for a fmal; thus hr mat provids all th hlp that is ndd. In
lbl antiprdator stratgy for largr yarswhn pry rsours ar vry high, omptition twn mothrs
and
ng groups bam th st stratgy thirsoon-to--funddaughtrs is low, and thir dughtrs ar not ford
l. Fortunatly,with largr siz also to disprs.Thy thrtrprovid nighorirrg mls with additionl mating
244 Soil Behuior d Soiobiolog
opportunitis, and polygyny rsults. For th largst-odid spis, how- or food rsours oftt
vr, fmal prpartum invstmnt is vry high, and thy nd all th hlp p r o d u d d i r i n sl i
thy an gt. To nlist th support of othr adult mals that must hunt o- failing to prdu an
oprativly to aptur larg pry' a dominant fmal not only kills th off- forming whn ondit
spring of othr fmals in th group ut also mats with thir mats so that But whn thy form'
ths mals hav as if thy ar th sirs of th dominant fmal's young. as many studis hav
Hn, dpnding on siz-dtrmindmtaoli investmntsand nds' so- If family omposit
ial systms of anids an vary from polygyny to polyandry, and thy som- hang. Baus los
tims us th srvis of juvnil or dult nonrdrs to hlp rar spring rarly attmpt
offspring. sx. Thus omptitior
As ths xampls illustrat, th omparativ mthod nfitd trmn- nitis rmains low. If,
dously y th dvlopmnt of soioologial thory that mphasizd th thn from the prsp
importan of fmals and thir rlationship to th nvironmnt in strutur- will likly produ .
ing soitis.But asptsof kinship and dmography also om into play. As mat and rmains in
amilton's rul illustrats, th strongr th dgr of rlatdnss among offspring of this pir,
rlativs, th mor likly altruists ar to nhan th rprodutiv opportu- dgr to whih th r
nitis of kin whil inurring osts assoiatd with diminishd prsonal r- ratd will dpnd on
prodution. Thus in th prvious xampls, it is not surprising to find that ship twn th tw
th oalitions that form among lionssswhn thy ar protting kills and prodution mong l
among male lions whn thy ar dfnding mating opportunitis with f- valu of 1 whn th r
mals ar formd most oftn among full silings (Pakr 1986); nor is it sur- whn rding is om
prising that th hlprs jakal adults kp to rear additional offspring are podution ours wil
thmslvsth full silings of th offspring ing raisd (Mohlman 1986). by dominants if suo
In gnral, strong kinship lowrs th thrshold for th apparan of altruis- nats if thy lav; (3
ti and ooprativ havior and may sustantially afft th osts of living and (4) th magnitud
in groups. rdrs. $hn oppo
Th strutur of jakal soitis not only highlights th importan of kin high, th dominan d
sltion ut also illustratesth fat that mny soitisonsist of adults of th nfits of having
diffrnt gnrations' som of whih ar for offspring that ar not nssar. breder will slt
ily thir own. any insts in th ordrs Hymnptr and Isoptra and (Rv and Ratniks 1
aout " of birds and mammals show srrh systms omposd of oopra- options lswhr and
tiv brdrs (mln 1997)' Up to this point th mrging synthti thory of damag on th sta
soiality has largly tratd all adult fmals and mals as indpndntdi- .!7rg
1992), Groov
sion makrs; rsponding t ologial onditions shapd th st rspons (Faaorg et a|. 1995),
for fmals, whih in turn onstraind th st rsponss of mals. But in 1994) have shown, do
many soitis mature offspring rmain and help parnts rar additional off. likly whn hlprs ar
spring. Ultimatly, undrstanding why individuals forgo prsonal rprodu- kin sltion.
tion to hlp los rlativs rprodu povids insights into how th alan But as might p
btwn dspoti and galitarin soial rlationships, as wll as th dynamis and laving disions t
of familis, is dtrmind y intrationsmong ology, kinship, nd domi. qustiond whthr d
nan (mln 1'995). groups (lutton.Brok
In most spis, offspring lav hom for rding. But in many soial ativ brding soitis
insts, primats, and othr mammals and irds, on sx rtins tis to par- tions asd on kinship :
nts, somtims prmanntly' rating multignrational familis. Gnrally, suh as mutualisms tha
offspring postpon disprsing whn rding opportunitis ar limitd ls- that mrg as mrka
whr. lgial onstraints assoiatdwith limitd rding sits, mats' om into play.
So il B ehtlior d So ci lbiollg z+\
: th lrgst-odidspis' how- or food rsoursoftn mak th option of stying hom and hlping parnts
:y high, and thy nd all th hlp produadditional silings ttr than disprsing arly and almost rtainly
rr adult mals that nrust hunt o. failing to produ any offspring. Thus marry family groups will unstal,
inant fmi not only kilis th off- formingwhn onditions ilre poor and disnding whn onditions good.
r l s o n r t sw i t h t l r i r m a t s s o t h t But whn thy form, intrgnrationaloopration should ommonpla,
:s of th dominant fmal's young. as many studis hav shown.
:taoliinvstmntsand nds' so- If family onrposition hangs, howv, thn soial dynamis an also
gyll to polyandry, and thy som- hang.Bause los inrding oftn has dltriousffts' rtirrdoff.
adult nonbrdrs to hlp rar springrarly attmpt to mat with a prnr or los rlativ of th pposit
s.Thus omptition with th sam-sxdparnt for rprodutiv opportu-
partivmthod nfitd trmn- nitisrmains low. If, howvr, a parnt dis and is rplad y a nonrlativ,
ogial thry that mphasizd th thn fom th prsptivof nondisprsingadults, nw mirting opportunitis
; h i pr o t h n v i r o n t n n ti n s t r u t u r - will likly produ onflit. Similarly' if a nondispsing adult aquirs
lmographyalso om into play. As mat and rmains in th group and limitd rsours ar onsuled y th
: th dgr of rlatdnss among offspringof this pair, thn omptition wi1l hightnd.In ithr as th
n h a n t h r p r o d u t i v o p p t u - dgrto whih th rprodutiv ativitisof th nondisprsingadult ar tol.
iatd with dimirrishd prsonal r- ratdwill dpnd on th stngthof th dominan-suordinrrtionrlation-
lls, it is not surprising to find that ship twn th two. Reproductiue she, desris th distriution of
i whn thy ar protting kills and rprodution among rding adults within familis and an rng from a
ding mating opportunitis with f- valuof l whn th rdrin harg garnrs all rprodution to a valu of 0
s i l i n g s( P a k rl 9 8 6 ) ;n o r i s i t s u r - whn rding is ompltly egalitarirr.Th dgr to whih sharing ot r-
)p to rar additional offspring ar produtin ours will dpend upon four onditions: (1)th rrfitdrivd
ing bing raisd (Mohlman 1986). y dominants if suordinats stay; (2) th rprodutiv suss of subordi-
sholdfo th app;rranof altruis- natsif thy lv; (3) th gnti rlatdnssamong potntial ordrs;
tstantiallyafft th osts of living and (4) th mgnitud of th domirran rlationshi btwn potntial o-
brdrs. \ff/hn opportunitis for th suordinat to rd lswh a
nly highlights th importan of kin high, th dominan diffnamong rdrsis small, kinship is low' and
t many soitisonsist of adults of th nfitsof having hlps to th dominant r high, thn th dominant
for offspringthat ar not nssar- rdrwill seleted to pay high sti?lg inntiu and p itttiz,
rs L{ymnoptra and Isoptrzr and (Rvand Rtniks 1993) to nsur that a hlpr hlps sin l-rlprshav
suh systmsomposd of oopra- options lswhr and, aus of narly qual fighting aility, an inflit
int th mging synthti thory of damag on th stalishd rdr. As studis on -atrs(mln and
ralsand mals as indpndnt di- Wrge 1992), Groov-illd anis (Vhrnamp 1978), Galipagos Hawk
:onditionsshapd th st rspons (Faaorget a|. |995|, and mongoss (Crl nd \Wasr1991; Kan t al.
th st rsponssof mals. But in 1994) ave shown, dominant rdrs making ..onssions'' to hlprs is
and hlp parnts rar dditiona| off- likly whn hlprs ar not los kin sine thy gain littl indirt nfitvi
ndividtllsforgo prsonal rprodu' kin sltion.
ovids insights irrto how th bln But as might b xptd, variants to this st of ruls that shp staying
.lationships,as wll s th dynamis and laving disions of hlprs our. lutton-Brok and ollagus hav
; mong ology' kinship, and domi- qustiondwhthr dominants hav full ontrol ovr rprodution within
gloups (lutton-Brok t al. 2000). Thir studishv shown that in oopr_
forbrding.But in many soial ativ rding soitisof mrkats, lvls of onssions do rrot fit prdi.
n d i r d s ,o n s r t a i n st i s t o p a r - tionsbasd on kinship and th nds of dominants' As a rsult, othr ftors'
nultignrtional familis. Gnrally, suhas mutualisms that nhan rprodution for vry group mmr and
ldingopportunitis ar limitd ls- that mgas mrkat groups gt lrg (lutton-Bok t l. 2002.), ma
l w i t h l i m i t d r d i n g s i r s ,t n a t s ' ominto play.
246 Soil Behuior nd Soiobiolog
a sukr dos tt
Additional Soial Glus b-product mutulis
of what a partnr d
As mrkats illustrat, not all soitisnd hld togthr y th glu of kin- Studis on a varit
ship. Rlationships uilt around oopration rathr than altruism or asd Long-trm studis of
on mutualism and riproity an hav just as strong an fft and provid Pakr 1991) and St
th opportunitis for diffrnt typs of soitis to volv. Rlationships ar y adopting individu
uilt upon rpatdintrationsthat an involv omptitivand ooprativ ilarly, strong rlation
lmnts. Manipulating th natur of prsonal rltionships and thos of fantiidal mals or
ompanions should also dtrminth xistnand haratristisof many prdtors providd si
soitis. ing alon.
Trivrs (1972a) Was th first to suggstthat riproity ould play an im-
portant rol in shaping soitis.If individuals Wr to aid a strangr y in-
rasing its survival or futur rprodution at som ost tO slf in trms of Sial Variants al
diminishd rprodution' thn suh havior ould only maintaind in a
population if th ripint wr to vntually rpay th osts inurrd by As th xampls of fa
th altruist. This onpt has n formalizd as th prisoner's dilemm, dnsity and th intn
whr two individuals hav two altrnativs-ooprat or dft. utual trritoris with suffi
oopration, usually y staying tru to ah othr dspit attmpts y au. rsponsibl for favori
thoritis to ajol on Partnr to tufn on th othr, provids a highr r- of hlprs. Thus dm
ward (R) than if thy both dft and ar punishd /P/. If only on partnr sx ratio and ag stru
turns on th othr' hwvr, then th siz of th tmptation (T) nssaryto vitl ats, will shap
rak th partnrs' oopration nals th hat to do st of all. on- spis variation as
vrsly' th partnr who rmains ooprativ to th nd is th sukr and pl, wr ag spifi
rivsth lowst rward (S/.As long as payoffs ar ordrd > R > P > s, thn high rding s
dfting is favord sin it provids th gratst short.trm nfit. If, tionships would gn
howvr, suh situations our frquntly and th nmr of individuals in of motality Wr r
a soity is small, thn pairs of individuals will find thmslvs intrating ommon. Sin som
tim and again. In suh a stting, dfting is no longr th st stratgy. th variation in anid
Rathr, oopration dos bst' and th tati tht oftn provids th grat. uls an hav importa
st long-trm rWard is tit-for.tt (TFT; Alrod and amiltorr 1981)' in Th synthti modt
whih intratants ooprat initially and ontinu to do so until a partnf nvironrnntal fatur
dfts. If dftin ours' thn th othr partnr dfts and ontinus to mals. In many ass1
math th havior of th initial dftor. If th initial dftor again oop- systm of soial org
rats' thn th rspondr is instantly forgiving and ooprats as wll. TFT onditions. In quids
an rsist invasion y a stratgy of pur dftion. But sin it is qually as transitory groups wh
rwarding as a stratgy of always ooprating, TFT an suprsddby populations mov fr<
suh a stratgy via drift. Sin a purly ooprativ stratgy n invadd hang. As was foun
by th slfish stratgy of dftion, th stratgy of tit-for-tat annot truly Stats, whr food an
an SS, ut it oms los. diffring nds an
Th logi of this argumnt suggsts that struturing soitis on rl. aras ar ford to s
proity alon will diffiult. Kinship an always provid a strong foun. spond y dfndingtl
dation sin it rdus th nd of fully rpaying dlayd nfits. By and harmlike soiti
oopating mutually to solv ommon prolms, partnrs may nfitto Soial variation' ho
suh a lrg tnt that th tmptation to dft rmains low nough that Variation in sial str
Socil Bhuior d Soioliolog 247
a s u k r d o s t t r t h a n a d f t o r . . W s t - r h a r d( 1 9 8 9 ) t r m s t h i s
b-produt mutulist us oopration is univrslly st rgardlss
of what a partnr dos.
hld togthry th glu of kin- Studison a varity of spissuggstthat mutualism an shap soiality.
t rirthrthan altruism or asd Long-trm sttrdisof lions y Pakr (Pakr and Ruttan 1988; Shl and
as stongan fft and provid Pkr 1'991')and Standr (1992) showd that only y hunting togthr and
:isto volv.Rlationshis r by adopting individul rols wr lions liklv to ing down lrg pry. Sim_
l v m p t i t i v n d o o p r a t i v ilarly,strong rltionshipsamong fmalsto prott us from attaks y in-
lnal rlationshipsand ths of fantiidl mals or to hold on to pry in th fa of othr marauding
n and hartistisof mny prdtorsprovidd simultanous nfitsthat old not obtaind y at-
ing alon.
l r t r i p r o i t y u l d p l a a n i m -
lls wr to aid a Strangr y in-
at som ost to slf in trms of Soial Variants and Altrnativ Stratgis
r ould only maintaind in a
lly rpay th osts inurrd y As th ampls of fmilis with ooprtiv brdrsshow, high population
zd s the prislr's dilmm, dnsity and th intnsifid omptition it ngndrs in th sarh for sr.rital
s_ooprat or dft. utual trritories with suffiint food or haital urrows is th fator ultimatly
h othr dspit attmpts y au- rsponsilfor favoring th stalishmntof oalitions nd th rruitmnt
,h othr, pror,ids highr r- of hlprs.Thus dmographi ftors, suh as popultion dnsity, as wll as
unishd /P). If only on partnr sx ratio and ag stutur that rsult frorn diffrns in phenotyp-spifi
th tmptation (T) nssary to vital rats, will shap pattrns of soial organization and lad to within-
hat to do st of all. on- spisvariation as ologial irumstanshang. If mortality, for xm-
iv to th nd is th sukr and pl, wr ag spifiand high for prrprodutivfmals than for mals'
ryoffsar odrd T> R > P > s, thn high rding sx ratios (mals/fmals)wuld rsult, and mating rla-
gratstshort-trm nfit. If, tiorrshipswould gnrally om mor polyandrous. If s.spifi pattrns
nd th numr of individuals in of mortality wr rvrsd, howvr, thn polygyny would om mor
w i l l f i n d t h m s l v si n t r a t i n g ommon. Sin so of ths nrortality onrns wr also rsponsil for
g is no longr th st stratgy. th vriation irr arrid soil strutu dsrid arlir' ths mortality shd-
'i that oftn prvids th grat- ulsan hv irpotant onsquns.
lrod and Hamiltorr 1981), in Th synthti modl of soiality that is mrging shows that a varity of
:ol-ltinu to do sl until a prtnr nvironmntalfturs influn th pattrns of soility xhiitd y ani-
p a r t l . l d f t s n d o n t i n u s t o mals' ]n many ass ths pattrs ar flxibl, and spis an vary in th
[ h i n i t i ld f o g a i n o o p - systmof soial organization thy hiit dpnding upon nvironmntal
,ing and oopratsas wll. TFT onditins. In quids' for ampl, although fmal asss typially liv in
ftion.But sin it is qually as transitorygroups whos mmrshiphangs (Runstein1986,1994) wen
ting' TFT an suprsdd y populations mov from arid to msi aras, soial rlationships an also
prativstratgyan invdd hang.As was found on an island off th ost of th southastrnUnitd
:gyof tit.for-tat annot truly Stats'whr food and Watr ar aundant and los togthr,fmals with
diffringnds an oals into prmannt groups. als, whih in arid
a t s t r u t u r i n gs o i t i s o n r i - arsar ford to stalish trritoris along routs to and from watr, r-
r lwys provid ir strong foun. spond y dfnding ths groups muh lik mals of horss, thir los kin,
' rpaying dlayd nfits. By and harmlik soitismrg.
'olms'partnrs may n{it to Soil variation, howvr' is not limitd to diffrnsamong popultions.
dftrmains low nough that Vrition in soial stratgisan volv within populations sin not all indi-
248 Soil Behuior d Soiobiolog
orintation, groups ould mov from disorganizd swarms to highly polar. is no on animal
izd onformations. In fat' strngth of alignmnt playd a ritial rol in un- grs y oopra
drstanding ladrship, information transfr' and onsnsus dision making thos of most t}
within groups vn whn informd individuals did not know th quality of and mor variti
thir information or whthr thy wr in th majority or minority (ouzin diffus. But this .
t al. 2005). not apply to thos
nssarily invali<
on of th mos
From Animals to Popl an plain a n
rothr's phnorr
Humans ar animals, so it sms slf-vidntthat volutionary prosss posd to thir ov
should shap human soiality. First prinipls, spiallythos that undrli strat8y appars
th synthti modl of soiality in nonhuman animals, hav stimulatd and qual, a parnt sh
organizd th study of human soial havior. Not surprisingly, onntions as with a sibling's
among partiular faturs of soial volution and partiular aspts of hu- spially tru in
man natur hav n idntifid. Nowhr is this mor apparnt thn in th (Flinn 1981; Iron
ara of mat hoi. In rodnts gns assoiatd with th majr histoompat- rih' and as a rst
ibility ompl (MH) afft ll rognition and immun funtion. During rais their hildr
strus fmals assoiat with mals that diffr gntially from thmslvs' philandring is hi
whras whn thy ar prgnnt, thy assoiat with fmals that ar gn- is ommon' and l
tially similar. Assssmntis via odor and suggststhat MH-speifi assoi- vn if offspring i
ations ar volutionarily adaptiv. Divrsifying th gen pool of offspring sister's hildrn t
should improv thir immun funtion, and assoiating with los kin during ship thory, mn z
prgnany should rdu osts and inras nfits assoiatd with soiality th st of a ad j
.sfdkind
(Pnn and Potts 1999). Whn and ollagus(1995) prformd a Applying st.t
study to s if human fmals showd similar sual prfrns, h found man soial hav
that womn in th frtil phas of thir yl idntifid th odor of T-shirts biology sought to
from mals with whom thy shard H gnsas lss ..pleasant''than thos havior of isti
of mals with whom thy shard fwr gns. Apparntly th sam undrly- hop was sltiv
ing fors and mhanisms mployd y animals may oprating in hu- that had opratd
mans whn it oms to hoosing mats. But y no mns do ths rsults likly to tru' i
dmonstrat that prosss that shap th soiosxual havior of animals for idntifying h
will nssarily shap th havior of humans in similar ways. Unfortunatly, among and tw
th links twn animals and popl ar oftn prsntd as univrsally tru most animal soi
whn in fat thy show variation dpnding on th natur of sltiv fors. losly to thir n
By ignoring th ontingny of sltion, laims of univrsality of human so- als an mas
ial institutions and thir immutal natur ar likly to fals. As many of Prhaps spur
th ampls of animal soiality plord rlir hav dmonstratd, partiu- psyhologists lso
lar typs of soial rlationships and soial systms thriv only undr a limitd prsptiv, but t
st of nvironmntal onditions. Why, thn, should W xpt that only on sought to idntify
st of human rlationships or only on typ of soity should volv? lutionary tim. Fo
It is somwhat ironi that politial thory itslf is a human onstrut with anisms tht our
ah prediatd on a diffrnt st of assumptionsaout human natur. on- ostraizing heats
srvativ philosophis oftn dpit humans as fundamntally slfish, ompt- solv suh prol
itiv, and somwhat invariant, whil mor liral philsophis oftn assum usd on rprodu
that humans ar flxil and adaptabl.As is th s with all animals, thr mhanisms that
So iI B huior d Soio bioll91, z . \ I
rganizd swarms to highly polar- is no on animal natur. Aggrssion and dominan ar offst to varying d-
]nmntplayd a ritial rol in un- grslry ooperation and altruism. What sts human soitis pat from
r, and onsnsus disilrrmaking thosof most thr animals is thir ornplity. Mo lvls of orgenizatiorr
Juals did not know th quality of and mor varitisof rlationshipsan form, thus making intrations mor
th mjority or minority (ouzin diffus'But this dos not man th ruls that strutur animal soitis do
not apply to thos of humans. Prhaps thy ar insuffiint,ut th ar not
nssarilyinvalid.
on of th most omplling ampls whr volutionry uls of soiality
an plain novl st of human rlrionships is th so_alld mothr's
brothr'sphnomnon, in whih mn invst in thir sistrs' hildrn as op-
.idnt tht evolutionary posd to thir own hildrn (Alxandr 1979a, 1979)' At first glalr this
Prosss
pls,spiallythos that undrli stratgyappears to violat th most si ruls lf kinship. All ls ing
mn animals' hav stimulatd nd qual,a parnt sharson vragtwi as many gnswith its own offspring
v i o r .N o t s u r p r i s i g l y' o l ] n t i o n S as with a siling'soffspring. But in som soitisntlt all ls is qual. This is
.Wst
tion and partiular aspts of hu- espially tru in marry Afrian soitis, suh as that of tlr Tongir
is this mor apparnt than in th (Flinn 1981; Irons 1981). Th land upon whih ths hortiulturalists liv is
i t dw i t h t h m j o r h i s t o o m p a t - rih, and as a rsult, fmalsn driv nough rsoursfrom thir laor to
tion and immun funtion. During raisthir hildn witlrout th aid of thir husrrds.In suh ulturs fmal
diffr gntiall from thmslvs, philandringis high' and fmals an initiat divor. As a rsult, ukoldry
soitwith fmals that ar gn- is ommon' and mn ar not likly to faths of thir putativ lrildrn.
suggststhat -spifi assoi- vn if offsprir.rgar only half sis, mn will mr losly rlatd to thir
'sifying th gn pool of offspring sistr'shildren than to thir own ptativ offspring. Thus, aording to kin-
rd assoiatingwith los kin during ship tory,mn r havingvolutinrily rationlly sin thy ar making
s nfitsassoiatdwith soiality th bst of a d job and invsting in th losstgnti rltiv vilal.
and ollagus(1995) prformd a Applying ost-nfitand gn-aountingprinipls to th study of hu_
rnilar sul prfrns' l.rfound mn soil hvior is th ornrstonof soioiology. uh of rly soio-
:yl idntifid tlr odor lf T-shirts bioiogy sought to plain humar"rity's soial roots nd thus studid th soial
..plasant''than thos
) gnsas lss haviorof isting ultures minimally impatd y lrVstrnsoitis.Th
ns.Apparrntly th sam undrly- hop was sltivprssursthat urrntl1,opratd Wr th sm as thos
l animals may oprating in hu- that had opratd in tlr past. Although th assumption of ontinuitv is un-
But b no mans do ths rsults likly t b tru, it dos not invlidat th undrlying soioiologil sarh
:l soiosxualhavior of arrimals for idntifying how nvironmntal prssurs shap htrnran rlatitllrships
rans in similar ways. Unfortunatly, among and twnss and within and twnfamilis' Morovr, s irr
: oftn Prsntdas univrsally tru most animal soitis,individuals in ths non-!strnizdsoitisar tid
ing on th natur of sltivfors. ioslyto thir nvironrnt.hus logil, soial, nd podr-rtiv vri-
laims of univrsalityof human so- alsan rasurdto idntify pattrns of ustion.
rr ar likly to fals. As many of haps spurrd on y th Darwinian invstigation of human soiality,
Larlirhav drnonstrtd'partiu- psyholgistsaiso ga nrirringhunrn soial havior from an daptiv
l systmsthriv only undr a limitd prsptiv'ut thir approah was fundamntally diffrnt aus th
rn,should We pt that only on soughtto idntify how natural sltionshapd th human psyh Ovr vo-
.pof soityshould vo[v? lutionarytir. For psyhologists,th r.rltimatqustion ws, ,.Ar th mh-
ory itslf is a human onstrut with nisnrsthat our rains us to mak disions aout hoosing lnats or
mptions out human natur. on- ostraizinghats th ons that w woul] pt if thy wr .dsignd' to
ns s fundamntallyslfish' ompt- s o l v s u h p r o l m s ? ' ' ( B a r r t t t a l . 2 0 0 2 ) . T h u s w h i l s o i o i o l o g i s t sf l -
r liral philosophis oftn assum usdon rpodutivoutomSto masuradaptation and oftn igrrordth
As is th as with all anirals, thr mhnismsthat undrli assssmnt and dision making' volutionary
2.'2 Socil Bhuior d Soiobiolog
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(t h lIogi 20: 4 10-4'].].
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\.ctRsrb d Ntu Thor' Biolog 46z 5-57.
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197 4. P arent-offspring onfl it. mri Zoolo gi st |4: 249_264.
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d uoltil of Behuiour, 11'9-141. 4: 1-33.
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.,T. Sk,
F. Bttns,
andA. J. Ppk.
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)rtionin rrids.In D. I. Runstin
spts lf Soil t.,uolutil:Birds d
ton UnivrsitPrss.
g systmsirr irds and rarmls.
th human lin, and Hakl (1s6s) imagind and namd a spifi sris of
forms from aplik to humanlik.
uriously, thir ap-man thory (as it was latr alld) was sriously hal-
lngd during th first svn dads of th twntith ntury. Ths hal-
lngs inludd th viw tht th last ommon anStor w shard with th
n animal world rsmldth tarsir mor losly than any ap (Jons 1918,
|926, |929). other arly twntith-nqurysintistsronstrutdth last
ommon anstor as mor monkylik than aplik (Straus 1949) or as som
..pr-ap''without long fingrsand uppr.ody spializationsof liv-
kind of
ing aps (.g.,Napir and Davis 7959I. Bausmodrn humans appar to b
so divrgnt in shap and bhavior from othr nimals, th tim of divr-
gn of th human volutionary linag was oftn givn as quit anint.
..ap.man'' thry ontinud to hav spPortrs
Th nintnth-ntury
(.g.,Grgor \9 |6, 1926, 7927a, 1927, 1'928,19 41 sashurn1 95 1 ), ut
lows rasonalonfidn in r- until th last thr dads of th twntith entuy ths wr rlativly
rord dmonstratsthat humn- lonly vois (Flagl and Jungrs 1982).
/ars ago and aumulatd mor Among th many important hangs in viw and mthod in human volu-
r i m .A s D a r w i n a n d h i s o n t m - tionary studis during th last fw dads of th twntith ntury was th
ws lar vidnof dsnt with appliatin of molular iology. Immunology providd th first insight. By
ion of this is th pansion of th omparing th strngth of th immun systm,s rspons y on spis to
s that wr larly humanlik in proteinsof anothr spis,Goodman (1963\ quantifid an stimatof gn-
3o had ap-sizdbrains that wr ti rlatdnssamong primat spis. Th rsults wr shoking at th tim
ans.Just aftr2 million yars ago to most omparativ anatomists.Contrary to prvailing undrstandingat th
rlf th siz of Homo spis, and tim, humans wr mor similar to Afrian aps (himpanz and gorilla)
L rain siz appars in th fossil than Afrian aps Wr to th Asian grat p (orangutan). Monky spis
r s a l l2 0 0 2 , 2 0 0 3 , 2 0 0 5 ; D l s o n wr far mor distantly rlatd. Mor sophistiatdimmunologial mthods
(Sarihand.!ilson 1966) and DNA hyridization (Hoyr et a|'.1964) con-
rtionistswr knly intrstd in firmd ths arly rsults and rvald that hangs in som maromoluls
.Wilson
i n i n t r p r t i n g o m p r t i v m o - (.g.,alumin) wr rlativly onstant through tim (Sarih and
v' vry fw prmodrn human fos- 1967).onstany of hang prmittd stimatsof tim of divrgnof vo-
63). By th turn of th twntith lutionary linagsas long s th rat ould aliratd y a known dat of
..molularlok.''
tan'' ntrdth onsiousnss of divrgnand thus provid a asis for dvlopmntof a
'Wilson
l turn of th twnty-first ntury, Sarihand (\967) usd th dat of 60 million yars for th divrgn
l thir way into th world's mus- of svral mammalian ordrs to alulat th rat of hang in alumin.
had grown sptaularly (Dlson From this thy alulatd that th divrgn of th linags that ld to hu-
7wartzand Tattrsa||2002. 2oo. mans and Afrian grat ps ourrd 4.2 million yrs ago. Susquntr-
; of th rord is du to th intlli. finmntspla th dat somwhr twn 10 and 5 million yars y most
rs who find th fossils and dtr- urrntstimats(Kumar and Hdgs 1998; Kumar t al. 2005; Holms t al.
\LS. 1989;hn and Li 2001; Arnason t al. 2000; Pilam and Young 2004;Pat-
trsont a|.2006; Glazko and satoshi 2003; Stauffr t al.200]). !7ith
th advnt of DNA squning it is now lar that humans ar vry losly r-
latdto th Afrin grat aps)and th tim of divrgnof thir volution.
ary linagsis rlativly short from th prsptivof mammlian volution.
rristhat popl shard a ommon A sond important hang in th latr prt of th twntith ntury was in
rn with any othr animal. Thomas th mthods of omparing morphology of spis. Molular approahs
Ldvoatfor an aplik anstry of annot applid to fossiis muh oldr than 70'000 yars (Kraus t al.
l 1. Speis,dats,ody size, rain siz,nd postriotoot siz in arly hominids
;
65
95 294
:Y
rrori tugesis 6 +l lJl 329
Ardip itb eus kd|| 5.2-5.8 .)/
u strlop i tll us msis 4.z_.9 ; 428
A ust r l op it h t's [r nsi s 3.9-3.0 45 29 151 4; 4; 460 2..4 1.7
us trlop it ll cus fri nus .0_2.4 4l 30 18 115 452 448 516 2.7 2.
Prnthropus ethiop ius 1-7 11
41 47 688
Prthropus boisi 2'_I.4 49 t+ 17 r24 521 514 756 2.7 2.7
Prnthr ops rt:lbustus 1.9-1.4 40 .)z 132 110 530 523 588 3.0 2.2
Astrlopitbes grbi 2.5- 4s0 446
Hrllllo bilis 7.9-r.6 -)/ ; 131 100 612 601 478 .6 1.9
Hlmo rudolfesis 2.4-1.6 60 51 160 150 752 76 572 -1.1 1.5
Homo ergstr 1.9-1.7 66 56 180 160 871. 849 - ) // 3. 0.9
Homo eretus 1.1-0.01 66 56 I,050 1,01.9 402 4.0 1.0
Horo bidelrgesis o.6-0.2 77 56 1,\94 \,156 389 4.2 0.9
Late H omo n.ldrtb.llsis o-2-o.o3 77 66 ,t o. -/ .l
7.62
58 7,414 33-5 4.7 0.7
.l
omo floresisis 0.09-0.01 29 106 3B0 78 2.6 .8
Homo sIpies xtant 58 49 t75 16,|' 1,350 4 s.8 0.9
1. Taxtlnlmysdon Klin (1999)with dditionsfronr Brunt t al. (2002),Snutt l. (2001),Brown t al. (2004)'and Hail-Slssi t l. (2004).
2. l)ats (in millilns<lfyrs g) r from KIin (1999),,ith additilnsfrom Brunt t al. (2002),Snutt al. (2001),Brrlwn t al. (2004) and Hil-Slssi
t l. (2004)'
. Bod mss stimts(in kg) ar flmHnr (|992-)eptfor th fol|wing:. tugsisis fr<lmNktsuksat al. (2007)'whr 41 is th rrrid-
point <lfthir rang and thy did not spifys; . msisml is from Laky t al. (1995);. lsisfmalis lulatdfrom th rtio of ml and
fml in . frtlsis;. rgstr'H. rts,H. heidlbrgzs,and |ate H. drthIsisr from Ruff et al. (1998); and H. florsizsisis from Brwn t
l. (2004)
4. Stturstimts(in n.r)ar from Hnr (991) pt for . tugeesis'whih is from Nktsuksat al, |207)'whr 13l is th midpoint of thir
.Walkr
rng and thy did nlt spify s . rgstr,whih is from Ruff nd and lat . drthlsis'whih r
(1993); H, rtus,H. hidIbrgesls,
fronl Ruff t l. (1998);and H. |lorsiesis, Brown t al. (2004).
whih is frln.r
5. V is ranil paityin fror sourslistd in MHnry (1994a;1994)u,ith th addition of S. thdsls(Zollikofr t 1.2005);,L.444-2 (540
) to . frsis(imbl,rsonlommuntatin);Stw 505 (51.5) to . fl.ius(lnrlyt t. 1998);P. thi<.lpius (Walkt al. 1986);KGA 10-525
(545 ) to P. bisi (Suw t a|. 1997|; and BOU-VP-12/1 0 (4.50) to . grhi (sfaw ct aI. |999). ' rtus,H. hidlbrgsls,and |ate . dthl-
sis ar from Ruff t 1.(1998), and H. |loresizslsis from Brlwnt l. (2004).
6. Brin wight is alulatdfrom formula (6) in Ruff t al. (1998).
7. Postanintoot ar (in mm2)is th sum of prdutsof bual-lingualnd rsio-distllrrgthsof l,., ', r"rdM. irnd is takn from MHnr |1994)
with th ddition of S. tchdsisstimtdfrom TM266-02-154-1 reportdin Brunt t l. (2002);(). tugsisfrlm Snutt al. (200.lI;' Ar. kdbb from
il-SIssi t al. (2004); r. rmidus frtlmlihitt al. (1994); . msisfrom I,kyt l. (1995); H. H.
rtus, hidlbrgsls,nd lat ' n-
drthIsisfrorn Wlpoff (1971,1982I; and H. |lorsisis from Brown t al. (2004).
8. Q is th nphlizationquotintalulatds ritmss dividd y (11.22xbody mass().'6) from Mrtin (1981).
9. MQ is th mgadontialquotintdrivdas ostnit-t tooth ar dividd by (12.1.5bodymass()86) from Hnry (1988).
260 Hum uolutil
Bfor Hominir
ll0mo
reus
our los gnti r
{n'.,,./
origin of our volul
r"l
natly, th fossil r
panzs' and humar
bttr known. Disr
rvaling. In sdim
forms appar that ]
(Pilbam 1982). Siu
uniquly orang-lik
profil of th fa t
faturs that among
Th fa of th As
th Afrian lad (g
(flakedston of ths living sp
tifds may not' y th logi,
unknown)
ost palontologis
qualify uropan f.
ago), ouropitb,
(1'2.5_| million y
--\
as part of th Afri
Sahelanthropus
thadensis Afria for ]\4ion
spars' but dilignt
Figur 1. A working hypothsisof volutionryrlationshipswithin th human pitheus (16 to 14 r
fossil rord,rdrawn from R. G. Klin. 15 million yas ago
umn uolution 261
4y is th only guid to undrstarrd- lins of dsnt. Homoplasy is prvasiv in human volution (Hnry
ionshipsan infrrd from simi- 1996|.Becaus Hnnig's mthod fouss on th haratristis of volution-
.rotwitlrotrt arful arralysis of tlr ary divrgn, it is oftn rfrrd to as ldistis' ld in this voaulary
logistshv alwas n awar of is a uniqu volutionary ranh f th tr of lif d6nd y shring uniqu
l mak omparisorrsof form only traits that ar not shrd in losly rlatd ranhs. nnig alld suh
1 ntry. shard uniqu traits ..synapomorphis,'' whih is usually translatd as
..shard
roah to omparing morphology of drivd.''
rr-rdid not nssarily imply los Th powr of this logi pvds volutionary iology (Hull 1988) and
u to sharing priritiv haratris- hlps ronil th findings of molular iology with th intrprttion of
liologial lationships.Aspts of th fossil rord. As a rsult, th trms usd to lassify our biologial rlativs
now rflt our tru gnti rlationships. Th old tradition plad grat aps
"olioithcus, for ampl' rsm-
.osof nrodrn aps (Kurtn 1972), (orangutans'gorillas, and himpanzs) into th family Pongida and hu-
stral trits and not to th volrrtion mans into on-rirrida'but th tu gnti affinitis for a rlassifiation
lts and not prsnt in othr losly that rognizs th Asian grat ap as th family Pongida and th Afrian
: du to paralll or onvrgnt vo- lad as th family Hominid. \/ithin th family Honrinida ar th su.
l nsto.Th wings of irds' ts, familis Grillina (gorilla), Panina (himpanz), and Hominina (hu-
ausr thy volvd from th s:rm mans).It is a usful rmindr that w ar tightly nstd in th Lrshof lif's
usd to dsribth phnomnon oganiforms and that th origin of our uniqu linag is vry rnt.
v o l v di n d p n d n t l yi n s p a r a t
BforHminins
ll0mo Artifat raditions
f/oresiens 1"
,t". st"'* g"
la .
J Upper PaIeo|ithi
our los gnti rlationship to th Afrian grat aps implis an Afrian
n(........-
[ )i - iddle stone Age & origin of our volutionary linag. Fossil disovrisonfirm this. Unfortu-
ff::'|'
\j!g]g|g!gg]i. natly' th fossil ord of th origin of th Afrian lad (gorills, him-
A [-) panzs'and hunrans)is poorly doumntd.Th Asian lad of our family is
ffi ."uteanffi ttrknown. Disovris in th foothills of th Himalyas ar partiularly
tt 13
rvaling.In sdimntsthat dat twn aout 12 and 8 million yars ago'
v
,.'"'r'i,
lParanthropus 1 . 6 5-
forms appar tht rsml tlr ofangutan in mny spifi faial faturs
|
J,.'",|^
d7\ oldo*n fl) (Pilam 1982|. Siupithcus is th gnus nam that inluds fossils with th
\_)/ \,
uniquly orang-lik faturs of narrow spir twn th ys, ..prowlik',
2.5
profilof th fa low th nos' onstrution of th palt, and matly othr
fatusthat among th living spis our in th rnguta n (Pongt)'
Th fa of th Asian Iade (Siupithelts and Pogl|ontrasts with that of
,,.^".:*,*, th Afrian lad (grillas,himpanzs,and humans), ut th rsmlan
, of ths living spis may du to sharing primitiv haratristis and
( f | a k ds t o n e
theUs atifts may not' y th logi of ladistis, usful in dtrminingtru rlationships.
unknown)
ost palontologists apt ths rsmlns as suffiintly r"rniquto
qualify uropan fossil gna suh as Dropithus (8_13 million yas
I-'
.
ago),Ournopitheus (9_10 million yars ago), and prhaps Pirolpithes
l.
(12.5-1million yars ago) (Moyi-Soli et a|' 2004; Bgurl and rwrd 2005)
as part of th Afrian lad (Bgun 1994;F|eag|e 1999). Th fossil rord in
Afria for ion mmrs of Hominida (Afrian aps and humans) is
spars' ut dilignt attmpts ontinu to promis nw disovries' en.
within th humn
nary rltionships pitbus(16 to 74 million yas ago) and rlatd gnra (qutoris, 16 to
i5 million yars ago) appar to part of th Afrian lad with living nrm-
262 Humn ulllttio
brs, ut ah has its own spializationsthat mk thm lss attrtivas for sltion that fav
andidts for anstry of livirrg spisof ominid (rossin and Bn- g r s .A n r h h o m i n i
t 1997\. of th formn lltag
T.h palonvironmntwithin whih our nstorsvolvd was inflund howvr, Sbeltbr
profoundl y drmati flutuationsirr world limats ovr th past 7 millio t h h u m n l i n a g .I
yars (Vra et a|. 19951Bromag and Shnk 1999; Potts 2007; Pikford mans. It has normou
2006; Bhrnsmyr2006).Intnspriods of old tmpratursin highlati. ps or arlir mm'
tuds rsultdin pisodsof drought in th tropis' Thes apprd:rt16_13' having n Issdv
6.5-5, nd 2.8 million yars ago' and ginning at 2.5 million yars ago' th h disovr of S
arth prind21 yls of glaiation in highr lvations and ltituds n s so f i h l B r u n t
aompanid y rdution in rainfall in quatorial lands, spiallyAfria. Chadian studnt who
Haitats fvord y hominids (anstorsof th Afrian aps and humans) Afrian Rift, whr
ontratd and xpandd. Th fossil rord of hominids in th iddl nd minds us to mind{l
Lat ion is frustratingly inomplt, and almost nothing is known of th Almost all of th .
linags tht ld to himpanzs and gorillas. Suprisingly, muh more is omes from th Afri'
known of th fossil rord of our own rnh, th Hominina. vnts'many lt i<
mnts layrd y vol.
nrurally ouringp<
Th arlist Hominins of ash in thiopia, K
th arly part of th s
Rogr-ritionof th rlir mrrsof Homirrina is guidd by undstanding ourrd.
gaind from moltrlr iology and ladistis.Th molular lok rakets Th fist haptr in
th im of divrgn somwh twn 10 and .5 million yars y most to just lss than 6 mill
urnt stimats.By th logi of ladistis, th arlist astor of humans 2001; Pikford et a|,2
an rognizd y th prsnof traits uniqu to th linag.This is ob- this spis omind '
vious ut is asily onfusd y naiv viws of pr-Hnnigin omparativ appar to adaprd
morphology that ontinu to plagu human volutionary studis.If it looks J u n g r s2 0 0 8 ) .
.W.
lik an ap' it is not nssrily ludd from th human ralm. now Slightly latr in tim
know f n traordinary rord of an aplik-oming-lrumanlik s- thiopian spimns rt
qun of fossils.Th arlistmmr of this squnlolks vry aplik.By Gabril t a|.200l; H
ldisti logi, that is atly what on would pt. with rrori tugs
Th arlist andidat for mmrship in th sufarrrily Holinina most rspts,ut it d
Shlnthropus thdsis (Brunt et a\. 2002). This prious fossil skull ths is anin tooth
frorrr had was found in sdirnntsthat ontilind funa linkd with databl sil or modrn aps an
sdimnts in thiopia nd North Afria 7 to 6 million yars ago. Its ovrall the ability to hyprxtt
apparan is quit aplik' inluding an stimatdrain siz slightly low It livd in a losd wo
th avrgfor living himpanzs (Tal 1). But y th logi of ldistis,it
sharswith latr hominins uniqu taits that dfin our linag.Th mostim-
potant of ths is th anir-rthat, unlik that of ps and primats in gn- Austrlopithecus i
rl, dos not projt or war against its lowr prmolar. This is a signifiant 4.2-2.5 Nlillion Y
fatur tht ws rognizd as an important diffrn twn aps and
popl y many, inlding Darwin \|872\. h fssil vidn f
Small nins ar signifiant.Darwin (|872) notd that whn human n. Sh lt brops t h di
stors first stood rt' thy frd thir hands and ould randish wpns yars ago thr appar
fo thrat nd fight, thus no longr rlying <rnlarg anins. Small nins cus nlnlnsls that pos
arry spial signifian,ut it is not too diffiult to imagin oth rasons ( L a k ye t a I . 1 9 9 5 , 1 9
Humn uolution 26
that mak thm lss attrativ as for sltiontht favrs rdution to rliv th xpns of growing suh dag-
Hominida (rossin and Bn- grs.Anothr hominin novlty sn in S. tbdensis is th forward plamnt
of th foramn magnum' In ovrall morphlogy (Zollikofr t al. 2005),
r anstorsvolvd ws inflund howvr, Sblthroprzs is aplik, as rfits a spis los to th origin of
rld limatsovr th past 7 million th human linag.Its rain is minusul omprd with that of modrn h_
hnk 1999; Potts 2007; Pikford mans.It has normous brow ridgs that r vn largr than thos of modrn
s of old tmpratursin high lati- aps or arlir mmrs of th gnus Homo whr row ridgs appar aftr
j tropis.Ths apprred at 16_13, havingn lss dvlopd in arlir spisof our linag.
nning at 2.5 million yars ago' th Th disvr of Shlthropus is du ttl the prsistn,skill, and lvr-
in highr lvations and latituds nssof Mihl Brunt and his tam' inludig Djimdoumaay Ahounta' th
:quatoriallands, spilly Afria. hadian studntwho found th skull. Its loation 2'500 km wst of th st
of th frian aps and humans) Afrian Rift, whr almost ll th othr alist hominins wr found, r-
rd of hominids in th iddl and minds us to mindful of th limitd tnt of ou smpl.
Lndalmostnothing is known of th Almost all of th vry arlist fossil rord of our volutionary history
rillas. Surprisingly, muh mor is oms from th Afrian Rift Vally. Baus of xtraordinary gologial
nh, th ominina. vnts' many lat Mion to Rnt trrstrial fauna ar prsrvd in sdi-
mnts layrd y volani ashs that an datd by th radioativ day of
naturallyourring potssium-40 to argon-4O.Fossils urid bnath layrs
of ash in thiopia, Knya, and Tanznia povid most of th vidn for
th arly pat of th stoy of human volution arrd th ontet in whih it
nininais guidd by undstanding ourrd.
tis. Th moiular lok rakts Th first haptr in this story ols from fossils nath ash layrs datd
.n 10 and 5 millin yars y most to just lss than 6 million yars ago in th Tugn Hills of Knya (Snut t al.
:s,th arlistanstor of humans 2001; Pikford t al. 2002; Sawada t al. 2002). Namd rrorin tugesis,
s uniqu to th linag.This is o- this spisomind aplik tth and forlims with humanlik thighs that
w s o f p r - H n n i g i a n o m p a r a t i v appar to b adptd to ipdality (Pikford t al. 2002; Rihmond and
an volutionarystudis.lf it looks Jungrs2008).
.W
l from th human rirlm' now Slightly latr in tim (5.8_4.4 million yas ago) is th olltion of th
r aplik-onrirrg-humanlik s- thiopianspinrnsrfrrdto as rdipithcus (Hal|.Slssi2001; Wold-
his squnlooks vry aplik. By G a r i l t a l . 2 0 0 1 ; H a i l - S l a s s i t l . 2 0 0 4 ; W h i t e t a l r .| 9 9 4 , t 9 9 5 ) . A s
'uld xpt. wit rrori tt,tgsisand as ptd in n arly anstor' it is aplik in
p i n r h s u a m i | yH o m i n i n a i s most rspts,ut it dos shar rtain ky traits with latr hominins. Among
2002). This prious fossil skull thsis a anin tooth that is lss projting and lss pointd than that of fos-
lntaind faun linkd with datal sil or modrn aps and a ranil s that is shortr. Its tos apparntly had
'to
6 million yars a]o.Its ovrall thaility to hyprtndlik thos in ipdal humans (Hail-Slassi2001).
stimatdbrain size slightly low It livd in a losd woodland nvironmnt (WoldGaril t l' 1994.200t\.
1). But v rh Iogi of ladistis, it
t dfinour linag.Th most im-
t h t o f p s n d 1 r r i m a t si n g n - AstrIopithecusand othr Hominins
lwr prmolar. Tlris is a signifint 4.2_2.5illion Yars Ago
.tant diffrn twn aps and
rniodntal rmains of H. rdolfesis (-1.9 million yars ago) as longig Th gologilly lt <
to that spis' thn the mor humanlik ody proportions and hip arhit- dmonstrats th fallay
tur first appar in this spis just aftr 2 million yars ago (MHnry and progrssive stags. Populi
offing 2000). Both of thsspisar transitional,with som primitiv and othr populations unlss
som drivd haratristis of latr Homo. Postranial assoiations ar rit-
ial hr aus ody siz appars to vry diffrnt. Homo hbilis was
vry small odid (35 kg), and H. rudolfensis was lrg (55 kg) (Hnry Arhai Homo to H
1'994|.Saling hk-toothsiz to ody wight shows that thy oth had r-
vrsd th trnd of vr-inrasinghk-tooth siz. Rlativ rain siz x- Larg-ind spistl
pandd, spially in H. bbilis' 600,000 yars ago and in
Asolut rairr siz pands furthr with th apparan ol 17.ergsterlr- Nandrtals dominat E
tus at last 1.8 million yars go, ut ody siz also inrass,so rltiv 3 0 0 ' 0 0 0 t o 4 0 , 0 0 0 y a r si
brain siz apparntly Was not dramatially pandd (l/lkr and Laky H' spis appars 6rst in
1'99).Th arly Afrian form of this spisis oftn rfrrdto as th spis 1999, 2000). Th arlis
H. rgster to ontrast it with th wll-known Asian sampl of H. eretus latr am th hallmark
(.!ood1991; Spoor et a|. 27|. Body size and spially hindlim lngth su-Saharan Afria for
.Walkr
rah modrn proportions in this spis (Ruff and 199). oth .}IBrartyand Brooks 20(
synapomorphis with |ater Homo inlud a furthr rdution in prog- spis sprad out of Afri
nathism, a rdution in th postglnoid pross that is fusd with th tym- gntially swamping lo
pani plat, and rdution in th asolut and rlativ siz of th hk tth. Gol 2007). Th spra
Brains ontinu to xpand, and hk tth get progrssivlysmall during rly (50,000-46,000 yar
th volution f th gnus Homo. aly longd to modrn l
old as 40'000 yars (Bowlt
from Asia who migratd
First Disprsal out of Afria Alaska prhaps 20,000 to
of most of th Amria
Th first apparan of human populations outsid Afria appars to hav Gol t al. 2008). Pop
ourrd soon ftr th apparan of arly Homo in Afria. Fossil Hoo r- thousand yars ago from tl
mains from th Dmanisi sit in th Rpuli of Gorgia ar most likly old .. 300 and Nw Zea|and
than 1'.7 million yars and ar vry muh lik thos found in Afria attrib. 2007; Hunt and Lipo 2006
utd to H' erts/bbilis (Gaunia and Vkua 1995; Gaunia t al. 2000; Thnologial dvlopm
Vkua et al. 2002; Lordkipanidze t a|. 200 5, 2007 ). omo rtus oupied pa, ut it rtainly shows
parts of tropial Asia prhaps s arly as 1.8 million yrs ago (Swishrt al' rud ston tools prsist fol
|994). Th at hronology is still unrtain, ut somtim aftr this and r muh mor rent. Hu]
bfor 0.5 million yars ago some populations hd adaptd to lif in th tm- l s st h n 0 . 2 5 " o f t h i r v
pat limati zon of northrn Asia (Ant6n and Swishr 2001). linge.
An arhai spisof Homo lft tools on th island of Flors, Indonsia,as Th historial distrtuti
arly as 800,000_700,000 yars ago (Brumm t al. 2006; Morwood t l. viopd only in th past 40,
1998)' and in surprisinglyyoung sdimnts(95,000-12,000 yars ago) a nw ezs quippd with Uppr
spis, H. floresisls, has n rovrd (Brown et a|. 2004; Mrwood Afrian homland, first to ,
et a|. 204,2005; orwood and vn Oostrzee 2007).It has a small ody urop (Klin 1999). This
(29 kg)' minut ndoranial volum (380 ), and arhai ody (Larson t al. genti homognity found
2007; Tohri t al. 2'007).It is most likly a dsndantof an arly disprsal with that of our own los
of arhai Homl that undrwnt insular dwarfism (Brown t al. 2004), al- dgr of gnti htrogn
though sm hallng this intrprtation (Argu et a|. 206; Jao t al. that in Homo spies (Ga1
2 0 0 6 ; H r s h k o v i t ze t a | . 2 0 7 ) ' tions to loal onditions ar
Humn uolution 267
l million yars ago) as longing Th glogially lat ourrn of suh a primitiv spis of Homo
dy proportions and hip arhit- dmonstratsthe fallay of rgarding human volution as passing through
nillion yas ago (MHnry and pogrssivstags.Populations adapt to loal onditions and divrg from
sitional,with som primitiv and othrpopulations unlss hkd y gn flow.
P o s t r n i al s s o i a t i o n s r r i t -
ry diffrnt. Homo hbilis was
sls was larg (55 kg) (Hnry Arhai Horo to Horno spies
ight shows that thy oth had r-
) O t hs i Z .R l t i v r a i n s i z x - Largr-raind spis of arhai omo gin to appar in Afria y
600,000 yrs ago and in urasia somwhat latr (lark et a|. 1'994).The
l apparan of H. rgster/er- Nandrtls dominat urop and wstrn Asia from approimatly
dy siz als inrass,so rlativ 3 0 0 , 0 0 0t o 4 0 , 0 0 0 y a r s a g o ( K l i n 2 0 0 3 ; K u h n 2 0 0 7 ; K r a u s t a | . 2 0 0 7 ) .
. pandd (Walkr and Laky . spiesappars first in Afria somtim for 100,000 yars ago (Klin
i is oftn rfrrd to as th spis 1999, 2000). Th arlist tras of Uppr Palolithi tool thnology that
own Asian sampl of H. erets latr am th hallmark of arlist mmrs of H, spies first ppr in
and spially hindlim lngth su-SaharanAfria for aout 90,000 yrs ago (Brooks et a|. 19951
(Ruff and !alkr 1,99). othr Brarty and Brooks 2000). By aout 50,000 yrs ago populations of our
a furthr rduion in prog- spissprad out of Afria to Asia and thn urop, apparntly rplaing or
ssthat is fusd with th tym- gntiallyswamping loal arhaic Homo populations (Anikovih 2007;
rd rlativsiz of th hk tth. Gol 2007). Th sprd Was apparntly rapid in Asia, as vidnd y
l get progrssivlysmallr during early (50,00046,000 yars old) arhaologialtras in Australia that pro-
aly blongdto modrn H. spis populations found thr in sdimntsas
old as 40,000 yars (Bowlr t al. 2003). Amria Was sttld y immigrants
from Asia who migratd aross a land ridg that onntd Siria and
Alaska prhaps 20,000 to 15'000 yars ago' ut th sussfulolonization
of most of th Amrias bgan aout 12,000 yars ago (Dillhay 2003;
s outsid Afria appars to hav Gol t al. 2008). Popl first rahd som of th Paifi islands svral
Homo in Afria. Fossil Hozao re- thousandyars ago from th ast, arriving at th arqusas Islands y aout
of Gorgia ar most likly oldr .o. 300 and Nw Zea|and y aout .. 1200 (Lum and ann 1998; Finny
lik thos fund in Afria attrib- 2007; Hunt and Lipo 2006).
kua 1995; Gaunia t al. 2000; Thnologil dvlopmnt in human volution appars to rrari in
l5,27\.Homo eretusoupid pa'ut it rtinly shows a pattrn of alration(Klin \999). Relativ|y
8 million yars ago (Swishr t al. rudstntools prsist for ovr 1.5 million yars. Finly workd lad tools
tain' ut somtim aftr this and ar muh mor rnt. umans hav had agriultur' itis, and writing for
lns hd adaptdto lif in th tm- lessthan 0.25% of thir volutionary dvlopmntas a sparatmammalian
i n a n d S w i s h r2 0 0 1 ) . linag.
th island of Flors, Indonsia, as Th historial distriution of human gnti varition pprs to hav d-
mm t a|. 2006; Morwood t al. vlopdonly in th past 40'000 to 50,000 yars whn populations of H. spi-
(95,000_12,000yars ago) a nw s quippd with Uppr alllithi tool thnology sprad out of thir
d (Brown et al. 2004; Morwood Afrian homland' first to Asia and y at last 30,000 yars ago throughout
;terzee207).It has a small dy urop (Klin 1999). This rent olonization ounts fr th rmarkal
), and arhai od (Larson t al. gntihomognity found among living mmrs of our spis ompard
a dsndantof an arly disprsal with that of our own losst ilogial rlativs, P and Gorill' whr th
jwarfism (Brown t al. 2004), al- dgrof gntihtrognitywithin ah spisis man tims gratthan
L (Argu t a|, 20Q Jao t al. that in Homo spil1s(Gagnu et a|. 1999; Ruvolo 1997). Rapid adapta-
tionsto loal onditions and th random snti hanes that n our in
268 Hum uolutio
small isolatd populations hav ld to svral onspiuous phnotypi mammalian avrag (th
hangs, suh as skin olor, hair form and olor, and ody proportions' ut t i o n s k p t u p w i h i nt
ths diffrns appar to rlatively rnt altrations. Th gographi sis (Brown et a\. 24),
distriution of skin olor is losely orrlatd with th intnsity of solar radi- ndowd with modrn r
ation suh that tropial popl in Afria, Asia, and th Amrias hav grat s u j o f r i n r o r g e n
dnsity of mlanin pigmnts in th skin for prottion gainst tissu and aly profound hangs t
mtaoli damag. Highr-latitud popl ar dpigmntdmost likly - othr uniquly human fa<
aus ultraviolt radiation from th sun atalyzs th synthsis of vitamin D ti hang that nald
in th skin and aus of othr fators (Jalonski and haplin 2000). D- haviors ginning aut ]
pigmntation of th skin in high-latitudpopls appars to tru of Nan-
drtals (ulotta 2071 Lalueza-Fo et a\. 2007). Body proportion varis
gographially aording to tmprtur' so that popl in hot limats hav volution of Humar
longr lims and lss gloular trunks to allow gratr hat dissipation
through th skin (Ruff 1994). ptions to this pattrn an oftn - Bipdalism is th first mz
plaind y th prsn of ultural daptations that allow prfrntial sur- first hominins wr mos
vivl and rprodutiv suss indpndnt of gnti daptations. Th first odis without th spi
H, spis in urop, fo xampl, had ody uilds haratristi of tropial mark of our linag (Tho
popl (thy am from Afria), ut thy had thnologis far suprior to possily uropan radiat
thos of th loal Nandrtls. Thnologial innovation is oftn th rason win's wods, ..to liv som
..a hang in its
for population xpansion and plays a pofound rol in dtrmining why r- mannr t
tain physial varitis of H' spiens oupy gographi rgions and displa ditions of its native oun
arlir indignous popl (Diamond 1997). th human lad proa
volutionary novltis. l
that podud th ipd
volution of th Human Brain dfin th human grad
transformations of th tr
Th aumulation of volutionary novltis through tim rvals th ontinu- fossil rord. It may t
ity of human volution. Th dmonstration of this ontinuity is a triumph of rats so that th lngth of
volutionary iology. It furthr onfirms Darwin,s viw of dsnt with mod- of th hallu. h xtin
ifiation. Th pansion of th rain in th hominin linag is on of th most pithus appars to hav l
dramati dmonstrations of this. Absolut and rlativ rain-siz xpansion big to (Kohlr and
follows tim rmarkaly wll, as shown in al 1. Th arlist speis for rmidus will hlp larify <
whih rain siz is known, . fresis' has an avrag ndornial volum ipdlity in our linag.
of 48 (asedon 4 spimns),ompard with modrn himps at 400 ' arly volutionists, rr
gorillas at 500 ' and H' spis at 1,350 . Th ndoranial volum for . akl (1868), prdit
grhi \1 spimn) is rportd to 450 , for . fricus (7 spimns)' rain. How' why, and i
4 5 2 , f o r P . b o i s i ( 1 s p i m n,)5 2 1 c c , f o r P ' r o b u s t s( 1 s p i m n ) 5' 3 0 uniqu ostur is a favor
, for H. hbilis (6 spimns), 6|2 c, for H. rudolfsis (1 spimn), 752 2004). It is possil tha
' and for . ergster ( spimns), 871 . Howvr' at last on spisof ipdality, givn th fat t
Homo (H. |loresislsl managd to surviv on a rmot island until 18'000 wih folim suspnsio
yars ago with n ndoranial volum of only 380 (Brown et a|.204). ipdal, this ody plan a
Brains epand during th Plistonfrom 914 twn1.8 and 1.2 mil- quadrupedal mammals
lion yars ago to 1,090 twn550,000 and 400,000 yars ago' 1,186 c (Rodman and Hnry l
btwn 300,000 and 200,000 yars ago' and aov 1,300 aftr 150,000 Thr has n a pul
yars ago (Ruff t aL. 1998). Rlativ to stimatd ody wight, th fossil primitiv traits rtaindin
spisgo from slightly aov th siz sn in modrn aps to 5.4 tims th On pol mphasizsth 1
Hum L'olutio 269
svral onspiuous phnotypi mammalianavrag (th olumn lald V in Tal 1). Not all plpula-
olor, and body propotions, but tinskpt up with rin pansion, s is dramirtiallyshown H. flrlrsien-
nt altrations. Th gographi sls (Brown t al. 2004), ut all tant populations of H. spies ar fully
d with th intnsity of sola adi- ndowdwith modrn airrs.Unfortunatly th fossil rord is mut ()n th
\sia, and th Arrias hav grat sujtof rain rorganiztion,whilr is pity usthr wr prsum-
for prottion girrst tissu and aly profound hrrgsthat aompnid th dvlopmnt f sph and
, ar dpigmntdmost likly e- othruniquly hurrranfaultis.A strong i1san mad for a major gn-
,talyzsth synthsis of vitamin D ti hangthat nald H. spis to spak nd prform modrn human -
haviorsginningaout 50,000 yrs ago (Klin and dgar 2002).
lalonski and Chaplin 2000). D-
oplsapparsto tru of Nan-
I. 2007). Body proportion varis
;o that popl in hot limats hav volution of Human Walking
o llow gratr hat dissipation
j to this pattrn an oftn x- Bipdalismis tlr first major grad shift in hominin volution, r.rtth vry
ations that allow prfrntial sur- first hominins wr most likly gnralizd hominoids with rathr plik
rt of gnti adaptations. Th first odiswithout th spializationsfo ipdlism that latr am th hall-
dy uilds haratristiof tropial mark of our linag (Thorp et a|. 2007). Thy wr part of th Afriirn and
' had thnologisfa suprior to possiblyuropan radiation of lat ion hominoids that am, in Dar-
..to liv
ial innovation is oftn th rason win'swords, somwhat lss on trs and mor on th ground'' du to
.oundro1in dtrminingwhy r- ..a
hangin its mlrn of prouring susisttl'or to a hang in th on-
ly gographirgions and displa ditionsof its nativ ountry'' (Darwin 7872, p. l.]5)' h fist memrs of
th human lird proaly will rognizd in th fossil rord y sutl
volutionary novltis. Th major gnti/dvllpmntaltransformtions
that produd th ipdally adaptd trunk nd hindlinr morphology that
dfinth human grad proly appard latr. Th pis squn of
transformations of th trunk and hindlim may nvr known from th
ls through tim rvals th ontinu- fossilrord. It may that parts of th ipdal ody hangd at diffrnt
n of this ontinuity is a triumph of ratsso that th lngth of th plvi lads rdr'rdarlir than th addution
)winviw of dsntwith mod_ of th hallux. Th til-ltand distintl nonhonrinin ione ape oreo-
: hominin linagis on of th most pithusappars to hv had a rdud plvi lngth but rtirrda divrgnt
a n d r l t i v r i n . s i z x p a n s i o n ig to (Kohlr and Moyi-So|' 1997). lr full analysis oi rdipithus
in Tal 1. Th arlist spis for rmiduswill hlp lrify our undrstandingof th natur of th traI-rsition to
las an avragndoranial volum bipdalityin our linag.
dwith mdrn himps at 400 ' arly volutionists, inludirrg Lmark (1809)' Darwin (7872\, and
l . Th ndoranialvolum for . akl (1868)' prditd that ipdality prdd the pansion of th
, for , frins (7 spimns), brain. How, why, arrd in what haitat an aplik anstor adoptd this
uniqupostur is a favorit topi of spulation (Kingdon 2003; Mnr
, for P, robustus (1 spimn)' 530
H' rudolfsis\I spimn), 752 2004)' It is possil that many aplik spis in th ion zrdoptd
. Howvr, at last on spis of bipdality'givn th ft that th ap bodv pln is daptd to nroving in trs
v on a rmot islnd until 18,000 with forlim sr-tspnsion and not to f6int quadrupdal walkirrg. on
only 380 (Brown t al. 2004). ipdal,this body plan an mov with an ngti ffiiny qual to that of
> m 9 t 4 t w n1 . 8 a n d 1 . 2 m i l - quadrupdalmammals and muh gratr than tht of quadrupdal apes
) 0 a n d 4 0 0 , 0 0 0 y a sa g o ' 1 , 1 8 6 ( R o d m a na n d n r y 1 9 8 0 ; S o k o l e t a L .2 7 | .
Thr has bn .r puliar polarization of viws aout th maning of th
, and abov 1,300 aftr 150,000
' stimatdody wight, th fossil primitivtrirs rtaindin th postrni of rly spisof ustrlrlpithus.
n in modrn ps to 5.4 tims th on pol mplrasizsth bipdal spilizations(Lovjoy 200.5,2005), and
270 um uoltio
ar uniquly Homo (Bra
th othr pol lls attntion t() tl.l many primitiv haratristis (Strn
tivly nlagd hip ioints
2000). Viwd in gand prsptiv,th divrgntintprtatil1s an sn
haps rlad ro long-dista
as diffrns irr mphasis. Both amps agr that all spis of AustrIo-
H' spies. Th disover
pitheus wr ipdal. hat implis tht thsspisdid not lim lik aps.
volution in that it has a ]
Thy did, howvr, rtain morphologia1 fatursssoiatdwith arboral.
rlativ forarm lngth (l
ism for at lst :r rillion yars nd a hip arhittudiffrnt frol that of
latr spisof Homl. That implis so diffrnfro-tmldrn humans in
gait and linring ility, ur thy did not walk :lnd lim lik ps.
Th volution of Hr
Th publishd hominin fossil rord dos not yt hav a tru intrmdiat
stag twn an aplik and a humanlik ody (Hnry and Jones 2006I.
xpt for rlativly sma
Th postrniur-lrof ustrll1lithesrtins mally primitiv aplik traits
lar to thos of many of th
ut is fundamntally rorganizdand highly spializdfor ipdality,unlik
of our sufamily livd il
that of any ap (Hnry 1986). hr is a profound altratiorriIl th gn-
thir dits wr simil l
ti tmplat that produd th short plvi lad, addutd hallu, and othr
limaxing with th appa
ipdal traits of this hominin.
ago), th hk teth 1
Th diffrn twn Astrlopithes'spositional and loomto -
Although Darwin and his
havior nd that sn in latr spisof Homo ;rmlarrwith th disov.
fossil rord would vnt
is of postranil fossils assoiatd wit . rgtstr/erets(]il/illrkrand
..StrppingYoth'' (KNM WT 15000). Thr minin nrgdonrsA . fri
Laky 199|, partiularly th
rlativ to ody siz. But
ar striking diffrnstwn th plvis and fmur of all spis of us-
molars with onomitnt
trIopithus and those of H. ergster/rets,with th laftr apparing
hks, and attahmnt :
muh mor humrrnlik (Mnrv and offing 2000). This proal,vrgistrs
proimat trend throug
a nrajor shift in adaptation twl-lstrlopitbttsand Homo. From this
hk-tooth siz from sn.
prsptiv'ustrlopithe,/sapprs t :l ipd with fr hirnds ftlr arry-
ntttsisnd . frcttsis
ing ut st daptd to short distans and rvith a halthy ppriation of
hug in P. boisei and P.
trs for safty, fding, and slping. Th longr fmora and mlr human-
so that from H. bbilis at
lik plvs tht appar 1.9 million yars a!]o wit omo mark th gin.
siz oms progrssiv
ning of an impotant hang.
l a t i v h k - t o o t hs i z
From th urrntly availal fossil vidn' thppar to e two grad
mgadorrtiaq l u o t i n ti s s
shifts in th postranial volution of th hurrrzrrrody. Th first Was th tran-
lar and first two molars
sition from gnralizdhominoid to hominin ipdalism that is doumntd
spis od wight. Ir is
in . msis' . fresis' . frius, and . grhi (Ward 2002). Th
rl indition of rlativ
diffrnsin postrnial morphology among thsspisar diffiult to as-
m g a d o n t i a t h o u g ht i m
sss usof tlr inompltnssof th fossil rrd.All shar ]rivdfa-
t o p r o s s g a t rq u a n t i
turs with H{lnll, ut onl1' . zlfresisarrd . frius ar omplt nough
rvrsl of th trnd in m
to mak dtild ompaisons. [n som rsptsthsspisar rmarkaly
to oth ditary hang (i.
similar rlativ to othr spis of Hominoid;r. For xampl, thy shar a
orporation of mr rna
similar mi of aplik, humanlik, and uniqu fatursin thir wrists, hips,
for mastiation.
and kns (MHnry 1986). Th,vapparntly diffr in th ratio of forlimto
hindlim joint sizs,howvr, with ' frinus ppaingto mo :rplike
vn though it is latr in tirrrnd mor Holo-|tke in raniodrrtalfaturs
(N4Hnry and Brgr 1998; Grn et a|.207). Both spisapp to shar
Rmaining NIystri
omination of spializdipdal traits, ut not xatly lik thos of mod-
Th divsity of living m
rn humans, in faturs of th forlim ssoiatd with liming (Strn
i s h d l a t i v t o w h a t i t r
2000). Th sod grad shift in postranial volution took pla out
it is lar tht thr wrr
1.9 milliln yars ago with tlr 21pparan lf rnr.rhmo humalik hips that
Hum uolution 271
ar uniquly Homl (Braml and Lirma 2004)' Long fmoa and rla-
iy primitiv hartristis (Strn
tivlynlargdhip joints mark a signifianthang in loomotion tht is pr-
v r g n itn t r p r t t i o nsa n s n
haps rlatd to long-distan, ffiint striding mor lik that sn in modrn
rgrthat all spis of ustrIo-
H. spies. Th disovr of ' grhi revea|sth omplity of postranial
tsspisdid not lim like aps.
volutionin tht it has a humanlik ratio of fmur to humrus ut an aplik
fatursssoiatdwitlr aroral-
rlativforarr lngth (Asfw et a|, 1999).
arhitturdiffrnt from that of
diffrnfrom modrn humans in
walk an]lim lik aps.
s n()t ,ytlrav a tru intrmdiat
Th volution of Human astiation and Dit
ody (Hnry arrdJons 2006).
pt for rlativly small anin tth' th arlist hominins had tth simi-
ains mar-ryprimitiv aplik traits
lar to thos of many of th ion aps (Ungar 2007). Ths arly mmrs
ly spilizdfor ipdality, unlik
of our sr'rfamilylivd in woodland haitats (Pikford 2006). Prsumaly
a profound itration in th gn-
thirdits wr similar as wll. But ginning aout 4 million yrs ago and
: lad,ddutdh[lu, and othr
lima-ingwith th apparanof Prthropus boisi (2._1.4 million yars
ago), th hek tth pand through tim (onstantino and .il7ood 2007).
s's positionl and loomotor -
'zo anrlarr rvith th disov- AlthougDarwin and his ontmporarisprditdmuh of what th human
fossilrord would vntually rval,no on antiiptd th disovy of ho-
h H. rgstr/rttts(Walkr and
minin mgadonts. Afrian aps and modrn humans hav small hk tth
Youth'' (KNM $T 15000). Thr
rlativto ody siz. But rly hominins had rlativly hug molars and pr-
s and fmur of all spis of us-
'rtrrs'nith h lett pping molarswith ono-litantlygiganti jaws, alvolar bon, uttrssdfa and
hks'and atthmnt aras for th hwing muslatur. Thr is n p-
:fing2000). This proaly rgistrs
proimat trnd though tir in sussiv spis of inrsing rlativ
rlopithezusad Homo. From this
hk-toothsiz from small in rdipithecus rmidus to modrat i . -
a ipd rvith fr hands for arry-
mnsis and . frsis, ig in . fricnus, P. robustt'ts, and . grhi, and
nd with hlthy appriation of
hug in P. boisi and P. ethilpius. Th trnd rvrss in th Homo linag
llngr fmora anl mor human-
so tht from H. bbilis and H. rudolfnsis to modrn H. spis, th rlativ
s ago with Homo mark th gin-
siz oms progssivly smallr. On way to xprSs ths trnds is with
rlativhk-tooth siz, as is shown in th last olumn of al 1. Th
n'th:lpparto two grad
rngadontialqr-rotintis simply th asolut siz of th lowr sond pmo-
uman rrdy.Th first ws th trn-
lar and first two molars dividd y thir ptd siz drivd from th
inin bipdlismthat is doumntd
spisody wight. It is not intndd to pris ind ut mrly a gn-
zs' and . grhi (.!fard 2002). Th
ral indiation of rlativ siz. It is rasonalto assum that th inras in
ong thsspisar diffiult to as-
nrg;rdontia through tim is rlatd to ditay spializations(i..,th aility
fossiIrord. All slrar drivd i_
to pross gratr quantitis of low-nutrint and tough-txturd foods). Th
d . fricus ar omplt nough
rvrs;rlof th trnd in mgadontia in th volution of omo may rlatd
sprtsthsspisar rmarkly
to oth ditary hang (i..,lss lian on tough-tturdfoods nd th in-
noida. Fo ampl, thy shar
orpotionof ror mat in th dit) nd th prossingof foods with tools
niqu fatusin thi wrists, hips,
bfrmstition.
rtly diflr in th atio of forlim to
us apparingto mor aplik
H om o-|ik in raniodntal faturs
Z007).Both spisappar to sha
Rminingystis
i' ut not atly lik thos of mod-
h divrsityf living mms of th suprfamily ominoida is impovr-
assoitdwith liming (Strn
ishdlativto wht it was in th past. vn with th limitd fossil sampl,
ranial volution ttlok pla alut
ir is l that th Wre many kinds of aps and hunrns long ago. or
of muh mor humanlik hips that
272 umlt uolutio
sis await disovry. Thr wr proaly many volutionary xprimnts Argu' D., D. Donlon,
in th varid haitats of th Afin ion to liston (Suwa t al. irophali',g
2007). Although th urrnt sanrpl of fossil hominins lads som to th im. u o l u t i l n5 l : 3 0 - j
Arnason,U., A. Gullrg
pression that thr wr only a fw hominin linags (.!fhit t al. 2006), it
of pimatdivrgn
far mor likly that our family tr will turn out to quit ushy (Sarminto o f m t l d nh u m . r l l s
et a|.2007). Spisnams may nd to multiply to aommodat th divr. Asfw, B., . !7hit' o. I
sity, although a alan nds to maintaind twn xssiv splitting Austrlopithusgt
and lumping. 284:62-9-635.
Bgun,D. R. 1994. Rlt
It will b wondrful to rsolv th qustion of knukl-walking in human
intrprttions as
volution(Rihmond and Strait2000,200l; Rihmond t al.2001). Living
Athropolog37: 11
spis of th Afrian ld of Hominoid inlud thr knukl-walkrs B g u n ,D . R . , n d . V . \
and a ipd. Th odds ar that th first hominin Was also a knukl-walkr, nw iddl N{ion
ut th arlist hominins known so far ar ipds and limrs without sp- Bhrnsmyr' A. . 2006
ializations for knukl-wlking. Prhaps thr funtionl and dvlop- Bowlr,J. ., H.Johnst
nd N. A. Spoonr.i
mntal prosssthat pdispos larg-bodid aps to om knukl-walkrs
hangsat Lak ur
whn thy adapt to haitats that rquir mor trrstriality. If so, that appar- Braml,D. ., and D. E
ntly odd gait may have volvd in paralll irr two o more linags. of Hotl. Ntur 43
Thr may hv n svral pansios of populations out of Afria dur- Bromag,. G., and F. S
ing favoral limati onditions that populatd parts of urasia and thn Hum uolutio.N
am xtint. It is sdutivto ir-rragin Boks,A. s., D. . Hlg
only on triumphnt puls of mi.
R . G ' K l i n ,W . J . R i l
gration that sprad humanity for th first tim throughout urasia' Th
J . V n i s' n dJ . .
sprad of H. spies appars to hav happend in a short amount of gologi- sitswith on point
l tim' ut rhai spis of Hlmo wr poaly muh lss adaptabl B r o m ,R . l 9 . ] 8 . h l
and mor vulnral to th svr limati shifts that ourrd vywhr 77-379.
ovr th past svralmillion yars. Loal xtintions may hav bn ommon Brown,P., . Sutikna,.
R . A . D u .2 0 0 4 .A n
throughout th hanging gographi rang of arly hominins. Bfr 50,000
lors,Indonsi.Na
yas ago humirns Wr an insignifiantlysrall part of th vrteratfauna. B r u m m , 4 . ,F . A z i z , G . D .
It is tmpting to swing to xtrm viws of pssimism or optimism. Th h- Kuniawan,D. R. H
minin fossil rord is limitd and inmplt, ut it is also rih and onsis- Flosirnd its impli
tnt. Its ovrall quality is onsistnt with Darwin's viw of dsntwith B r u n t , . ' F . G u y , D . P i
modifiation. It maks it vry unlikly that, for xampl, signifint n. A. Bauvilain,. Blor
J . D j x , . D n v s ,P
phaliztion gan for th volution f ipdalism in ou linag.n.
D. Graads,T. Lhm
fliting intrprtationswili always pat of th sin,but datis a sign G . o u h l i n ,O . o t
of intelltual halth and hlps ensur that idas ar groundd in aurat . Spnt,N4.Shu
osrvationsof ratrialvidnand pris logi. ..Zazzo, and . Zo|
hd, CentlAfria
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.Vood'
B. 199|. Koobi Frlr Rsrb Proit.o|' 4, Hlmiid ril Rmis. of groundhogs found ,
Oxford: larendon Prss. and prairi dogs ar k
Zollikofr, . P. ., M. S. Pon d Lon, D. . Lirmn, F. Guy, D. Pilam,
and tularmia (rai
A. Likius, . T. kay, P. Vignaud, and M. Brunt. 2005. Virtual rnial
nods that ould indi
rnstrution of ShlthroDs tchdsis. Ntttr 44: 7 5 5_7 59 .
a diffrnt dirtion.
As it happns' th
hiago, whr it
ulats in wild animal
a los rlativ of th .
a shipmnt of Gamiz
prairi dogs had n
link in th hain of i
forgd (D 200; G
though his lovd pl
halth workrs ar tr(
munity against monk
Th ase of this par
monkypox outrak
sons who am in on
pt faility (D 200:
loolltionof ust r lop ith us
PhsiIAthrrlpoklg 45
281
282 uolutilr Biolog of Disse d Dr,ii ediie a
smallpo, ut th rsmlanof th monkypo virus to its ousin, whih is mismth wn th
widly livd to hav n radiatd from th wild by a wrldwid vai- volvd and adaptd'
nation ampaign in th 1970s, is frightning.Th as also rvalshow mod- t w n h i g h f i n s si n t |
rn mdiin and publi halth us th framwork of volution to solv Ns 1991; Swyngh
potntial halth riss.This has n alld Darwinian mdiin (so) - hart disas (Swyngh
aus diagnosis and tratmnt dpnd on mthods of volutionary iology to and prosttanr ((
sluth th undrlying auss of immdiat symptoms (\/illiams and Ns fts (Glukman and
1991'; Stearns 1999).Inftion with monkypox was onfirmd y viral iso- hv nprimry r
lation' pidmiologial invstigation,and finally DNA analysis, and family Th rnt worldwir
rsmblans among po viruss trad this partiular virus k to Afria. a m p l t h t p r v i o u ss t
A similar story an told aout th oronavirus that auss svf aut gns that aus dia
rspiratory syndrom (SARS) and sparkd a puli halth pani in ality, howvr, fw a
2002_2004 whn inftions sprad from hospitals in Asia to North Amria. l a t i o n t o d i s s :t h
In a mr ight months gnti nalysis of th nwly mrgd SARS virus first g l o i n ' a n d m l a i r
ld to its idntifiation, thn to intrmdiats in ivt ats sold in wild-gam on, nd th story dl
markts in hin (Ruan t al. 200), and vntuallyto potntial rsrvoirsin populations (Hill and
a t s ( L i t a l . 2 0 0 5 ; . W . a n g t a | . 2 0 6 ) .o n a t r a g i a l l ym u h l a r g rs a l t h and inftious disas
human immunodfiiny viruss (IVs) that aus AIDS hav also n a s p t so t h n v i o
trad by gnti analysis to wild primat populations in Afria (ahn t al. immun rsponss .n
2000; Klet al. 2006). l 9 9 9 ; W i s sa n d M M
s o n 2 0 0 6 ) . F i n d i n gt h t
Although I aknowl
tiology tinary for tht |
m d i i n t h a t r q u i r
Darwinian mdiin sarhs for th volutionary origins of siknss-thos d p t i v i n h L r m n si
things that mak lif nasty, rutish, nd short. Darwinin mdiin suds off th world of m
y looking for disas origins in iologial adaptations of humans and th al sltion ut lso h
miros that plagu thm-thos traits that allow sm to surviv and r- of ommon dsnt fr
produ bttr than othrs. On asi tnt of Dawinian mdiin oms 1999iBuIl and !ih
from adaptationist thinking, that disassaris aus humans ist in a r m a I a d p t i v t o h L
Disas, howvr, is l
ial wold in whih a.
Darwinian mdiin dsribs n volutionary approah to aring for idal ut ar rlativ tr
the sik, and for publi halth in gnral.In this sns,Darwinian md- i t i s .T h i s W s t h p o i
iin looks to th adapttions' and maladaptations' of oth humans and y Gou|d and Lwon
th miros that aus diss.For miros in partiular' Darwtntan stan' W an undrs
mdiin must aount for rlativly rapid gnration tims of patho- sns, in th adaptiv
gni miros, and high rats of mutation and gnti rominatlon tm upright t th to
that lad inxoraly to novl adaptations and th mrgnof nw dis- movmnr that dos l
ass. Both pathogn and host, howvr, also must undrstood in why w hv svnvt
trms of volutionary history' homology, and phylognti onstraints Th answr ro rhis qu
on adaptation. Drwinian mdiin, thn, taks a dynami viw of oth and vrtbrats and in
inftious and gntidisass,whr svralvolutionary historison- inhritdfrom our an.
vrg within individal patints on th on hand, and in human popula- adaptiv nmr in th
tions on th othr. passdOn to Us as i l
and Lwnttn 7979).
lttdii Llolutionry'Biologl, of Diss d Drii dii 28.]
:ypovirus to its ousin, whih is mismath twn th urrnt world and th world in whih our spis
th wild by a worldwid vai- voivdand adptd, and that lrumans sr.rffrfrom sris of trd-offs -
;.Th aslso rvls how mod- twnhigh fitnss in th past and rdud fitnss irr th prsnt (.Williams arrd
ramwork of volution to solv Ns ]991; Swvnghdar'rw2004; Glukrn and Hanson 2006). Ositv'
Dawinianmdiin (so) - hartdisas(Swynghdauw2004), tpe-2,diats (Dianrond 200), rst
thodsof volutionry iology to and prostat anr (Gravs 2002\, goitr' iodin dfiiny, and irth d-
syptoms (\/illimsand Ns fts(GlLrkmnnd Hnson 2006), nd aging (\Williamsand Ns 1991)
ypo was <rnfirmdy virl iso- havn primary targtsof this kind of thinking.
inally DNA analysis, and family Th rntworldwid inrasin typ-2 diatsprovids a omplling -
s partiularvirus ak to Afria. mpi that prr,ious sltiorrflr tolrating starvzrtion rrrily hav sltd fo
rnavirus that auss svr aut gnsthirt aus diatsin a food-rih nvironmnt (Diamond 2003). In r-
kd a puli hlth pni in ality, horvvr,fw ass doumnt ongoing natual sltion in humans tr-t
'spitalsin Asia to North Arnria. rlation tl disas:th s of sikl-ll anmizr,low-fur.rtioningo hmo-
r nwly mrgd SARS virus fist gloin, and malria rsistanin su-Saharan Afria is ssntiallv th only
s irr ivt ats sold in wild_gm on,and th stor dos not hold up gnrallv outside a fw wll-doumntd
n t u l l yt o p o r n t i lr s r v l i isn populatins(Hill and tulsky \999). As will showrr latr' oth gnti
a tragiallymuh lrgr sal th and inftiousdisasshav multipl auss that inlud nutrition and othr
tht us AIDS hv also n sptsof th nvionment' gnti sLlsptiility,mrvoni dviopmnt,
looulationsin Afria (Hahn t al. immun rsponss' and pathogrr divrsity (Burn and \/hit 19721 Stearns
1 9 9 9 ; s i sasn d M i h a l 2 0 0 4 ; G r a h a r n t a l ' 2 0 0 5 ; G l u k m a n a n d H a n -
son 2006). Firrding th aussof disasrmairrsa diffiult hallng.
Although I aknowldg that natural sltionis th most powrful volu-
tinary for that lds to adaptation, I t:rk a diffrnt tk for Dirrwinian
mdiirrthat rquirslss spulatilnaout whirt may or may not hv n
tionary origins of siknss-thos adaptivin humns in th past. vn so, volutionary iology has muh to
ort. Drwinian mdiin suds offrth lvorld of mdiin via undrstndingnot onlv adaptations y ntu-
l adaptationsof humans and th ral sltionut also hrologis,traits tht ar shrd y organisrnsaus
rt allow some to surviv and r- of ommon dsntfrom thir anstors (Williams and Ns 1991; Starns
lt of Dawinin mdiinolns 1999;Bu|Iand Wihmn 2001). larly oth gnti and irrttiousdisss
aris aushumans xist in a ar malildaptiv to humns in that moridity and mortality rdu fitnss.
Disas,lrowvr,is an insapalonsqunof an vr.hngingiolog-
ial world in whih adaptatior-rirrrdfitrrssr not asolr.rtly tund to sorn
ionary approah to aring for idalut ar rlativ to what might our in a world of onstraind possiil-
. In this sns,Darwinian md- itis.This Was th point of th farous ritiqu of th adaptationistPOgln
laptations,<lfoth humans zrnd by Gould and l,wontin 0 yars ago (Gould and Lwontin 1979). For ir-r_
: r o si n p a t i u l a r ,D a r w i n i n stan'w an undrstand th svn vrtra in our nks in a funtional
pid gnrationtims of pato- sns,in th daptiv tra.l-offs tlr:rtkp our orry skulls :rnd nrvous sys-
'ion and gttiromination tm upright at th top (font) of our odis whil mainti.rining11iilityof
s and th mrgnof nw dis- movmntthat dos not damag our spinal ord. But this dos not tll trs
:r, lso mllst undstood in why lv lrv svn vrr'rot six lr ight (Williars rrd Ns 1991).
y , a n d p h y | o g n t i o n s t r a i n t s Th answr to this qustion rsids in our volutionary history as hordts
l, taks a dynami viw of oth and vrtratsatld in th dvlopmntalProgram nodd in th DNA w
, l v o l u r i l n a rhy i s t o r i s o n - inhritdfiom tlur anstors.Svn nk vrtr:rmay lrv rpresntdn
n hand, and in human popula_ adaptivnurr in th past, ut this aus is lost in history, and th rsult is
passdOl1to us ils a plrylogntionstraint of our ody plans (s Gould
nd Lwontin |979\.
284 uolutkr Biolog of Diss d Drii diie t
rsonsidrsth volutiorr of dis- A primary ampl is th shitling antigns (surfaprotins that stimulat
tins tht prinotlr natural immun rspollssvia spifi antiodis) of influnza A viruss. \V r all
within a framwork of phlogn- familiar with anrrual flu outraks' though prhaps not with th pandmi
ors to ginthinking lik popula. Spanishflu of 1918-|919 tat killd mor than 40 million individuals. Flu
:ss out varial populations o{ outraksour worldwid ah yar On a lss dramti sal as th viruss'
ls, in arr ologial sphr that vr-hangingsurfa antigns avoid immunity stimulatd v prvious inf-
tte 1972 L1rrg 2000; \/'An- tions and nssitatonstant updting of flu vains (Bush et al.1999;Fer-
:diur|sin will grtl enfit guson t al.200). lnf]unz viruss r rmarkly sirrrpl:ight RNA
ls' frlmhuman popultior-rsthat strndsod for intrnal protins tht surround th RNA, an trnal pro.
s of mirosthat liv within on tin apsul, polymrass (nzyms)fo viral rpliation, and 3 protruding
a I . 1 9 9 9 1S t a r n sa n d r t 2 0 0 1 ; protinsthat at as antigns' 1 1 protins in all. Th 3 surfa protins ar
two asptsof th urrnt lruman hmagglutinin(HA) fo inding to host lls, protin hannls that aid RNA
populatilnsiz and volution of invasion into th host ll (M2), and nurminidas (NA) to hlp nwly
l t i v t o o u r o w n g n t i s a n d formd Partils saPth host ll. Gs oding for HA and NA ar sp-
ially prorr t() muttion' and otlr A arrd NA ar rprsntdy multipl
antignisutyps within grorrps of rltd hosts. For instn, watrfowl,
shor irds, and poultry, th main rsvoir and spillovr hosts for th in-
flunzaA viruss, hav at last 16 HA and 9 NA antigni sutyps (olsn
)rsnta rsour for miroil et a|, 2006). Th urrnt virulnt avian influnza sutyp is thus dsignatd
th hurnanpopulation has grown H5N1.
l i l l i o n i n t p a s t d a d ,a n . hl u - lfithin ah sutyp, howvr, HA nd NA gns ar subjt to rgular
f tlr plant's iorn:rssand us a antignidrift in that flu viruss umult nugh muttions to bom
t p r - r(tV i t l u s k t a l . 1 9 8 6 ; D i l y lssasily rogr-rizdy th imnrun systmsof prviously infted or vai-
pultionshav pandd aus natd individuals (Bush t' a|. 7999: Frguson t al. 200). Th implia-
t us from th miroial world, tions for human halth ar normous, oth in trms of prditing how long
-rt mortality and longr lifspan vainswill rmain fftiv (D. J. Smith 2006) and in th possiility of
rg 2000). Th othr sid of th mutations or antign shuffling twI-lvirus sutyps,whih ould lad to
tion of food. But dd to this tlr hangs in viruln and transmission twn hosts (Baignt and .
h provnaility <lfr,iruss,t_ uley 200; Kuikn t l. 2006\. h grt far, r>follrs' is that th H5
tt to nw nvironmnrs.Summd hmagglutinin will hang influnza's tnsmissiility in humans, for in-
imls and plants onstitr.rta vast stan)y volving HA affinity frorn lls dp in th lungs t lls that lin
l otat with wild rsrvoirs of our throats and noss. Rathr than sttrdass assoiatdwith posur
to inftdwatrfowl or poultry, a nw form of HSN1 ould transmittd
I r g u l Iyi r u m v n t u r l l l l t i m i - y dirt ontt twn popl, lading to disas outraks in human
again nl again y miros tht populations.
its and lnovmnts.F<lr-rpl,
ti intrmdiatstwn oron-
GNls oF MUAloN
k r si n h i n a . n d h u m . t r l si:t p -
som volution in ivt ats' not Many wll-known human disass rsult from mutarions in our gnoms.
2006). uh of this an pr- utation is also invital, a onsqun of mistaks in opying th DNA
ldiinvia th adaptiv potntial that for th most part kps th rod straight during ll yls that rnw
pathogns(Ldrrg2000). i- our odis and mak our ggs and sprrr.Sot mutations :rspontanous'
i tha ous, hv n shown tim somar nvironmt-ttally aused,ut st ;rssurdthat mutation nvr sleps.
by rrturalsltion nd provid Individualmllttions ur at rltivly low ats,ut th numbrs ar tiltd
str.rdis of vlution (t 1998; in favor of muttions apparing in high absolut frqr.rny.Prvaln of
ommon gntidisasslik diats,musular dystrophis'and ansan
286 uolutior Biolog of Diss d Drtaii dii
n via thos inxol mutations al. 2005). Rgardlss,svralothr muttions in th R5 rptor also pr-
ni gror'rps (Frank 2004; Tishkoff sist,and it appars that sltionvia rsistanto inftions, rathr thn g-
arisin somwlrrbtwn 1 in ntidrift' provids plausil xPlanation for th high frquny of mutant
ior-rs. Givn a humn popultion allls(Blanpain t al. 2000; Galvani and Slatkin 2003; Novmbr t al.
0,000 gns and 6 billion s 2 0 0 5 ;S a t i t a l ' 2 0 0 5 ) .
h grrrationy th thousands, if Finally, inftions and gnti disassoftn o-our within th sam pa-
an trd and prdited within tints,and Darwinian mdiin an also nompass this rodr prsptiv
iffr in fr1ur-rY twn popu_ on human halth. Patints may siknd y pathogns, ut othr aspts
rhritddiss. of thir gntis and volutionary history (thniity) an mak individuals
pt gnsand hmful. on- prdisposdto inftion or to gratr pathlgnsisand mor svr disas
issausd y th inaility to on inftion sts in (ook and Hill 2001; \Wissand MN4ihael 2004).
21us of mutarions in th PH For instan,snr llls oi HLA gns of th immun systm inras sus.
as hs varial phnotyps, of ptiilityto a whol varity of pathogns, inluding thos that aus lp-
t, ut it n r,oiddif phny- rosy, tubrulosis,malaria' and AIDS (Hill and Motulsky 7999; Dan t al.
:d individuals (afully rd th 2002)' vn mor profound is that diss fom ommon human maladis
r than 400 rnr.rtntPH llls lik dngu'influnz,malaria, and turttlosissr'rltsnot only from dit
ln idntifidin various human danag from th pthogns bt also fror ovrrtion of th immun sys.
lr unusually high phnylalanin tm whil attaking thos pathogns (Graham t al. 2005). Thus th roots of
cs a rpid tst for diagnosis and disasar ompl and in many instans grow from dp in our gntis
dvlop. nd irmun rsponssto inftions,ppropiat or not.
'riss r hiddn in th huntn
f n < l r t h rn u o p nd s n t i n -
IDS, nd many of thm ar ho-
omosms)flr a trrutationin th
Diagnsis
]D4+T lls of th immurr sys_
What asptsof ar for th sik and for publi halth distinguish th Dar.
'5 rptorto ntr D4+T lls
winian approah to disas?Th largst distintion is tnding mdial ar
lf hlprT lls that normally r-
yondindividual patints to onsidr human populations in th ontet of
r lllapsof th ir-nmunsystll1
thir onntionswith th miroial world nd th patints' gnti and vo-
\32 ausof dltion of 2
lutionary history. This an b don without ompronrising immdit trat-
ulltional protin that prvnts
mnt to allviat pain and sLrffringand in many ass an provid mor
lls. R5-A32 is found in 5"%
rapid diagnosis, tatmnt' and rovry. A typial anrpl aiss ln trat.
:ntially asnt from st Asit-t,
mnt of fvr and iron dfiinvin patints rvith inftions,ausoth of
Dnt l. 2002)'
ths may rprsnt part of an adaptiv dfnsiv rspons to slow th
td for inrsd frqunv of
growth of batri (Williams and Ns 1991; Nse and.!7illiams 7994).
as n suggstdthat high fr-
Trating th symptoms of inftd patints ould hav th undsid fft
sltionin th past, spially
that inftions prsist longr, with gratr potntial for tansmission.
n wr oth omr-rronand da1l
Th following dision tr slrows atwys through this roadr ontt
ns and until th lat 1700s (Gl_
and hopfully an lad physiians not only to diagnos and trat disson
rvith surmising sltion in th
th asi symptoms and auss ar known ut also toward gloal undr-
laguor smallpo hypothss.In
standingof disas:
as ommon in CR5-dfiint
) 0 4 ; l v i n t a l . 2 0 0 4 ) , i n h L r m a n 1. Is this an inftion or a rrgnitlondition?
ls of th R5 rptor also x- 2. If it is an inftion, is this prson othrwis hlthy?
nd th R5-^32 mutant alll a. If othrwis unhalthy, is th patint ompromisd y poor
yasgo) skltonsfrom svl nvironmnt (.g.'poo dit and nutrition, smoking, or oupational
ght to hav invdd (Humml t h a z a r d s )?
288 uoltior Biolog of Dises d Drtuini dii
aids (lysin, hydro
. If othrwis unhalthy ut with a good nvironmnt,ould th
triian, ut in his l
patint b gntillyprdisposdto inftion (.g.,poor immun
first dtt GA1 via
systm)?
a gnti tst asd
If
. othrwis halthy, should symptoms tratd (.g.,nausa,fvr,
whr th Amish or
nasal ongstion)'or should this inftion allowd to run its
of uropan dsn
ours with minimal intrvntion?
rdud among Am
3. If it is an inftion' is this pathogn uniqu andlor ommon in humans,
Rognizing this
or is th pathogn's sour from othr animals (alld zoooti dises;
.$olfe in. In part this kir
s th rviw y et a|. 207|.
.\ihr sionals undr th st
a. did this patint ontt th inftious sour' and how was
th silvr_ulltya
th pathogn transmittd (..g., y dirt ontat or y an arthropod
in th 1930s and 1j
vtor lik a mosquito, fla, or tik)?
rdisovry of this t
b. How ontagious is th pathogn:should patints isolatd until
thrapis and th r
thy ar n longr inftious?
losis that wr thol
. Do w know th inftious miro's losst rlativs?l/ill thraois
Andrson 1999)'
to fight thos inftions hlp this patint as wll?
d. Aftr w idntify whthr th pathogn is a atrium, a virus, a
protozoan' or a fungus, dos th inftious agnt hav rsistanto
Tratmnt and.
antimiroials?
4. If this is a ongnital disordr' what is this prson'sfamily history and
Darwinian mdiint
thniity?
th patint in front
a. Is th individual from a small population with a history of marriags
prditiv and prr
btwn los rlativs (.g.,first ousins or losr)?
th mdial world,
. Is this disashighly prvalnt in othr ppulations with th sam
mbra oth natur
thni origins?
th pross that tr
5. !ill a thrapy or intrvntion hav an fft yond this patint?
.!7ill translation humans
a. thrapy slt for highr pathogniity or viruln in th
ntions to th natu
miros (.g.,fftsof imprft vains)?
.lfhat sin of volution
. kind of sletion prssur will antimiroials rat, and what
dvloping tratmr
will th trad-off twn intrvning with a drug now and
sn latr, som .
failur of thrapisin th futur if drug rsistanvolvs?
th ost f ignoring
ass.
fftiv us of this dision tre prsupposs a thnologial world whr
laoratory diagnostis asd on modrn iohmistry and gnomis are
availal for rapid srning.lThis tr also prsums that mdial and ADAPA|oN
halth profssionals larn th naturl history nd ology lf pathognsand
If on onsidrs mi
parasits in th ara whr thy prati, and that thy undrstand th thni
aus rsistan to i
divrsity of th population thy srv.
rmrgn of pr.
Th work of Dr. D. Holms Morton mong th Amish of Pnnsylvania
Andrson 19991 L<
and Illinois provids lar onntion twn iomdial rsarh and
multipl-drug rsist
linial prati (Blkin 2005). orton and ollagus first diagnosd and
y aquiring drug r
thn idntifid th undrlying auss of prtzl syndrom, a form of rral
This is tru for Nels
palsy with Svrmntal rtardtion and thn arly dath in fftd hildrn
gonorrha and dir
(Strausst al. 2003). Similar t PKU, prtzl syndrom is glutari aiduri
Similarly' multidug
(GA1)' ausd y m'-rtationsin th dgradation pathwys of thr amrno
uolutionr Biolog oi Dises nd Duini diine 289
dii
ids (lysirr,hydoylysin, and tryptophan). Morton is primarily a pdia-
od nvironmnt,ould rh
triian,but in his lini in ntral Pnnsylvarriah pursud th thnology to
inftion(.g.,poor immun
firstdttGA1 via nondgrded protins in rin sampls and thn dvlop
a gntitst sd on mirohips. Th mutation has n trad to urop,
ns b tratd(.g.'nausa, fvr'
whrth Amish originatd, and ours at low frquny within populations
tin allowd o ttn is
of uroparrdsnt. \With srning and dit, disas prvaln has en
rdudamong Amish to a third of th prtratmnt rat.
iqu and/orommon in humans,
Rognizing this rodr ontt is an intgral part of Darwinian mdi-
animals(lldzoooti disese;
in' In part this kind of training was at on tim providd to halth profs-
sionalsundr th sujt of gographi mdiin, ut it was swpt asid in
ftioussour' nd how ws
th silvr-ulltyars immditly aftr th disovry of antimiroial drugs
rtontat r y an arthropod
i n t h 1 9 3 0 s a n d 1 9 4 0 s ( B u r n ta n d W h i t | 9 7 2 ; W . A n d s o n 2 4 ) . h e
rdisovryof this ranh of mdiin oms on th hls of failurs of drug
luld patints isolatd until
thrpisand th rmrgnof inftious diseasslik polio and turu-
losis tlrt wr thought to hav n onqurd (Grrtt 19941 Lvin and
; losstrlativs?Will thrapies
Andrson 1'999).
int as wll?
3nis a batrium,a virus, a
)tiousagnthav rsistanto
Tratmntand Thrapy
this prsorr'sfamily history and
Dwinian mdiin asks hlth profssiorrlsto look yond symptoms of
th patint in front of thm and to prati mdiin in a way that is oth
r t i o nw i t h a h i s r o yo f m i g s
prditivand pmptiv(Zrhouni 2005; ulliton 2006)' Th hallng to
t s i n so r l o s e ) ?
th mdial world, inluding iomdial rsrh and drug disovry' is to
rr popr-rlatinswitlr th s-t
mrotlr rratural sltiorrand ommon dsnt (horn<llogy)as prt of
th pross tht translats iomdial rsh to linial prati. In this
fft ynd this patint?
translation humirns annot st thmslvs apart from thir volutionary on.
;niityor viruln in th
ntions to th natural world, inluding thir own gntis and iology. Th
rins)?
siof volution is highly pditiv,and it is unthial for thos who ar
antimiroilsat,and what
dvlopingtri1tmntsto ignor knowldg of volutionarv iology. s will
ning with a drug now and
.ugrsistanvolvs? b snlatr, som of ths pitfalls an only disovrdin retrospt,ut
th ost of ignoring volutionry iology in mdiin has n high in som
aSs.
ossa thnologialworld whr
i o h n r i s t r a n d g n o l n i s a
rlso prsurnsthat mdil and ADAPA|oN
:y and ology of pathogns and
If on nsidrsmiros' th ida of natural sltionis asily aptd-
d that thy undrstandth thni
us rslstanto antimiroial drugs has apidly volvd and rsultd in
rmrgIlof prviously ontroild pathogns (Grrtt |994; Levin and
long th Amish of Pnnsylvnia
Andrson 1999; Lederrg 2000). In som ass atria hav volvd
]twn iomdial rsarh and
multipl-drugrsistan,partly ausof novl mutations ut just as oftn
d ollagusfirst diagnosed and
y aquiring drug rsistanvi DNA hng from othr atrial spis.
zl syndrom, a form of rral
This is tru fo |'Jisseri,th atium rsponsil for sually transmittd
n arly dath in fftd hildrn
g<lnorrha rrd dirtly trarrsrnittdatrial mningitis (Snydr t al. 2005).
ll syndrom is glutari aiduria
Similarly, multidrug rsistan has volvd in th flsh-ating Stpblocous
l a t i o n p a r h w y so f t h r a m i n o
2 9 0 uolutionr Biollg of Diss nd Drtuii edii
urus' whih is oftn aquird in hospitals. In its nw and imprvious form ar not yt availal
this ommon skin atrium auss svr disas of skin and soft tissus' simulation studis
somtimslading to dath (Dikma t al. 2001; ady and ov 2003). gl drug until rsi
on th othr hand, it is implausil that humans will volv nw mha- gratst halth r
nisms of dissrsistan,at last in th asnof som kind of rgular and th sam ptints
atastrophi hallng that would svrly rdu fitnss of thos who lak with th lowst tr
gnti prdisposition for fighting inftions. Again' this is not to say that son 1.999;'Bull an<
humans hav always n unafftd y th powr of natural sltion- Again, possil ou
rnt gnomi analyssshow ongoing sltionin rain siz (Mkl-Borov ontrolld human
t al. 2005) and rnt sltion for adlt latos mtaolism in uropan provid dfinitiv l
and Afrian populations (Tishkoff et a|. 207). But th disparity of gnra- As notd for inf
tion tims of miros nd hmans spurs natural sltion in favor of mi ult, although sv
ros, whih will xprin sltion at multipl lvls: within individual u s ( . g . ,m a s l s
hosts, twn human hosts, and twn humans and pathogn rsrvolrs possiility that 5
in ommnsal or wild populations of othr animals. human pathogn' :
An intriguing possiility is that our immun systms ar urrntly undf for widsprad usi
sltion via immunopathology, disas usd y ovrrsponsy our im. in will provid s
mun systms to What ould othrwis mild inftions (Graham t al. fortuntly' this
2005). our immun systms ould sn as a sris of trad-offs twn sltion within ht
rognition of slf and nonslf, dfns against invading pathogns, and th high ts of path
potntial damag to noninftd tissus whn laring invadrs from inftd ally' imprft v
tissus. In this as sltion would toward an intrmdiat phnotyp mission to sap ]
that alans trad-offs. Gnomi analyss of immun-systm gns' om- individuals (Gand
ind with primnts in modl systms that furthr dmonstrat how ths halth impliationl
kinds of trad-offs funtion, will provid an ativ ara of rsarh for yars must tak into a
to om (Graham t al. 2005). and tht within-l
Thus modrn mdiin still finds itslf on a shifting landsap, and w on- (Starns and rt
tinu to writ th haptr in human history suggstd y Joshua Ldrrg loal outraks or
(2000)' whih h alls ..our !its vrsusThir Gns.''volution in miros pathogns in th ft
will nvr stop, ut w ar ginning to apply stratgis spifially dsignd Finally, muh ht
to halt thir transmission, slow rsistan to antimiroials, and improv mnt that pathog
vains. Th first issu is prhaps th simplst: hygin an halt pathogn slows. transmissio
transmission. This was immdiatly apparnt from th famous pidmiologi- viruln id om.
al sluthing by Dr. John Snow in London, who in 1854 rmovd th handl uropan raits i
of a puli watr pump on Broad Stret in Soho and provd that a loal the virus was intro
holra pidmi am from a ontaminatd wtr supply (!7ills 1996). Sim. sap. Lowrd vir
ilarly, plagu and th Blak Dath' artsd y th batrium Ysii pstis is sond ampl
and rsponsil for som of th most infamous human pandmis, hav sis that alans vi
largly wand as human disassin aras whr rodnts and thir flas ar r- nition of viruln
movd from human living quartrs (Gag and Kosoy 2005). In trms of vo. within th host, to>
lutionary iology, hygin irumvnts th opportunity for sltion for host, and, finally,
pathogns by simply rmoving hains of inftion and prvnting transmis- Neisseri meigitt
sion (Galvani2003). transmission is fror
Slowing volution of antimiroial rsistan is on of th largst publi ally asymptomati.
halth hallngs w fa, ut most work in this ara is hypothtial aus mission and is atu
ritial data that ould distinguish whih stratgis would most fftiv optimal viruln
diie F,uolutionrBiolrlg of Dises nd Drtuii ediin 291
. In its rrwand imprvious form ar not yt avirilal (Lipsitlr 200l). Rgardiss,mathmatil modls and
dissof skin and soft tissus, simulation studis indiat th;rt in liu of attirking ah mio with a sin-
20I;adyand ov 200). gl drug until sistanrahs a lvl that ompromiss puli halth, th
humans will volv nw mha- gratsthalth nfitswill gaind by dploying omintions of drugs in
snof som kind of rgula and th sam ptints at t sm tim or targting for ontrol thos pathogns
rdu fitnss of thos who lak rvith th lowst transmissi.lt-t rzrts(Bonhoftrt a|. |997: Lvin and Andr_
rs. Again, this is not to say that soo1,999 B u l l n d W i h m a r r2 0 0 1 ; L i p s i t h 2 0 0 1 ; D . L . S m i t h t a l . 2 0 0 5 ) .
h powr of natural sltion- gin, possi1Outomslravnugh ontingnisthat orrly tim, arfully
:tionin rain siz (kl-Borov ontrolldhumarrtrials, and laoratory pimntswith modl systmswi1l
latosmtaolism in uropan povidedfinitiv 11llswrs.
t07).But th disparity of gnra- As notd for influnza A' kping fftiv vins orr hand an diffi_
natural sltion in favor of mi. ult' although svral vains hav rmaind fftivvn aftr dads of
multipl lvls:within individual us (.g.,masl.svins). T.his qustion has arisn again baus of th
h u m n s n d p a r h o g nr s v o i r s possiilitytht H5N1 avian influnza ould mrg as a dirtly transmittd
animals. hurnpthogtr,ird it is not f:rsilto provid prftly matd vains
lun systms ar urrntly ndr for widspr:rdus. Thus th hop is tht vn an imprftly mathd va-
" r s d y o v r r s p o n s y o u r i m - in will provid som lvl of prottion nd slow virl transmission. Un-
: mild inftions (Graham t al. fortunatly, this dos not tak into aount that pathogns undrgo
as a srisof trad-offs etwn sltionwithin lrosts that tluld favor mor virulnt pthogns (dfined y
inst invading pathogns,and th high rats of pathogn rplition alising ll r tissu damag). Spifi-
:n laring invadrs from irrftd ally' imprfr vains mav alrs pathogns to volv oth fastr trans-
rd an intrmdit phnotyp rnissionto sp prottd irrdividuals and high viruln in unprottd
; of immun.systmgns' om- i n d i v i d u a l s ( G r - r d o n t a l . 2 0 0 1 ; v a n B o v n t a l . 2 0 0 5 ) . A g a i n , p u l i
at furthr dmonstrat how ths halth implitiotls of using virinsthat only Partially potet individuals
l ativ ara of rsrh for yars must tak int() ount that sltion or-rperthognsoLlrs at svral lvls,
and that wirlrin-host volution gnrlly Slts for gratr viruln
a shifting landsp' and w on- (Starnsand rt 2001; Galvani 2003). Rlasing partial vins to stm
y suggstdy Joshua Ldrrg loal outaksluld hav th unintndd risk of sltingfor mor virulnt
ir Gns.''volution in mirobs pthognsin th futur.
lly stratgisspifiallydsignd Finally, nrlrhas n nrd f th volutiln of viruln and th argu-
to antimiroials, and improv mnt that pathogns should volv lowr viruln ausdamag to lrosts
plst: hygin an halt pthogn slows transmission (wald 1994). mpirial support for th optinral-
nt from th famous pidmiologi- virulnida o-tsfrom a frv xampls lik th rdud dath rat among
who in 1854 rmovd th handl }-uopanrits ilrftdwith nryoma virus in Austli in tlr yars aftr
Ln Soho and provd tht a loal th virus was introdud t.lontrol th r:ritsthat wr rvaging th land-
l watr supply (Wills 1996). Sim- sap.Lowrd viruln in th trium that auss diphthria in humans
r y th trium Ysini pstis is a sond anrpl (brt and Bull 200). I}utagain, th trad-off hypoth-
[amous human pandmis, hav sis that alnsviruln and trnsmission assumsa rlativly simpl dfi-
rr rodnts and thir fls ar r- rrition of virln, wlrilr is tually a fuIrtion of pathogn rpliation
nd Kosoy 2005). In trrs of vo. within th host' tiity of rh pathogn to th host' immun rsponsof th
r opportunity for sltion for host, and, finally, whih tissus ar afftd. For instarr' th atrium
. f t i o na n d p r v n t i n gt r a n s m i s - |'lisserimenigitidis auss mningitis whn it infts spirrl tissus, ut its
transmissionis from inftions of th uppr sPiratory trt, whih ar usu-
:anis on of th largst puli ally asympt<lmati.hus virr.rlnin N. migitidis is r-rnlatdto trans_
Lthis ara is hypothtial aus missionand is lly a spillovr f inftions within hosts. From th ida of
tratgiswould most fftiv optimal viruln oms th notion that disasould avoidd if pathogn
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Afria and urop. Ntr Gtis 39 \4.
haratristis of l
van Bovn' M.' F. R. Moo, J. . P. Shllkns'H. . d lkr, nd M.
Krtzshmar' 2005. Pathogn adaptation undr imprft vaination:
thy hav daptil
Impliations for prtussis. Prodigs of th Rol Soit lf l-odo, Sris B Howvr, thr is
272:1617-1624. (i..,how th forr
Vitousk, P. ., P. R. hrlih, ' H. hrlih, and P. A. Matson. 1986. Humn ss whry natt
pproprition of th produts of photosynthsis. BltlSie 36.. 368_7.
diffrnt forms in
Wang, L. F.,Z.L' Shi, S. Y. Zhang, . Fild,P. Dszak, and B. T. aton.2006.
tion annot pla
Rviw of ats and SARS. mrgig Iftious Disss 12: 1834-184'
.Wiss, plaining th difft
R. A., nd A. J. Mihae|. 24' Slialand nvironmntal risk fatsin
th mrgn of inftious disass.Nture dii 10: S70-S76. ontribution to th
.!illims,
G. ., and R. . Ns' 199]r'T dawn of Darwinin mdiin. Thr is a tnd
Qurtrl Ruieul lf Bbklg 66: 1'_22. an gnratd
Wills, . 1996. YIlo Fur, Blk Goddss: Th oulutio of Peo1llnd
indfinit divsit
Plges. Rading, A: Addison-Wsly.
!olf, N. D.' ' P. Dunavan, nd J. Diamod. 2007' Origins of mjor human assumption that I
inftious disass.Nture 447 : 279-283. sis of two typs (
Zrhouni, . A. 2005. US imdial rsarh:Basi, translational, and linial spis morpholo1
sins.Jourl of th mri edil Asslitil 294: 13.52-13.58. into distint, not (
onstraints on tht
som forms n
amin in dtail
obsrvational, ant
m p l . A l l t h r a
tion of ontinuou!
dii
1005.Stgi inrritions
istane.Prodigslf th
in
Byondth Darwinian
_1.5 8.
| .S a u n d 2s 0. 0 5 ' o l l l p a r i v
:strvlt thptlrognr
Paradigm:Undrstanding
1 - 218 .
Dises.Nw Yrk: oford Biologial Forms
:althnddiss:
Work in progrss.
Brin Goodtlli
r' andD. H. ortorr.2003.Typ I
7 ptints.
AmriJourlof
\cthirurgiBIgi104:
lssons' highhops.FD
;/faturs/2000/5 00-gn.html. Th qustion that I shll min in this ssay is th rlation twn form
ls of iogography of l-ruman
, Centics36: S21_S27' and funtion in volution' and how th rigin of diffrnt forrs is undr-
ght'. C. Bitt.J. S. Silvrmn' stod. Form hs vo aspts:slrpor strutur' as in tlr morphologv of or.
osr:rn, . Irahim,S. A. omar, ganisms; and pattrn in tinl tlr lrirvilr,suh s th movmlrts of
ad,J. K' Prithard,G. A. !ary, individual orgarrisms(swimming' walking, flying) or th spatial pattrns of
o no f h t r m a n
l a t a s p r s i s t n i n
olltivs,suh as floks of irds or faging olonis of soial insts.Ths
+0. htrtristis of spisr oftn rfrdto olltivly as hratrs' and
. . d l k r ,a n d .
thy hav dirptiv, funtinl signifi:.rn for tlr survivl of tlr spis.
Jr irnprftvaination:
RI Soit of Lolldo, Series B Howvr, thr is a distintion twn undrstandingth origin <lfa form
(i..,how th form an ris i organisms during thir volution) anl th pro-
d P . . N 4 t s o n 1. 9 8 6 . H u t r l ssrvhrynatttrl sltiorrats t hang th rlativ frqunisof th
sis.iSic6: 68-373. diftntforrrrsin poprrli.rtinausof thir survir,lvaIu.N21tulsl-
D a s z a k ,a n d B . T . t o n . 2 0 0 6 .
tiorr annot xplain how any form origir-rats, and it n usd <lnlyin x-
s Disss|2: |84-1840.
plaining th diffrntil aundan of haratrs' onntd with tlrir
t n d n ri o n m n t a rl i s k f t o r s i n
ediill l0: 570_576. ontriution to th fitnssof ganisms'
l n o t ) ritn i n l r l d i i n . hr is tndnyam()ngvolutionirrvbiologiststO ssumtht Ily ti)rnl
an gnratdas a rsult of random r,ltriationin tlr gns,rsulting in lr
luoltio of Popl d indfinitdivrsity of possil forms for volution tl work with. This is th
assumptionthat I shall suggstis mistkrras a gnrl prinipl, ol-rth a-
}07.origins of mjor uman
sis of two t,vps of vidn.Tlr first f. tlrs tlms for osrr,tion lf
isi' translational,zrndlinial spismorphology, whih shlwsthat flrtnsgnratdduring volution fall
:soitiln 29 4: 1 52-1 5 8. into distint, not ontinuous' lasss.An impliation of this is that thr ar
onstraintson th possil forms of organisms and tlrir havilr,so that
sonrforms n gnratd,,lril orhrs annot. TIr as tht I sll _
trrrrinin dtil ms forrrplant morphollgy, sin w hav rimntal'
obsrvational,nd thorti:rlvidnflr distint sts lf pttrns irr this -
m1.ll. All thr ar rqtrirdto mak a onvining s against th assump-
tior.rof ontinuous vriation as a asi prinipl in volutiln.Thr r mn
2L)9
.00 Bod t|l Driin Prdigtn
h
pattrns' som of whih I disuss, ut th plnt ampl mks th grrral
poits larly and illustrats how th argumnt tat rlats to morphology
an pursud in otlrr :rses.
Tlr sond typ lf vidn that I prsnt flr onstraints ln tlr possibl
\
forms availal for volution oms from t ntur ilf th prossSthat
gnrt th forms. Ths ar th dynari ativitis whry diffrnt mor-
phologis and hviors ar gnrtedy intrationswithin orgnismsdur-
ing thir dvlopmnt and twrr.thrnin prdr.ringhirvior pattrns.
This involvs som lmnt of thortil modling, showing what typsof
prossS or'rld giv ris to t flrs rrd rr,lry hs sult in disrt,nol
ontinuous' variation in haratrs.
Th viwplint psntdh in no sl.lshallr.rgs t irr-rportano{ nat-
ural sltionirr volution. Whthr th flrms f organisms ar initilly gn-
ratd as distint pssiilitisor as a ontinuum of variations, sltionstill Figur 1. Th tr
in fushia 1middl)
oprats to influn th frquny of th diffrnt forms <lrdingt the fit-
nss that thy onfr on th individual mnrrslf a spis.Th important
point nrd is that tlr gnrativorigins of iologial strrrtursad pttrns
ar not to undrstood in th sarWay as thir diffrntial aundanin
diffrnt l,raitats.
.W.
ar still t from hilving anything lik a orrrpltdsription of th 90.. Tlr tlrird pt
prosssin organisms that undrli th origin of thir distintivharatr. ah nod nd a
istis. Hwvr, I liv tlrat w undrstnd suffiintl,vwll th asiprin- fom h prdin
ipls of ths gnrativ ativitis to draw som importnt lnlusions th growing tip oI
aout oth th distintions and th rlatilnshipstwnform and funtion turs y growth al
in iology. on rh sis of th xampls prsntd:rnd th gr-rrl prinipls
that govrn nonlinar dynamis, I shall agu that th prosssinvolvd
oRPHoGl
hav th onsqunthat som ftrrrs plssil,whil othrs are not. Un-
oF MoRP
drstanding th origins of th forms that dfin diffrnt spis thrforr-
quirs thrtw ltndstadth dynmi prosssthat gnrttlrm,whih A omrlrnsiv
r not th<lsof natr.rralsltion. Douady nd lrrd
a modl that sho
damntal ida is tl
Phylltais as a Slf-organizing Growth Pross mordium lvill arir
egir-rfalls low
A lassi ampl f form in flowring plants onrns th arrangmntof sourof inhiitio
lavsalorrgth stn-r(phyllotaxis).It has n rognizedfor lnany yarsthat ing. Howvr' th
P t t r n s r. h o u g ht h r r I s os o m i n -
t h r a o l l l y r h r i m a 1 ..( ) l n m ( ) n that as lavs gro.
frquntly found arrangmnts. Th rnajor arrangmnts r shown in Figure an initiatd. It
1. Th first of ths has th forr of a spirl whll on looks dorvn on th that rsults in th .
plant, illustratd hr y yua. Sussivlavs ou1.lyp()sitionsrotatd mristm and its g
throrrgh wll.dfind avrag angl (l37.5o) rlativ to th prvious laf. terns of laf arrar
Th sond pattrn is alld wlrorld, in rvhil-rtwo Or tnor lvsarlst ondary ons. Tlrt
ah nod. h lavsat ah sussivnod ar rotatd so tht th lavsar pattrning poss
loatd ov tlr gps lf tlr prvilusrvhorl. Th xampl slrown is fushia' l trms, a of
whih has two lavs at ah nod' with rotation of sussivrvhorls through dynami prossl
Bed th DriLl Prdigm 01
<a omplt dsription of th 90.. Th third pattrn is knorvn lls listihtrs,in r,l,lrilrth
is a singl laf at
gin of thir distintivharatr- ah nod lld irh sussivlf is loirtdon th opposit sid of th sfm
d s u f f i i n t Iryv l lt h a s i p r i n - from th prding ln. Th gnratolf all ths pattrns is tlr lnristm,
lW som importl-lf onlusions th growing tip of th plant' wh th lirvsar produd as distint stru-
ships twnform and funtion tursy growth and diffrntiationlf Ilsin intration with on anothr.
:sntdand th gnral prinipls
]gu that th prosss involvd
MoRPHoGENE| FILDs As GNRAoRs
ossil,whil othrs i1rnot. Un-
oF oRPHoLoGY
tirldiffrntspisth|.llr_
sssthl1tgratthm, whih A omprlrtrsivxaminatitln of tll gnrativpross in th rristmld
Douady nd oudr (1996) to synthsizth omponnts of this pross intl
a modl that slrtlws how th thr typs of phyllotis ould our. Th fun-
damntlida is that th mristm is a dolain of grwth in whih a laf pri-
,owthPross mordium will aris only if th onntration of n inhiitory fill in this
rgion falls l<lwll thrshold valr-r.ah rrw laf that dos form ats as a
nts onrnsth arrangmnt of sourof inhiition to nighoringtissu' prvntinganothr lf from fom*
ln rognizdfor mn-:rrsthat ing. Howvr, th irrlriit<lr influn drass,ith inrasing distrr,so
r r s . h o u r . ht h : t I s s r t l r i t r - that as lvs erlrv irnd mov aprt' th ar rgions whr laf formation
rrangmntsar slrown in F.igur an irrititd.]t is th pattn of intrlrtionsin this slf-orgnizingpross
al whn on looks down on th tht rsultsin th rrangmntof lavs.Dpnding on th initial stat of th
lavsoupy positions rotatd mristmand its gomtry' th pross gllertson of th thr n-re jor pat-
5") rlativto th prvious laf. trns of laf arragmnt as th primary pattrn and othr pattrns as s-
hihtwo ()r mor llvs is at ondary ons. Th trm usd in iology ro dsri this typ of sptial
] ar rotatd so tht tlr lvs ar pattrningpross irr dvloping organisms is lttorpbogtifild. In thni-
l. Th ampl sho,rr is fushia, al trms' h of th liffrntpattns that ariss from th slf_orgirrrizing
rion of sussivwhorls thrugh drramipross ls alld an ttrL1t()r, rvhil-ris th stal soluti-lllto th
02 Beod the Drluiin Prdigm
)DUFoRs WlH
s
:haratristimthmatial ordr in
S o m a m p l so f t h s r g i v n i n
]mntsof lvsin ruri ls,
,. Th numrs on th
lvs fr to
rtd' 1 irrgth ylungstlaf or th
',
and so on. Th sirals in this figur
,tontwith
ah othr. Now look
of sussivlavslolrg any spiral.
.hspiralsdfindy dottd lins and
c lins.This dfinswhar is knor,vnas
'o ampls
in th figur hv (5, 8)
rnd 2' rsptivly).Th nLrmrsin
wll.known mthmatial sris d.
lunrrof th sisis th sum of th
numrsof th sris dpnd on th
:hth nunrbrs1, 1, th sris is 1, 1'
ost plants with siral phyllotais
t u n r r si t l l i s s r i s ' s w s w i n
) o u a d n d o u d r ( 1 9 9 6 ) s h o w d
: m i n b r n n ho f h g n t i v p o -
th nlrrnr diffrns longirrg to
Figure2. Laf arrarrgmntson ranhs
n a t u r 's r r ha s ( 2 , 2 , 4 , 6 , 7 0 , 7 6 , of spisof onky Puzzl tr,
ruriecls.S tt for disr.lssiorr.
a n d ( 3 ,4 , 7 , 1 0 , 1 7 , 2 7 , 4 4 , . . . ) .
lranhy spifyingpartiular initial
101 Bod th Dt'ii Prdigm
onditions and pramtr vlus in th mdl of Douady and Coudr. Th and Coudr. Th |at
rar pttrns <lfr,vlrorldlaf arrangmntstht ar distintiv to partiular morphogn from la
spis ar lso gnrtd y th rndl lvlrrrspifi oundry orrlitions nw laf to form t a
and paramt vls:rr spifid, f<lringtlrs mods tl ppar' though vidn prsntd y
tlr a lss stal tl-ranth dtlllrirlnt ons in trms of pkig dnsitof dution of primordiur
l:rvs. auin that gtstransp
mation in its nighr
is thus rplad y rr
RLA|VE sABIL|Y oF DlFFRN PHYLLoAI FoRs
inhiitory tion y
A l l t h p a t t r n sg n r a t di n t h m o d l < l fD o u a d y a n d o u d r a r s t a l vtd so that instad .
flr srm rang ()i Lroundary and initial orrditions on th morphognti laf formation in a sl
fild in th tnristm, l)tlt som ar lss snsitiv to hangs in thsthan phogn tht indus l.
othrs' Agin, th ]onrinan t a t t r n s f F i g u r l a r t h o n s t h a t a r l a s t nw laf and its boun
snsitivt viritilnsin th gnrativonditilns in th modl, ad among nw laf inititation irr
ths th spial pattrns ar th most roust. This fild modl of th dy- a fild of inhiition
nmis of laf podution in th mristm thus provids us with a possibl rns whr th paks o
planation of how all th diffrnt laf pattrns an gnratdin highr port of th morphogn
plants; tht is, it givs us a plausil pittrr of how th diffrnt pattrns Th lgnt studi
ar possil forms of th sam dynami pross undr variation in th gn. mophognti filds iI
rativ onditi<lnstht an rlatd to diffrrttgn arivitis and ound- tions. larly thy d r
l l r v o n d i t i o n s l n t h n r r i s t m( m < l n o l tasn d d i t s ) .I t i s n o t n s s a r y o r modl of th whol
appropiirt tO tls th onpt tlf ntual sltion or diffrntil {itnss of dirtional-fild ompt
th harats to plain how th osvd rang of forms ould hvorig- tial loation of uin 1
i n a t d i n p l a n t s . W h t t h m o d l t l l s u s i s t h a t w i t h i n s o m p l a l r tl i n a g that is loalizd to th
lr linags thr volvd within th mristrn a dynami pross with th t r a n s p o r t i n ga u i n p i
gnrtiv haratristis dsid. This modl gnrats thos arrdonly vasular lls of th d
thos pattns lsrvdin known plant s-lis. Thr ar many pattrns n that undrlis th
tat w n inragirrtht might produ1,suh as a singl laf at a nod, to explaind in trm
trvo rotatd at 90" tO this ilt th nt nod, tlrr at th nxt rlttdrlativ morphognti fild. lV
to ths t.uvo,irnlthn rpat of this s]un.Ths ar nvr osrvd. aross ll walls, ll d
Biologial foms ar not sltd fror ny imaginabl st or i] ontinuum m i s t mt o p o d u t l
of possiilitis. Nonlinar dynami prosssof th typs that or'rrin d- osrvd in laf phyll
vlopmnt always gnrata disrt st Of forms assilto th dynami sk to provid n int
prosssinvolvl.It is not natural sltion that dfinsths pssibilitis, Pross. Clarly any r
ut th intrinsi l,vnamisof th gnrtiv pross in th dvlopingor. ahiv th sam rangr
ganism. Undrstnding ths rnorphognti prosssis what providsus nd thi r transt.ormar
,ith n undrstnding of th origins lf spis s distint forms assibl
w i t i n a l i n a g ( G l o l w i n 1 9 9 4 ) . DvLoPMN
WN oRl(
F|LDs AND MoLULAR GNRAoRs
I now turn to th issu
A rnodl suh as that of Douady and oulr shows what priniplsarr. plaining aspts of phy
quird t<lprlvid a andidat xplanatiln <lfphyllotais, ut it oprts t usd to dfin th ada
quit a hig lvl oi stration.Rnt stulisy Rinhardt t al. (2003)in- ism funtions in a par
diat th ntur lt th rrrlulsinvolvd in grrratingphyllltatipattrns with othr nrmrs o{
nd rv| fiId 1r.trirtrri
diffnt in dtil frorrrtlrat essumd y Douady t o t h h a r r e r so f n
Be'odtb Dru,,iittPrdigm 0s
modl of Doudy and oudr. Th
andoudr. Th lattr assumd an inhibitory
n t st h l t l r t ] i s i l l t i v t t l p r t i u I r influn,possily arrid y a
morphognfrom a laf primordium, that
I whn spifi oundary onditions drasswith distan,allowing a
.ingths nw lf to form t ritial distn from
a dvloping lf. Howvr' th
mods to ppa, though vidnprsntdy Rinhardt and ollagus
otls i trrs of paking drrsity of is that th influn is an in-
dution of primordium formation y auin.
A nw lf ts s a sink for
auin tt gts tanspotd away from t sit,
thus prvntin8 nw laf for-
mationin its nighhood.Th irrhiitio.'
Dou,jy and Coudr
jRN PHYLLoA| isthus rplad y rmoval of an indur from ",.u,n.dy
FoRs th nw laf rathr than dirt
inhiitory ation y a morphogn. The fild
:l of Drludyand oudr r stal rul is thrfor fftiviy in-
vrtdso that instd of prmissiv rgions
al nditions on th rlrplrognti of low inhiition that all<rwnw
laf fomation in a salar fiId, w hav lomins
is snsirivt hng.sin ths than f lvatd auin, a mor-
phognthat indus laf initiation, and
f Figur 1 ar th ons tht a lst transport of uin way fom th
n w l f n d i t s o u n d r i si n t h n r i s m
ondititlnsi th rnodi, arld along s r h l a f d v l o p , .p . . u . n . i n 8
new laf inititation in its nighorhood. Th
roust. This lild modl of th dy- dynami. i, -o.. .omplx than
a fild of inhiition austhr is now
mthus provids rrs with possil dirtionl olnponnt that gov-
rnswhr th pks of auin ur in th fild,
f pttrnsan grrrirtdin higlrr arisirrg from polarizd trans-
port f th rorphogn.
riturof how th diffrnt pattrns
Th lgant studis of Rinhardt nd ollgus
j p r o s su n d r v a r i a t i o n tk th qustion of
in th gn- morphogntifilds in plnts in sonre vry intrsting
l diffrrrtgn ativitis nd ound- and lrallnging dir-
tions.larly thy do not plain phyllotaxis,
ltsand diots).It is nlt nssary or whih quirs oth ohrnt
modl of th whol and an xprimntally
ral sltionr diffrentil fitnss of valid moulr dynami. h
dirtionl-fildomponnt tht undrlis uin
lvd rng of fortns ouid hav orig- flow is govnd y th spa-
tial loatin of auxin pumps in th ll mmran,,u.
us is that within som plant linag u, PIN1, a protin
thtis loalizd to th apial sid of lls in
lristm dvnai th orrtrlayrs of th mrsttn,
oss with th trnsortingauxin apilly' ut orintatd for
his modl gnrts tlros nd oniy asa1transport in dvloping
vasularils of th dvloping laf. hus
rrt spis. h ar many pattrns thr is fild of ..g,,l,to.y influ-
nthat undrlisth prodution and loalization
l L r ds,u h s a s i n g l l a f a t n o d ' of PIN1 in lls and nds
to xlaind in trms of molular orgaizing
ld, thrt th nt rottd rlativ fors in ordr to modi this
mophogntifild. Modling th los oupling
squn.Tlrs ar nvr osrvd. twnorintd pumplng
arossll walls, ll diffrntiation,and ,.'"in
a n v i r n l g i n l | \ t o r i l o l l t i n u u m flo- attrns througout th
mristmto prdu th ohrnt marosopi
rsssof tlr typs tht our in d- pattrns of lf arragmnts
obsrvdin laf phyllottrxis prsnts n intrsting
:t of flrmsssilto th dynmi hallrrgfor thos who
sk to provid an intgratd-r'rndrstanding
,tionthat dfins of ti, .nbu,]t morphognti
ths possiilitis, pross. larly n1, modl that sks
]tlvpossin th dvlopirrg or- ro plain ths phnomn ,nust
ah,ivth sam rang of intgratd undrstanding
]ntiprosssis wlrt provids us of phyllotati pattrns
. spis andthitransformationsas th modl of
Douady nd Corrdr does.
s distintfors ssibl
of lavs)and to th lif yl as a it is a disadvantag,it will nlt surviv. Howvr, it may not utility that d-
n to dsrith pross whry trminsdiffrntial aundan.
ir haitat aris in th ours of vo-
ary haratrs aris as a rsult of
(fittr)haratrs inras in fr- olltivBhavior in Ant Colonis
ls haratrs that af lss fit. Th
is ar thrforrgardd as having Th havior pattrns of orgarrismsar larly an important and intrsting
, so that th diffrntial aundan asptof thir volution. Th abilitis of fish to swim, of irds to fly, of in-
s adaptivvalu to th organism. If ststo forag for food' and of plants in tmprat limats to drop thir
of phyllotais pattrns in th lavs lavsin th autumn all ontriut signifiantlyto thir apaity for survival.
piral phyllotais is th pattrn with Just as thr ar diffrnstwn spisin thir morphologis, so thr
of highr plant spis,sin this is ar diffrnsin thir havior pattrns that allow us to rogniz thm as
stalishdhistorially y natural signifiant indiators of th distintiv lifstyls and haits of diffrnt
spis.I shall now tak a look at th way W an undrstand havioral pat-
angmntsin plants? A rasonabl trnsas iologial forms. For this I us an xampl from th study of dy-
n of athing light for phtosynth. nami ordr in an ant olony. Th qun and th young ants at all stagsof
m hav diffrns in th ffiiny dvlopmntar kpt in a sp within th olony alld th rood hamr.
lpnds on how muh thy ovrlap In this hambr thr a many workr ants that tnd th qun and th
rnthat th spiral pattrn illustratd young' fding and laning thm, moving thm out as th young matur'
l shading whn light is oming from and kping th rood hamr fr of dris. Rsarhrswho hav am-
arrangd in rows nssarily shad indolonis kpt in laoratrishav osrvdth ourrnof a rhythmi
uld a sis for undrstanding th ativitypattrn among th workrs in th rood hamr of rtain spis
s in trms of natural sltion and of ants. Whn individual workrs Wr osrvd, ithr in isolation from
r onsidrd,this proposal loss its thir fllows or at low dnsity so that thr wr fw intrations twn
of a day, with th sun oupying all thm, it was shown that thir havior was haoti in th thnial sns
r diffrnstwn diffrnt laf of th trm (Glik 1998). Th qustion ariss:how dos a rgular priodi
om minimal. Furthrmor, osr- rhythmthroughout a olony aris from th intrations of haoti individu-
shows that thr is no orrlation als?Whr dos th ordr om from?
axis, as might ptd if mai- Nigl Franks and ollagus(Franks et,a|' 1990) oservdthat th workrs
spiral laf arrangmnt.Plant adap- in a olony of ants f th gnus L|ltothor who wr tnding th qun and
i o n i n l a f s h p a n d p i g m n t a t i o n ; th young in th rood hamr xhibitd a wll-dfind rhythm in thir
ht frqunisavailabl. Th ovrall ativity-inativityyls. Roughly spaking, th workrs wr inativ for
ltivly nutral in plants, sin all about 10 minuts vry half hour. Th first assumption ws that ah ant is
rious light haitats.In this as dif- individually priodi in its ativity-intivity yl, and that in th rood
lf spiral phyllotais in highr plants, hamrths yls om synhronizd through intrations so that th
rral sltin of a funtional hara- whol hiits a rgular priodi yl. Suh synhronization is wll known
hyllotati pattrns work adquatly in th flashing of firflisor in fmal monkys oming into strus with th
rnspiralsprdominate aus thy full mon. Howvr, whn ol (1991r\xamind th havior of individual
l stlT . hr is noting suprising ntsfrom olonis of a spislosly rlatd to that studid y Franks t al.
' mans simply that w nd to tak (1990)'whih also hibitd rhythmi hvior in th rood hamr, h
ssiility and staility of diffrnt found that thir ativity-inativity yls wr thnially haoti: thy -
th frqunyof haratrsin speis hiitddtrministi haos, without any sign of a rgular rhythm. How'
:s whr th haratrs ar sltivly thn,dos a rgular priodiity merg frlm th intration of hoti indi-
diffrntial aundan is always x- viduals?
it is always assumdthat if a hara- This prolm was studid y Sol t l. (199) y onstruting a omputr
will not inrasin frquny' and if modlof th intration of ants in a rood hamr. ah individual modl
Bod th Drii Prdigm
hirvilr as hoti' in aordan sonrmryosor lrva might gt mor attntion than othrs simply y ei-
: nlodl nts intrtd with ah dntof how th workrs w distriutd in th ood hamr' Dlgado and
v i t ho s r v a t i o l li:i a n t i v n t i n - 5o16(1999)hav shown that whn this is modld y omputr simulation of
rlthassrsult in inrasd ativ- olony ativity, priodi pattrns do indd produ mor vnly distriutd
of moil nural ntwork in whih aIthan haoti pattrns. So w lrav a possil planation of why rhyth-
that ould mov round on a grid mi yls of ativity in th rood hmr ar bnfiial nd may s-
yas not possil to prdit th out- ltd:thy nhan th survivl hans of th yourrg y virtu of good a
lson to assumthi.rthaoti individ- whn thy ar dvloping, and sr inras th fitnss lf th olony. This is
,uld dvlop a ohrnt rhythnr. proalywhy ths ativity yls ar ommonly osrvd in suh olonis.
'. }.urthrmr,th model also on- Howvr, it is lar that natural sltion in no sns plains th origin
rilt into th modl: a rgular prio_ of th rhythmi ativity pattrn in th rood hamr. Th possil fun-
,.hrrth dnsity lf arrts xds tin of th rhythm plyd no rl in dsigning th modl' whih provids an
in his pimntlsrudis.Th p_ undrstandingof how rhythri ativity pattl]sa11mrg in n nt olony
ritialdrrsit1'; ir-rthnil tms' with th proptis osrvd. As statd in th omprhnsiv volum Seff-
..On of th
;ition,lik th suddn ondnsation orgiztio i Biologil Sstms (amazin t al. 2001, 89):
:ur'so that tln kind of ordr sud- rvlationsof slf.organization studis is that th rihnss of struturs o.
sk what kind of planation of th srvdin natur dos not rquir a omparal rihnss in th gnom ut an
risfrom th rptdappliation f sirnpl ruls y larg numrs of su-
units,''odling th ant olony as omplx systlll provids a dmonsttion
of th possiilityf rhythmi havior in th rood hamr. Furtlrrmor,it
R H FoRM
showsthat th dnsity rquird for rhythmi ativity is quit low (aout 20%
ramrannot rdud to th b_ of maimum dnsity)' as osrvd xprimntally,.rrrdth rhythm riss for
intrationsin any spifi ausal a wid rang of xitaility of th irrrs.This is what is rquird for ph.
of individual nts nd th ntu of nomnonto aris in volution: it must lur fairly radily ( assil)and
dit th priodi havior of tlr must roust in th snsthat th should a orrsidralrang of param'
iity is a onsistnt proprty of th trvlus in th rodl ovr whilr th phnomnon ariss. This orsponds
[ris a as of onsistny without to a wid rang of gnotypsin whih th proprty an aris. In addition, th
l ssof mrgnin omplx sys_ fatthat th rhythnr riss suddnly as a phas transition in th olony means
thortrrs will b disovrd that l_ that it is not sltdgradually, with progrssivlyinrsing dfiniion, s
.W
1 sstms.Horvvroths wr not thgadualistviw of volution would hav it. rtinly annot say that
rtd.What th modl maks lar is all iologialproprtisaris in this suddn, unxptdmannr' as do phas
nrgnt pross. Th mathmatial transitions,ut omplity thory is dmonstrating that this is a ommon
:hsproprtis witlrout plaining faturof th way iologial organization mrgs(sGoodwin 19941 So1
il of sintifiprodur: modls ndGoodwin 2000).
laniltitrnsof th phnomna.
ng various asptsof mrgent prop-
MRGN,.oRDR FoR FR''
trtiular,W an sk what rol natu-
of rhythmi ativity of wokrs in It is ths spontanOus'roust mrgnt proprtis that Kauffman (1'993,
ndto ask what furrtion rhythmi 1995)alld ..ordr for fr'' in volr-rtion:
th prolution of som orgnis-
rvsits sr:rvivl.Fnks t al. (] 990) mi morphology or lravior mad lssily th ourrn of onditions
thrshav osrvd tht if an ativ in omplliving systmstht allow th proprty to gnrated.Ths on-
r e t i s a l r a d yg t t i I t g t r n t i o n f o m ditionsris y a random sarh of possiilitisthat rsult from spontanous
o to anothr mmr of th rood. divrsifiationwithin organisms oth of gns and of dynami possiilitis,
rrr ativ at th sam tim, thy inludingth ourrn of haoti ativity' as w hav sn in ants' and
lrood irnd th qun so that thr is fromdivrsifiationof nvironmntl lnditions outsid th organism. Th
tivity pattrns rvr lroti, thn variationof on]itionswithin organisms and irr tlrir nvirol-nntsis what
t 1 0 Bttr]th Drtuii Prdigt
Varitywithin populations and th (\952), ithison (\972), and Jan (1994), Soial insts and thir organiza-
Natural sltion may plain th tion ar disussdin ol (|99|), Sly (1995)' and Godon (1999). volu-
f v n r i u sf r l r l n s h r h v r i s n i n tionary dvlopmnt (also olloquially known as vo-dvo) is th topi of
:hmiativity pattrns in th rood Gilrt et aL' (1996). I disuss ths issus, trying to put thm into a roadr,
rr'thrar situations in whih dif- thialprsptiv,arguing for a holisti prsptiv that rquirs a rthinking
trmsof th dynamis of th !]nr- of onvntional sin t ahiv tru ologial harmony and sustainaility,
of th diffrnt forms produd. In inNtre's Due: Iig ur Frgmeted ulture (Goodwin 2007).
voluti()n'w nd to undrstand
at produth distintivharatr-
r nd th fftsof natural sltion lBLIoGRAPHY
lrs. Th intgrationof ths aras of Arthur,w, 2002' Th mrgingonptualframworkof volutionary
a ohrnt dynami pitur of th dvlopmntal iolgy.Ntur 415z 757-764.
ionry dvlopmntliology. Camazin, S.'J.-L. Dnubourg, N. R. Franks,J. Snyd,G. Thraulaz,and .
Boabau.201. Self-orgiztioni BiologiISstms.Printon,NJ:
PrintonUnivrsityPrss.
ol,B. J.199\.Is animalhviourhaoti?vidnfrom th ativityof ants.
Proedingsof th Rol Soitof Lodo, SerisB 244:25-259,
Dlgado'J.'and R. V. Sol.1999.Slf-synhroniztion and task fulfilmntin ant
ntral onpts usd in iology to olonis.lourl of TheoretiIBklklg 25: 433441..
n. Th partiular thory of volu- Douady,S.,and Y. oudr.1996.Phyllotaxisas a slf-organising pross.PartsI, II,
t, ut thr has n a tndny to and III.Jourl of TheoreticlBilllg1'78:25 5-312.
Franks,N. R., S. Bryant,R. Driffith,and L. Hrnrik.1990.Synhronisation of
ling how organisms ar shpd and
haviourwithin th nstsof th ant Lptothorruorum.Bullti of
tion. A asi assumptirnhas n thmtilBiolog 52 597_612.
s e s u | ot f r n d o m v a r i i o n i n Gilbrt,S. F.' J. M. Opitz, and R.A. Raff. 1996.Rsnthsizing volutionaryand
ids''whih partiular forms surviv dvlopmntal iology.DulopmtI Biolog \7 : 357_372.
rm twnisting spis ar thn Glik,J. 1998. hos:b mzig Sielf the Uprditbl.London:
lss,whih thn omsth sulptor Vintag.
.!idnfld
Goodwin,B. . 1994.Holu the Leoprd hgdlrs Spors'London:
andNiolsn.
.om this y prsntingvidn that
2007. Ntur'sDu: Hlig r Frgmtdultur. dinurgh:Floris
f organismsoftn aris as distint Books.
of possiilitis.Furthrmor, th in- Gordon,D.NI. \999. ts t Work:Hotu lstolo ls rgized.New
lrms prldudan ontriut to th York: FrPrss.
, so that natural sltion is not th Jan,R.V. 1994,Phllotis: SstmtiStud i PItorphogesis.
mridg:amridgUnivrsityPrss.
iffrntialaundan.This vidn Kauffman,s. A. 1993. rigis of rder: Self-rgniztil d Sltkli
from primntlstudy, and from uolutio.Nw York: Oford UnivrsityPrss.
lhologisar-rdhavior pattrns an 1995.t Hom i the Uiurs.Nw York: Oxford UnivrsityPrss.
twnorganisms. MaynardSmith,J., R. Burian,S. Kauffman,P. Alrh'J. mpll,B. Goodwin,
uralsitionwith a disrt st of R. Land,D. Raup, and L..!olprt. 1985.Dvlopmntal nstraintsand
vlution.Qurtrl Ruitaof Biolog 60 265-287.
)istint foms stiil nd to tstd
inlli,A. 2003. The Deulopmtof iml Form: tlge,orpbolog, d
on rgardingfuntion an mad uolutio.amridg:mridgUnivrsityPrss.
ulting in diffrnt frqunis of th Mithison, G. 1972.Phyllotaxisand th Fibonaiseries.Sinc,196:27-275.
i. FIoWvr,undrstanding th pri- Rinhardt, D., .-R. so,P. Stigr'T. andl, K. Baltnsprgr, . Bnntt'
dsto distinguishdfrom sond- J. Traas,J.Frimi' and . Kuhlmir.2003. Rgulationof phyllotaxisy polar
I Want to pursu th issus furthr, auintranspoft.Ntur426:255_260'
Salazar-Ciudad' I.,J. Jrnvall'and S. A. Nwn.rn.2003. hnismsof pattrn
a z i n t a l . ( 2 0 0 1 ) ,A r t h u r ( 2 0 2 ) , formationin dvlopmntand volution.Deulopmt1'30 2027-207.
w-lssidisussilnlf onstrints is Sly,T. D. 1995. Th Wisdom of the Hiu. amridg,A: arvard Univrsity
lis is disussdin Snow and Snow Prss.
12 Bod the Drluini Prdigm
Th dvlopmnt of v
l t a l a h i v m n t st l
major hallngs:how
t i m t o p l a i n d 'l
ganism and nvironm
ths hallngs. Thrt
iota (spiallyif w t
thr ar tainl mi
Donoghu 2004; Dwl
somtims rathtakin1
1'996). vn so, w no\&
divrsity and adaptati
of organismsin popu
diffr, it drivs diffr
ariss in th population
portant part of any org
of diffrntiation. Diff
mntal htrognity.
volutionary ilog1
onrn planations
to xtintion y a mt
.!as
thir tintion a
m a l s ? S o m o n r n r
r s d m o r o | s ss
whn it stilizd or v
tant has mass tintio
nisms that gnrat ad
that produ vriation?
volutionary hang?F
i t t h a t s u h l m n t s.
phvllotais:Diagonal slits through
s <lith Rol Soiet lf Lldo,
Kim Sterln
t1.t
1 4 P h ilo soph of u oltionr Th o ught
othr essaswili hav mad iar' thsar important and diffiuit qustions' also in sltiorr.Tlr fi
ut thy sm to mpirial qustions. Sin philosphs do nt ollt sussful, but thy ar
data, what ould philosophrs hav to say aout volutionary iology tht puzzls aout th ntu
hlps us undrstand it? ow ould irlswrirrgsulr qustiot1sas ths rl- synthesis dpn1don :
vant to philosophrs'o proits? My prlit in this ssay is to ?lnswr yt it was r-rnlarjust h
ths qustions.I gin with th rlvanof philosophy tO volutionayi- hs th synthsis o]
ology; I orrludwith th rlvanof volutionary biology to philosophy. Nonthlss, th on
mor intratal irr th
within iologv. Th ml
Frmnt in volutionary Bilogy: onptual individualism, to th id
InnonLost ual organisms. For a
(1976) agud that sl
.Wiilianrs
h no-Dawinin synthsis w:rs dvlpd y th Ronald A. Fishr nd Dr'vkins
( 1 9 3 0 ) - J . B . S . H l d a n ( 1 9 3 1 ) _ S w lW
l r i g l r t ( \ 9 | , 1 9 3 2 ) t r i u m v i r a t i n mins its indir,idualism
th arly 1930s and was ompltd in Amria y Thodosius Dozhansky th fomidal wapon
( 1 9 3 7 ) , r n s t M a y r ( \ 9 4 2 ) , G o r g G . S i m p s o n ( 1 9 4 4 , 1 9 5 3 ) , a n d G . L d - plagi vrtrats-ar
yrd Stbins (1950) and in Britain y . B. Ford and his shooi (.g.,Fishr tion builds orplx si's
and Ford |947; ain and Shppard 1950, 1'952, 1954 Kttlwll t955, th form th pltform f
1956) for, during, and aftr \/orld.!arII. It bam th rivd viw of tion must linagsol
volutionay iology. lt was individualist, mirovolutionary,gradualist,and tags b prsrvd-tha.
sltiorrist.Th synthsisfousd on individual orgnisnrs:omptirivin. Yt organisnrs dtl not
trations tak plae etwnindividual organisms in loal popultions,not would sh vn if I
twn groups or spis. Individual organisms ar mor or lss fit; A lon of m would n
individuals prospr or fil. Th synthsiswas mirovolutionafy: th lrg- and ths mrks do not
sal hngs visil irr th fossil rord ar nothing ut th aumultionof dnt, I hl, itrst Oll }'
small-salehangs in loal popul:rtions.Th synthsisws gradualist:volu- thousands of gns in
tionary hangs in loal populations ar slow and inrmntl. orss lost of th gns I arry ar
thir tos and grw thi tth littl y littl and ovr long priods. And it was opid' that is, pliat
sltiorrist.han vnts piay :-toie in evolutionary hang. A potntilly ags rflt th diffrin1
favoral mutation might simply just nt happn, or it might happn to an mny gnrations and
organism that is fit ut unluky. But aording to th synthsis,natural sl- short and thin. }.or th l
tion is th lominant atlsal fator that drivs th volutionary dynamis of not itlst rpliators, th
populations. f]urrthi prospts
Th synthsislft fomidal thnial prolms unsolvd, ut th thory odis, th vlrilsin
of volution onivd this way smd rlativly onptually unprolm- tory of lif is tl s it as
ati. This onptual lirrity was not omplt. For mpl' undrstanding ing to sur th rsot
th natur of spis smd a prtiuiarly prssing prolm, for th vry Dawkins 1c76. 1982).
gradualisnr of th synthsis smd to srr!]gstthat th apparan of wll- his lin of thought l
dfind spis might an illusion gnratd y th lngth of our lif spans. to muh ontmporary
Thy sm stal and wll dfind only ausw do not liv long nough g n s I i t l na g n u i
to se linagshang and diffrntit.If w had n around to s th an- Thr sm ro ll t ls
stral at population gradually hang and diffrntiat into its array of gn sltiorr hav n
modrn forms, prhaps w would not think of lopards, jaguars, panthrs' l:rnstrspons wlul
and snow lopards as distint, namal units of natur. Thr wr othr is- tint and suprior altr
sus too. for xampl, whthr volution is gnuinly indtrministipro. prov thir rplition
ss ausof th rol of han. not iust in th snration of variation ut uild a wll-adptd
P h iklslph of ullutitlr h ought ll.t
.h
importantand diffiult qustions' also in sltion. fittr organisms in a population ar mor likly to
Sin philosophrs do not ollt sussful,ut thy r not rtain to mor sussful.Thus thr ',vr
Y aout volutionary iology that puzzlsaout th ll:ltur of fitnss nd its ausurlrll in volutiln. flr th
ring suh qllestiollss thse e rl- synthsisrlpnldon distintion tr,rr tlrl nd petd fitnss,and
projt in this ssay is to answr ytit Was unlar just how this distintlOnWas to drwn (So 1984).
: of philosoph,vto volutionary bi- Thus th synthsisof volutilnarythory did pos onptual qtlstions.
llutionary iology to philosophy. Nonthlss,th onptualgogrphy lf volutionary iology atlrmuh
mor intratal in th 1970s and 1980s as th synthsis am undr fir
within iology. Th most systmatihllngto th synthsishas brrto its
onptual individualism,to th ida tht the targtsof sltionar, lusivly' in1ivid-
ual organisms. For x:rmpl,G. . Willils (|966| and Rihd Dirr'"'kirrs
(1976)argud ht sltiondos not rll at rr irdividualorgenisr-ns t all.
.lopd y th Ronald A. Fishr \ililliams nd Dawkins laimd that tlr gradualisnr of th snthsis rtnd-
Tright(I91, 1932) triumvirat in mins its individrrlism. ompl iologil adaptations-sllsoy systms'
nriay Thodosius Dozhansky th formidal wpons of prdatlrs,th powr nd stranrliningof grat
mpson (\944,195), and G. Ld- plagivrtrats-ar uilt inrmntally' n small stp at a tim. Sl-
B. Ford nd his shool (.g.,Fishr tion uilds omplx systmsumulativly v prsrvingsmll improvmnts
i0, 1952, 1954; Kttlwll 1955, that form th platforrn for furthr small improvmnts. So th targts of sl-
r II. It am th rivd viw of tion must lingsof nar-prftopis-flr only thus an smll 1vlrn-
mirovo1utilnay, gradualist' and tgs prsrvd-thilt prsistthrlughmany vls <lfsltion.
i v i d u a l o g e t r i s n r s:o m p t i t i v i n - Yt orgtrnisnlsltrnt>tform linags'l' daughtr is not i1oP}.tlf m, nor
lrganismsin loal populations' not would sh b vl'l if I wr n asul rtur rproduing [r Ioning.
organisms ar mor or lss fit; A lon of m wlrrldnot a nar-prft()py' for lif's aidnts rark m,
was mirvolutionary:th larg- and thsmarks do not rappar on a lon' As a rsult of a hildhood ai-
] nothing ut th irumulation of dnt,I hav just ln y: a llnwould hav two. In ontrast' though' mny
.h
synthsiswas gradualist: volu- thousandsof gns irr my daughtr ar opis of gns in m; likwis' rost
slow and inrmntal.Horss lost of th gnsI arry r opis of parrrtalgns.Gns, unlik orgnisnls,.tr
and ovr long priods. And it was opid,tht is, rpIiatl,and thy do forrn prsisting linags.Ths lirr-
volutionaryhange. A ptntially agsrfltth diffring susslf gs:s()rtar dp ni tlshy. lasting
happn, o it r-niglrthappn to an manygnationsirnd isting in man'vo1rjg5in ah gnratiotl.othrs ar
ding to th synthsis,natural sl- shortand thin. Fo t most prt' thsdiffrnsar no aidnt:glls r
ivsth volutiorrarydynamis of not just rpliatos,thy ar ativ rpliators. Thir own h:rratristisin-
flunthir prospts for repliation, normally through thir influrrln
prolmsunsolvd, ut th thory bodis,th vhils in whih thy rid. Th ight Wy to think aout tlr his-
.lativlyonptually unprolm- tory f lif is to s it :ls a giant struggle tw linagslf gns,ah try-
plt.For xampl, undrstanding i n g t o s u r t h r s o u r sn d d f t l r r p l i a t i o n ( W i l l i a m s 1 9 6 6 , | 9 9 2 1
ly prssingprolnr, for th vry D w k i n s| 9 7 6 . | 9 8 2 \ .
ggstthat th irpparan of wll- his lin of thorrghthas aisd v,vdiffiu|t issus indd that r ntral
rtd by th lngth of our lif spans. to muh ontnrPorarphilosophy of volutionary iology. In wht snsis
ausw do not liv long nough gnsltilna gnuin altrnativ ttl th individualism of th s1'nthsis?
w had n around to s th an- Thrsmto at last thr answrs to this qusti()n'and th dfnlrsof
and diffrntiat into its array of gn sltion hv nvr rally pikd thir rspns. h simplsr and
ink of lopards, jaguars, panthrs' lnstrsponswtltrld to insist tht gn sltion is an mpirillr' dis-
nits f natur.The Wr othr is- tintand suprior altrrrtivto individualism. Thor-rghgns normally irrr-
i s a g n u i n l yi n d t r m i n i s t i p r o - provthir rpliation prospts y hlping tlr lth gns in an rryo
t in th gnrationlf variation ut uilda wil-daptd ody that will r thm all intl th nt .r.ltion'
)1() P h ilo soph of u olutior h ough t
thy do not always rpliat so ooprativly.For ampl, som gnsan think of th iologial u
gt to th nxt gnration only if thy ar in an organism of th right sx. Th O n s u h p h n o m n < li
gns in th mitohondri in my sprm lls ar doomd, for mitohondrial linags mostly rpliat
gnsdsndonly in th fmale lin. y darrght'smitohondril gnsr ls? Givn tt thy do.
all opis of gnsin hr mothr. In many irumstnsorganisms ar fittst of organisms, in partt
if th s ratio of thir offspring is 50-50 or thrabouts. But gns that d- singl-lld stg? Dw
snd via only on sx hav no intrstin th ody of whih thy ar a part yls, and, indd, s<;m
having offspring of th othr sx, and ths gns ar undr sltion to dis- is possil to formulat t
tort th sx ratio in thir favor, vn though this is maladaptiv for th or- haratristi llf yl fr
ganism of whih thy ar a part. Many suh sx-ratio distortrs ar known sltion maks that qu
(Burt and Trivrs 2005). S-ratio gns and othr so-alld gnti outlaws xampls of th advanta
prompt th following rspons: gn sltion is prditiul differt from Th issu of gn sl
th synthsis. Gn sltion prdits that phnotyps that prss th fittst pt of th dat aout
gns will om mor ommon ovr tim; th synthsisprdits that ph. losophy of volutionary
notyps tht hratriz th fittst organisms om more ommon ovr that oopration is ntl
tim. Although ths prditions ovrlap sustantivly, thy ar not idntial. al systms ar ompos
any hav rspondd y apting that whn gns ar outlaws' W to hav in ways tht
should indd s thm as targts of sltion,ut thy tak ths to x- qustion, thn, is how n
ptional ass that do not undrmin th gnral validity of individualism. tion on th individuals tl
Sltion dos t on gns' ut only in vry spial onditions. So a sond David Wilson' StphnJ
way of dfnding gn sltionis to apt that it is, frrth most part' pr. at on individuals, ut t}
ditivly idntial to th synthsis, ut that it appals to distint undrlying spis (Sor and \rVils
ausal struturs. Twl thoris an lad to th sam prditions aout o- vidualism of th synths
srvaloutoms vn whn thy posit distint undrlying ausl prosss. gadualism. Arding t
lliott Sor and David \7ilson, in thir hallng to th synthsis, apt lif ar just aggrgation
that individualist viws of th volution of oopration an mak th right srv volution in tto
prditions, ut thy argu that ths misidentify its ausal asis. oopra. rabit immun systmsl
tion, thy say, volvs only whn and ausooprativ ats nhn th oming lrgr in dry s
fitnss of th groups that ontain ooprativ agnts (Sor and Wilson all th mhnisms of
1998). Likwis, with rgard to gn sltion, fit gns ar fit aus thy that ar hanging in sm
t on thir nvironmnt in ways that nhan thir proaility of rplia- and lots of tim to au
tion. Th usual mthod gnshav of nhaning thir rpliation prosptsis tns of th fossil rord
by taming up with othr gns to produ vhils, whih in turn rprodu' In a famous 1972 ess
rpliating th gnsthy host. Most of thsvhilsar organisms,ut pr- fossil rord dos not st
haps they an hivs, nsts, and symioti allians too. Fit gns mak thy maintind that th
wll-adaptd vhils, whih in turn rpliat thir uildrs. Th fat that trpolationism to tru
gns typially adopt this indirt Stratgy of rpliation rings th prdi- spis do nlt hang
tions of gn sltion into lin with thos of th synthsis,ut th various spis om into ist
piturs of th ausal strutur of sltion (gn' individual organism) r- thy split into daughtr
main distint. dos dsri th typla
A third lin is to rgu that gn sltionhs huristi advantagsovr th r o | | | 9 9 l ) . B u r i | : l d r
synthsis.Stritly spaking,it is rrot a diffrntviw of th ausal struturo{ xtrapolationism. If v
volutionary pross;instad, it nals us to s possiilitisand prolms hangs in loal popula
that th synthsisonals. Dawkins took this lin in Th teded Pho- than long piods witho
tpe (7982), his most important dfnsof gn sltion.H gav many x- is tru that thr ar l
ampls of volutionary phnomn that ar simply mor salint whn w ass; poptrlations an
Philosoph1'of ullutior Thought )t/
ly. For xampl, som gns an tlrirrko{ th iolgizrltlnivrsas n nsml of ompting gn lings.
r an lrganismof tlr right sx. Th on suh phnomnon is th volr-rtionof th organism itslf. Why do gn
ls ar doomd, fo mitohondrial linagsmostly rpliat indirtl,v,via th ooprtiv onstrution of vhi-
tlgtr'smitohondrial gns are ls?Givn that thy do' why do vhiles hv th distintiv haatristis
lirltmstans organisms ar fittst o[ organisms, in partiular, a lif yl that involvs rprodutiotr via a
tlrrabouts.But gns that d- s i n g l - l l sdt g ?D a w k i n s p o i n t s O u t t h a t i t i s a s 1 ,t o i m a g i n o t h r l i f
th od1,of whih thy ar a part yls,lld, indd, sonr of ths ltrntivsirr al, rhough unommon. It
g'nsar undr sltion to dis- is possiltl forulatthis qustion out th oigin of organisms and thir
g this is maladaptiv for th or- haratristilif yl from withirl th prsptivof th synthsis,ut gn
h s-ratio distortrs ar known sltionmaks that qustion insapal, and Dawkins givs marry othr
nd othr so-alldgnti outlaws xamplsof th dvntagsof th gn-sltionprsptiv.
tion is prditiul diffrent from Tlr issu of g sltionand its rlation to individulism is iust On as-
pnotypsthat prss th fittst ptof th dt aout th units of sltionthat is now ntral to th phi-
; th synthsisprdits that ph- losophy of evolutionary iology. F,volutionary iologists hav onr to s
isms om mor ommon ovr that loperationis ntral to th volution of mplxity. ompl biologi-
rstantivly, thy ar not idntial. l systmsar onlposd of individuals, and ths :rroftn undr sltion
rat Whn gns ar outlaws' w to havin wys tlrilt undrmin th intgration of th olltiv. A ky
'in, ut th tk ths t - qustiol1'thn, is lrow nw lvls tlf oganizatiolrvolv in th fa of sle_
gnralvalidity of individualisrn. tion on th individuals tht mak tlrm up to go thir own way. lliott Sor'
r 1 s p i a | o n d i t i o n s .S o s o n d David lfilson, StphnJay Gould, and othrs hav agrdthat sltiondos
)t that it is, for th most part' pr- at on individuals' ut thy hav argud that it ats as wll on groups and on
rt it ppalsto distirrtundrlying spis(Sorand !ilson 1998; GoLrld 2.002).Furthrmor' iust as tlr indi-
o tlr sm prditions aout o- vidulismof th syntlrsishas isd diffiult npt:rlissus,so too hs its
stintundrlyingausal proSss. gradualism.Aording to th syntlrsis,lag-salpttrns in th history of
l r | I n gro t h s y r l t h s i s 'a p t lif ar just aggrgationslf small-salhangs. Biologists an atually ob-
lf ooprationan mak th right srvvolution in ation in loal populations: thy an osrv hangs in
idntify its ausal basis. oopra- ritimmun systmsas rits ar posd to a nw virus, or finh aks
us ooprtivats nhan th bonringlarg in drv sasons.Aording to th graduzrlismof th syr-rthsis,
:rtiv tlgnts (Sor and Wilsn all th rhanisnrstlf volutionar1.hang ar rtrvork in loal popultions
:tion, fit gns ar fit aus thy tat ar hanging in small Ways ovr a fw gnrrtilns. ! just nd spa
hn thir proability of rplia- and lots of tim to aumulat th rsults of ths mhanisms into th pat-
n i n gt h i r r p l i r i o np o s p t si s trnsof th fossil rord.
vl-rils, whih in turn rprodu, In a famous 1972 essaNils ldrlgeand Stphn Gould argud tlrt th
..trirpolationist''
:svhilsar rgatrisms,ut pr- fossilrord dos not sLlpport this itr.tr'In partiulr,
oti allianstoo. Fit gns mk thymaintaindthirt th tr of lif dos not hang nough ovr til for -
iat thir uildrs' Th fat that trapoltionismto tru (ldrdgand Gould 197z|. Thy argud tht most
y of rpliation rings th prdi- spisdo not hang signifintly ovr most of thir lif spans. Instad,
of th synthsis,ut th various spisom into xistnrlativl,vquikly and tlrn do not hng until
n (gn,individual organism) r_ th1'splitinto dughtr spisor om xtint. Tlr tnt to whih this
dosdsritlr typil lif histtlr,vof a spisis still t>pnto dt (ar-
n has huristiadvantagsovr th rolr|1997|.But if ldrdg and Gould ar right, tlrir pitur dos hallng
rntviW of th ausal strutre of trapolationisrn.If volutionary pilttrns r nothing ut th aggrgat of
s to spossiilitisand problms hangsin lol populations, w w<luldpt tO s gradual hang rathr
' this lin in Th teded Pbeo- thanlong priods without hang intrruptdy sudlnspisformtion. It
f gnslti<ln' H gav many - is trr-r that thr ar loal prlsssth:lt n rsult in stasis in partir.rlr
a simply mo salint whn w ass;populations an trk tlrir lritat v shitiing glgraphillyrathr
r
I
I
!
than adapting in pla; adaptations an havioral rathr than morpho- volvd in rthinking
logial and hn n invisil to us. vn so, if th fossil rord is nothing tual issus. Th nt
ut population-lvl volution summd ovr larg quantitis of spa and tant prolm aout t
tim, w would not xpt stsis trl its dominnt pttrn (ldrdg 199.5; biology.
Gould 2002).
Gould has dvlopd a nror radial ritiqu of xtrapolationism that r-
jts oth its individulism and its gradualism. In a lassi artil Lwonin volution within
dfind th ida of a Drii popultio: a populatin of individuals that
diffr on from anothr and xprin diffrntial rprodutiv suss, In th past dad oI
and in whih dsndants inhrit thir anstors' distintiv traits. Suh intgration of voluti
populations volv undr natural sltion (Lwontin 1970). Gould argud multifatd. It aris
that spis groups form Darwinian populations (Gould 2002; Gould nd onnts vnts in lo
LIod 1999). Spis hav dtrminat origins and ndpoints. Dughtr spis. It ariss in dt
spisrsml thir anstors:ronstrutingphylogny would e impos- playd littl rol in l
sil if this wr not so. Spis ar diffrntially sussful.\Within a r- twntith ntury. As
gional iota som spis linags will diffrntiat rihly, radiating into tal iology rmains hl
many nihs' whras othrs will rmain rstritd in numrs and logi- ar oth volvd anir
al pntration. Gould dos not pt spis-lvlsltionto plain th logial sins and t
adaptations of organisms, for ampl, thir prptual systms, ut h thy nd somhw
thinks that sltion will plain haratristisof spisthmslvs:suh about human agn
proprtis s gnti variation, population struture' nd gographi rang I shall rifly introdu'
.shat
ar visil to spis-lvlsltion and hn n th targt of spis- is gn? Tl
lvl sltion. twn th lassial g
Gould's idas aout spissltionfa srious prolms. For on thing' and molular biolog
thr ar no onvining as studis of spissltionin ation. Morovr, sial gn of th frui
thr ar srious thortial hallngs to his idas. It is y no mans lar fft and was trans
that th inhritan ondition on Darwinian populations is satisfid.It is quns with oth
tru that th organisms in a daughtr spis rsmlth organisms in th DNA squn-hor,r
parental spis.But th spis-lvlproprtis of th dughtr-its distri- snsitiv to the loal
bution and population strutur-may not rsmlr thos of its parnt at off and an vn
all. Furthrmor, in thinking of spis sussand failu, w my not asn of rprssor
abl to distinguish drift from sltion.Th sussor failur of indiuidul rgions nd not ;
orgnisms is not always th rsult of sltion.Som mortality is th rsult not passd on wh
of unhappy aidnt rathr than ill dsign. Som rganisms ar fund y Numann-Held 199\
luk, dspit thir dsign flaws. Hn ralizd fitnssdiffrs frorrr ptd strat away from thi
fitnss.Sussdu to systmatifatursof th population's nvironmntis and instad think of 5
sussin virtu of petd fitnss,as whn a swallow dodgs a hawk. Su- itan: unlss sus
ss du to frak aidnts-ing lown y a storm to frightr that is that mak thm su
hading in th right dirtion-is drift. But th distintion twnptd adaptivly ovr tim i
and frak auss of mortality dpnds on th istn of a population of flow of auratly o1
vnts that is larg nough for thr to pilttrns in thos vnts.Howvr, tionary iologists oft
spis population sizs ar so small and yls of sltin r s fw that formation aout th
W may not al to draw th distintitln twnsltivand aidntal gns is ditd y nat
mortality. gnftion and unsui
I hop tht ths ampls hav gun to xplain th rlvanlf philoso- w an think of th 1
phy to volutionary thoy. It is rlvant us tlr mpirial issus in- aot organisms an<
P h iloso1lbof uolutior Th ought 19
bhaviorrrlrathr than mlrpho- volvdin rthinking th synthsisar not larly sepaatdfrom th onp-
:n so, if th fossil rord is nothing tual issus.Th nxt stion makes th sam point in disussing on impor-
vr lrg qllantitis of spa and tant prolm aout th rlationship of volutionary thory to dvlopmntal
c o m i n n tp t t r n( l d d g 1 9 9 5 ; biology.
)rlattO onsidr ountrfatul s_ So thos who think that humans ar oth thinking ings and part of th
mor srious ttntion to al ph. naturalwlrld nd to plirin th ntur of mrrtlsymbols nd th rlation-
nts ar oftn undrdsrid. Thy shiptwna syrnol and its targt.Y/hat fats out Ptr'srain, his lif'
'al ass.orovr, many of ths and his nvirnmnt mk it tru that a partiulr intrnal fatur of his
i n t t t i t i < l tpl s.r t l y b a u s i n t u i t i v
mindis a sylnol for spidrs?Philosophrshav xplord a rng of rsponss
d s r i tsh t h o u g h x p i m n t s . to this qustion.An intuitivly appalig ida is that symols rsembl their
lvs S/.rr r e k-st|e tltransportrs targtsin th world. Ptr'smntal stat is aout spidrs if it rsmlsspi-
ruitis not th sam as th ontinu- drs.Tlris is a natural su8gstionif thtlughts ar lik imges, maps, or dia-
t is sy to tll ths storiS so that grars.Howvr, almost rainly it Will not do. ntal imags do rrot sm
,tuallytravling whil laving on's to b ojtivlysimilar to nduring thr-dimnsionalphysial animals; th
k t-rsm lik killing mahins appran of ral similarity twnpiturs and things is an illusiln gnr'
tn impostor. This snsitivity to th atlb1'thwav lrumans irrtptpiturs.Ptr'sspidr imag wili rsml
dtailsmattr. Vry th stting, and thrthughtsmuh mor than it smlsany animal. orovr, vn if
in th dtails' nothing ut th ral sommntal stats ar lik imgs, surly most mntal rprsntatlonsar
not imaglik.Whirt is th inragthat ospondsto th thorrght ..Aristotl
rot fully onsidrd th rr-rgof a- Wasth foundr of logi, arrd ulid tlr foundr Of gomtry''?
nntt'swork is important in this r- So th rlation twn symol and targt is not a rsrlanrlation-
Lsallowd philsophrs to rpatdly ship.But prhaps it is a usa] rlationship.Som vnts r natural signs of
a ontinuum y rrgltingth rang thrs.Think in partiulr of instrurnts.An instrumnr is ful gaug if
rg aut rationl gny' philoso- thris a rlial latiorrtwn th rading on th gug and th ful in
intntional agny with mrly m- one'stank. That is what maks a symol on th gauge man ..quartr full.''
lti<lnall1.
snsitiv rspons to our Soan intrnalstatt>fPtr is ir spidr syrrrlrolonly if it lialy ovaris with
natiityof artifts and insts. Dn- thrbinga spidr in his viinity. Th symol-targtrlationship is th rl-
y showstht thr must a rang of tionshiptwn n innr strutrrrand its nvironmntal orrlation: th
snsitivityis ompl adptation, featurof th nvironmnt that aussth innr struturto light up. But this
tror rigid systms in ir singl stp will prolynot do ithr. [n th first instan,this pitur applis i1t st to
prption.Lots of my win thughts our prisly aus thr is no
;y within philsophy is as a soure of winaout. But th runh prolm is that this piur sms to mak it im-
i) has pplid volutionry modls to possilfor us v to mk r-t-tistaks,
t surly w do. If Ptr is somwhf
Millikan (|989a,1989), David Pa- arahnophoi'arrd his frirrdsplagu him with lvr imittion spidrs, h
ptd to us th iologial notion of rvilloftn think ..Spidr!!''whn thr is no spidr around. H is misrpr-
hilosrlphv:th natur of mntal sym- sntinghis nvironmnt. Yt aording to th ovariation pitur t.,f th
nttiosof th ''vrld.But wht in th symbol.tagt rlationship' lr would no misrprsntirrgat all, ftlr his
livthat tigrs ar th most hand- ..spidr''
thoughts ovry with th atgory of spids or imitation spidrs.
vhat maks my tigr thoughts bout Hnthy ar n()tsymols of spidrs atir ll; thir targt is th roadr at-
lturswithin m,v rin a smbol of goryof spider look-liks (inluding ral spidrs, ur spiders, nd so
ig prolm for arry physilist thory forth).And sin tr thirrks..Spidr!!'' only in th fa of spidr look-
: nonistntljtsand abstrat o- aliks,h has mad no mistak.
rlatiorlshiptwn mind and th David Papinau and Ruth Millikan hirv suggstdan altnativviw of
rlationshipsto ngls or numrs. thrlationshiptwnsyrrrbolsand rhir targtstht is zrsdon iologil
v notid that tugh prolms aris funtion.Th iologial funtion of, say, th pattrn on oystrathrggs is
'rt spidrs,for thr is no sirnpl rl- onaImnt. Thos ggs are mouflgel.ggs that wr diffiult for pda-
spidrs ing prsrrttO th thinkr. tolsto spot against typil akgoLrndsof sand nd dris w mor likly
spidrs'for h n tak a moth for a to hath than thos that w sir to find. Tlris sltiv histoy is what
lughtsvn in th prsnof a spidr maksit tru to say that th iologial funtion of th pattrn is amouflg.
on any partiular gg th pi1ttrnhas that iofur-rtionven if it dos not
24 bilosoph1'lf uolutirlnr Tbtlught
lnt fr of any trrstrial prdator In onlusion, philosophy of volutionary iology ats as a typ of two-
its pry y smll. But ology and Way pump. It pumps philosophy into volutionary thory, for thr ar
mak pris quantitativ prdi- many diffiult prolems of intrprtation' larifiation, and integration that
, and on of thm is th fragility of aris out of th xtraordinary dynamism of th lif sins ovr th past
island iota to tintion. But gn. ntury.And it pumps volutionary iology into philosophy: a stram of -
'ionlssnor quantitativ:thy ar ampls,tools, and modls that an prhaps adaptd to driv forwrd phi-
ts smto disovr ausl mha- losophy's own projts. If you want to rad mor, I suggst you turn first to
lusion' ologial sussion, nd th witings of Rihard Dawkins: The tedd Phnotpe (1982) is his
fftsof ths ausal mhnisms finstand most sutl df of gn sltin;limbig ount Improb-
ry ovr spa and tim. So th dis- ble (1996) is a sustaind and onvining dfns of grdualism. Both are
into xptionlss rgularitis that autifully writtn' though Th tded Photpe is quit thnial. For
hosmhanisms.Thr ar gnr- Gould's viws, see Tbe Structure of uolutionr Tbeor Q002),It is p-
; animals tnd to gt smallr, and pallingly long ut rasonably radal, and a good dal of th purly histor-
a r m a n y x p t i o n st o t h i s g n r - ial matrial an skipped. Drtuiism uoluig' y David Dpw and
Bru \Wr(1995), is a good vrviw of th historial dvlopmnt of
nifiation of sin fits iology synthsisand post-synthsisvolutionary iology. Two ooks y Dan Dn-
dxd, or wr thought to indx, a tt' Drlui's Dngros Ide (1995) and Freedom uolus (2003), ar
was itslf rsponsil for th ph- splndidampls of valu of volutionary insights within philosophy itslf.
r hmistry it is likly that th rla- S d Deth: ltroductio to tb Philosoph of Biolog' y m and
proprtis is vry oftn omplx Paul Griffiths \1999), ovrs many of th issus of this ssay and othr top-
lvious in iology, for ll iology is in muh mor dtail.
l th modl of rdution suggstd
iral ftoris:thir iologial a-
lls.To that tnt physis rally is BIBLIOGRAPHY
;hip twn' say' th onpt of a
lar mhanisms rsponsil for th Burt,.' and R. Trivrs.2005. Gsi oflict:The Biolog of SlfishGeeti
lmts'amridg,MA: arvard UnivrsityPrss.
)trozygousallls turns out to
ain,A. J., and P. . Shppard.1950.Sltionin th polymorphiland snail paa
of biology is not a sris of revolu- nmoralis.Hrdit4: 27 5-294'
'logy oftn sms to manifst inr- 1952'The fftsof naturalsltionon body olou in th land snail paa
:ir rvisions and fals trails' ut th nmoralis.Heredit6: 217_231.
tis, or vn volutionary iology 1954.Natural sltionin pa.Getis39:89-716.
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ambridgUnivrsityPrss.
ins.In shrt, it is hard to oth
raraft,J., nd M. Donoghu'ds.2004. Assembligth r of Life. Oford:
n and apt a onption of si- oxford UnivrsityPrss.
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ls a onsqunhav aptd th 1996.limbing out Improbble.Nw York: Norton.
204.Th Astor'sTle: Pilgrimgto th Dtu of Life. London:
not quit up to snuff (Rosnrg
Widnfldand Niolson.
ontmporary who rjts th im- Dnntt, D. . 1995. Dn.ui,sDgrlusId:uoltkl d th igsof
th gounds that it fails to fit iol- Llfe. Nw York: Simon and Shustr.
vork in philosophy of iology ar 23.Fredomuolus.Nw York: Viking.
is issu:thy think that evolution- Depw,D., nd B. H..!r.\995. Dnuiism uoluig:SstemsDmis d
;l sin,and hn th physis- th Gllgof Nturl Sltio.amridg'A: MIT Prss.
Dobzhansky, T. \937. Getisd th rigin of Sples.Nw York: olumia
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,lphof Si67:214_218.
ipcis.Nw York: olumia
hilosoph 86 281_297.
Philosophoi Si 56: 288_302.
ms i t Llgi of Sitifi Disour.
two wll-kno,n nd <
Spnr, Ptr Krpol
soial thinkrs who in
trs in suh rnt o<
e: Dsigig Ntu
volution and Soity familiar popularizrs
Rathr, w xplor l
soity v sking o
NInfrdD. Lbihlr nd
Jn ienshin Dfinitions
B uoltio W ma
ral volutionary id
hang of population
volution and soity ar onntd in many diffrnt ways. First and for. and lad to divrg
most' humans' as a podut of volution, liv in divs soitis.Ths ar adaptd t thir nvi
an outom of volution, alit in mor ompl and oftn mor indirt ally, with variations <
ways than th voltion of dirt morphologil nd physiologial adapta. and spis nd popu
tions. Sond' th initial dvlopmnt of volutionary thory ourrd in stant thms: l[ o
spifi soial miliu onduiv to this prriular intlltual endavor, and prnaturalism as a (
susqunt dvlopmnt of volutionary thory has also rfltd its volving nvironmnt tlrt in
historial sttings. volutionary thory, lik any othr sin, has thus n vord ar etter dp
th produt of a partiular human soity.Finally, and as proaly th most tions that mattr for (
visil onntion of volution and soity, spially in th Unitd Stats,w larning btwen gn
find surprisingly prsistnt dats aout whthr volution or som form of mission.
.W
rationism aounts for th origin of spis (s ugni . Sott disusss also shar th r
in th main ssay ..Amrian Antivolutionism: Rtospt and Prospet'' in ory of volution is t
this volum). som tims in th o
At som tims, in som plas, and in som wys' soity has nthusiilsti- othr of rhsthoi
ally mrad volution; at othr tims nd plas and in othr was, soi- ity, struggl ;rrrdom;
ty has not. In th pross euoltio has mant diffrnt things, as has all part of volution
soiet..W thrfor start with dfinitions and thn look at soity's impat onsists of irrtronn
on volution in two ways: th way in whih soity in th sns of soial' ul- this ssay.
tural, and intlltual ontt has shapd volutionary thory, and also th B soit w dsig
soial rption of volution. Thn w turn th dirtion of influn around th rlvant lmnt
and addrss how volutionary thory today and in th past has ontriutd ft many othrs. In t
to planations of soity.In onlusion w distlssurrnt rlations btwn tingnt, nd populist
volution and soity in a forwrd-looking way. b taught in th sh
.W
do not fous on muh-disussd rlzrtionshipssuh s standard irrtrpr- mands rationist
tations of soial Darwinism or rtra th lristory of ways in whih soial ial olrorts ply dif
ommntators hav usd (and misusd) volr.rtion.Thr is suh rih litra- groups as w go lon1
tur in ths gnral aras that any gnral Googl or Amazon sarh will of (human) soitis
yild dozns of offrings, of whih arl Dglr's I Serh of Hm Ntur: mans any group of
The Decli d Ruiul of Drtuiism i mrin Soil Thoght (1992| or struturd way and '
Rihard Hofstadtr's Soil Drinism i Amri hought (1959) ar just 19751Bod and Rih
0
uolutio d Soit 1
assumptions of unitl
volutionary Thry Shapd y Soity soity shard th as
hang ovr tim tha
volutionary thory xists in rtain soitis nd is dvlopd y mmrs 2002; Rihards 200,
of ths soitis.That muh is ovious. Darwin is th mst visil ntral th intlltual nvi
playr, and historins hav shown th signifian of th fts that soially partiular vrsion of
Darwin ws uppr middl lass and a Vitorian nglishman. For ampl, progrssiv impliat
th British gntlman would prsumaly hav found it asir than th poor an pliitly and
son of a long-poor working-lass family to om to Darwin's viw of inhri. impat, just s soi
.What
t a n a s a p o g s s i v n d p o s i t i v f o r i n p r s r v i n gv r i t i o n o t o s was sn as
hrdity as naling rathr than onstraining. Th way Darwin saw th ntury also rflts
world from .S Begle was surly shapd y his ontt and valus, as his- ory (Bowlr 1983)'s
torins hav shown (Dsmond and oor 1991; Browne 1'995,2003;odg randomly arising vz
and Radik 2003). His world Was th rihly ut gntly ntangledank of an and a fous on th i
nglish gardn rathr than n arid dsert, a divrs and pungnt rain forest, ganism in its nviror
or an Alpin tundra, h of whih might hav ld him to diffrnt qustions prtations tht app(
and mphass. rformrs who wr
Furthrmor, as Darwin sholars hav dmonstratd larly, th British ti dtrminist or h
.silliam
ontt of natural thology shapd volutionary thory. Paly saw ary thory, and th
ah organism and ah typ as adptd to its pla in th world. Of ours' If th hritabl vari
it was Paly's Anglian God th Crator who had don th adapting in that prsumably influt
.W
as, but th fit of lif form to nvironmnt arrid ovr natly to Darwin's annot ovr
viw of lif. Tru, a Thomas Rort Malthus might point to th human o. ing soity, though
nomi dilmmas intrinsi in th tndny of vry individual to inasinto Hr th point is th
a population that rapidly outgrows availal rsours.But althus did not volutionry thory,
all into qustion that humans wr adaptd to thir nvironmnt. Indd, to ask and whih as
th tndny to produ too many individuals just givs natur th opportu- rtainty whr som
nity to hoose som ovr othrs. Or, as Malthus suggstd, prhaps man as wll as intrnal t
ould do th hoosing with wis population ontrol. At any rat, soial viws study voiution: nt
shapd th volutionary thory that mrgd,and th rsultswr optimisti. trol and manipulatir
For althus, population ontrol would mak lif ttr for ah prson and whol orgnisms, pi
..halthy,vigorous, and happy''
for soity as a whol. For Darwin, only th parts lik bhaviors
.$isman
would surviv and rprodu. A happy pitur inded (Rus 1979,2003; August
Hodg and Radik 2003). 1'892 Kimplsml
Anothr part of Darwin's Vitorian miliu that was trmly important nvironmntal influ
for his vntual formulation of th thory of natural sltionas an plana- trial of volution,
tion of volutionary hang was his los ontat ovr many yars with many ptition among his
animl and plant brdrs(Brown 2003). Indd,th los omprison - long visibl hrom
twn artifiial sltion (with morphologial and havioral hangs in pop- matrial units, was
ulations that ould indud y ontinud rding)and natural sltion qustions aout hov
(hangsthat would a rsult of th ..strugglfor istn''in natur) is a th fin dtails of th<
..long
major part of Darwin's argumnt'' Was part of a sinti
Th Grmn intlltual movmnt Nturphilosophi, with its romanti mirosopi osrv
onption of natur, rought othr prsptivs that shapd volutionary ah organism, and
.!ismann's
thory as it playd out in Grmany. Johann Wolfgang von Goth's organi- volt
ism that saw an unfolding of form' Fridrih Shlling and Gorg Hgl's impat on th inh
uolutiln d Soit 3
disus of partiular parts ould hav any ffton volution is takn in rtro- te 1970s to qustin
spt as putting that ida to rst' though it did nt atually do so. Instad, ( l d r d g a n d G o u l d l .
no-Lamarkism ontinud to find a pla in soial and politial ontxts mulation of morpholo
that mphasizd th importan of hang through ffort and will, suh as ..
with mor suddn
tht of Rihard Smon arly in th twntith ntury nd with a lin of lss othrs, as passing thro
wll-known suPportrs through susquntgnrations (Smon 1908). By trnl fators might
th sond half of th twntith ntury' politi4l figurs suh as T. D. port ompting intrpr
Lysnko gaind onsidral authority in th Stalinist Sovit Union' and not thir viws wr intr1
solly for distortd politial rasons. Lysnko's no-Lamarkism appaldto th validity of volutio
th hop that if only sintists ould ultivat sds in th right nvironmnt, A ntral point hr
thy ould produ nough grain to sav th popl from starvation. Thr t h o r y t h a t r s u l t dv
was trmndous lif in th powrs of sin, and vn though no- appard. Old-shool
Lamarkian idas hav latr n idntifid with Sovit-styl matrialism, at f o h a n g .N o t o i n i
last initially many of thsproposals mphasizdth rol of ultu and th young and fanid th
soial nvironmnt in shaping th fat of human soitis. hang. This dos nor
Som would insist that this was larly ad sin, and for thos fully fa. than thos who favor
miliar with ontmporary sintifi paradigms, it was. But for thos for hav not had trmnd
whom volution was not a ntrl part of thir duation and worldviw, Thy hav.Our point l
suh an volutionary thory that would pla th ausSof variations dirtly tists asord hav sh
in us and disus in rspons to th nvironmnt mad sns.It was not a Today w s anotl
long strth from Darwinian volution to sing tmpratur hangs as a volutionary iolog i
sorrr of nw variations through mutation and sltion and thrfor as th tionary iology. Popul
sour and aus of th origin of spis.Lysnko's vrsion of no-Lamarkism within population. Bu
promisd a spdirvolution, and sin volution Was sn as progrssiv,it isms do, and th way
thrfor offrd a fastr rout to progrss and improvmnt. Soial nds variations has importa
shapd sin' whih thn playd out in soity (Tods 7989\. hangs. wo notions'
.s
s anothr ampl in th so-alld modrn volutionary synthsis of hav widr soital ir
th lat 1940s and aftr. Partiipants at ky mtings, suh as te |947 intrations (Maynard
Printon mting' saw this as a tim of optimism for volutionary iologists t h a t n o t l l v r i a t i o n s.
(Mayr and Povin 1980). Th prwar tnsions in urop and th Unitd vlopmntal and phys
Stats may wll hav nouragd th sort of synthsis that Julian Huxly, as vritions tran
a politial ativist with th Unitd Ntions, sought in is uolutio: The r t i so f t h d v l o p i n
odern Stbsis (1942). Huly was not a major intlltual shapr of the pl interations amor
privd synthsisut ntral voi in oniving of it in that way (Smo- togthr ontriut to
ovitis 7996). Howvr, vn whil volutionry iologists suh as rnst dvo (short for volut
Mayr and Gorg Gaylord Simpson agrly mrad th ida of synthsis s i z s a n p i g n r i a n d
with volution at th or, alrady othr iologists wr pushing in ountr- This prsptiv has
dirtions.ll iologistsand gntiistsrd mor aout th intrnal work. ida of Iimitson what
ings of lls and thir ontnts than aout th biggr piturs of volution. ws first mad popul.
Rapidly inrasdfunding for mdial rsarhand mdial dmands of wars om'' and th nvir
hlpd driv wdgs twn spialtiswithin th iologial ommunitis. on intrtions , v nb
volutionary iologists oftn found thmslvsas omprativ spialistsin f i r s t m a n i f s t t i o n si n
natural history musums' whil mor rdutionist rsarh programs gaind rlvant within dvlo
priority in many univrsitisand spiallyin mdial shools. ally, th ompltion
Dats aout th ffiay of adaptation rval similar soial shaping of nation of th gnti p
volutionary thory. StphnJay Gould, Nils ldrdg, and othrs gan in a n d p i g n t i n a t u r
uoltion d Societ 33.'
l.ton volution is takn in rtro- th 1970s to qustion how far natural sltion an rally fft hang
t did not atually do so. Instad' (ldrdgand Gould 1,972).Isvolution rally allgradualism and slow au.
in soial and politil ontts ..puntuatd''
mulation of morphologial diffrns? Or ould hang
: through ffort and will, suh as ..plosions''?Prhaps som hangs ould nal many
with mor suddn
lth ntury and with a lin of lss othrs, as passing through a ottlnk to th opning yond an do. Or
lt gnrations(Smon 1908). By trnal fators might lad to puntuation. volutionary thory ould sup-
, politil figurs suh as T. D. port ompting intrprtations gf mhanisms' ths rsarhrs flt, though
h StalinistSovit Union, and not thir viws wr intrprtd y antivolutionary rationists as hallnging
ko's no-Lamrkism appald t th validity of volution.
r s d si n t h r i g h t n v i o n m n t , A ntral point hr is that this ritiqu and th rvisions in volutionary
h popl from starvation. Thr thory that rsultd wr, in part, shapd y th soity in whih thy
sin, and vn though no- appard.old.shol adaptationism and gradualism providd a slow pross
l with Sovit-stylmatrialism, at for hang. Not oinidntally, thos who wr alling for puntuation wr
rasizdth rol of ultur and th young and fanid thmslvs soial rformrs who favord rapid soial
uman soitis. hang. This dos not man that thy wr wrong sintifially any mor
'ad sin,and for thos fully fa- than thos who favord gradulism. Nor dos it man that thir hallngs
ligms, it was. But for thos for hav not had trmndous positiv impt on volutionary thory gnrally.
f thir duation and worldviw, Thy hav. our point hr is that th soity and th soial valus th sin-
th aussof variations dirtly tists asord hav shapd th volutionary thory signifiantly.
onmnt mad sns.It was not a Today W s anothr transformation of th adaptationist paradigm in
singtmpraturhangs as a volutionary iology in th form of a mrgr of dvlopmntal and volu.
and sltin and thrfor as th tionary iology. Population gnti modls hav mphasized gnti hangs
;nko'svrsion of no-Lamarkism within population. But gns do not intrat with th nvironmnt, organ-
o l u t i o nW a ss na s p r o g r s s i v ,i t isms do' and th Way gnti variations rlat to orrsponding phnotypi
s and improvmnt. Soial nds variations has important impliations for our undrstanding of volutionary
rcity (Tods 1989). hangs.Two notions, in partiular, that ar part of this nw paradigm also
modrn volutionary synthsis of hav widr soital impliations, namly, th idas of onstraints and of
ky mtings, suh as t'e 1947 intrations(aynard Smith t al. 1985). Th ida of onstraint mphasizs
timism for volutionary iologists that not all variations ar qually likly or vn possil. It stats that th d-
nsions in urop and th Unitd vlopmntal and physial oundaris of an rganism limit th ways gnti
lf synthsisthat Julian Huxly, as variations an trnslatd into phnotypi varitions aus of th prop-
ns, sought in ts u<iltio: Tb rtisof th dvloping systm. Ths ar, of ours, a onsqun of multi-
a major intlltualshapr of th pl intrations among gnti' ellular' and nvironmntal fators that
:onivingof it in that way (Smo. togthrontriut to organismi dvlopmnt. Th nw synthsis of vo-
rtionary biologists suh s rnst dvo (short for volutionar and dvlopmntal ilogy) thus also mpha-
ly mradth ida of synthsis sizsan pignti and intrativ viw of iology.
i o l o g i s t sw r p u s h i n g i n o u n t r - This prsptiv has many sintifi, ultural, and soital rsonans; th
rd mor aout th intrnal work. ida of limits on what is possil within th oundaris of a ntural systm
th iggr piturs of volution. ..population
was first mad popular in th ontxt of th worry about a
rrh and mdial dmands of wars om'' and th nvironmntal movmnt. Similarly, th growing mphasis
' i t h i n t h i o I o g i l o m m u n i t i s .
n intrations' vn twn distant parts of a larg osystm' has found its
lvsas omparativ spialists in fistmanifstations in this ontxt. But today ths onpts ar also sn as
tionistrsarhpograms gaind rlvantwithin dvlopmntal and volutionary biology, whr, quit ironi-
in mdialshools. ally' th ompltion of th many gnom proits-in mny ways th lmi-
n rval similar soial shaping of nation of th gnti paradigm-has ld to insights into th highly intrativ
ils ldrdg,and othrs gan in and pignti natur of oth dvlopmnt and volution (Hall 1,998)'
6 uolutio nd Soit
sltion as an pl
That volutionary thory has not n insulatd from soial influns is
207). Anothr influ
no surpris to historians of sin' ut it is oftn surprising to sintists who
natural history and
s sin as rlativly insulatd and progrssing aording to its own intr-
is of volutionary h
nal logi and rsponsS to opportunitis. It is also surprising t som soial
whil, in Vinna m
historians' who rarly study th history of sin and who tnd to tak si-
dvlopmntal mha
n as givn and to s it as shaping soity on oasion rathr than to s
th work of th iol
th intrations in both dirtions. Darwinism. as Dsmond and oor
primental basis fo
(1991) and Brown (1,995 , 2003 ) hav shown learly, is iust as muh a story
ontt in favor of a .
of Darwin in soity as of Darwinism in soity.
on volution (Przirar
All ths ampls
ity nvr Was jrrsta
Soity'sRption of volution
rlationship that in tu
ary thory in intrsti
Historians hav long dsrid th rption of Darwinism, in partiular, to
.shy
show th variations in ration. is volution takn as a hallng to s-
talishd valus in som ountris or loal soitis and not in othrs or to
volutionary Th
som rligions and not othrs? \/hy hav som groups radily ndorsd th
naturalism at th or of volutionary thory, as Germany did in th lat
Thr ar no a priori l
nintnth ntury, whil thr is hostility at som tims and in som groups
planation as long as
to th apparnt randomnss and lak of purpos that thy s in volution?
multitud of diffrnt
To what tnt is th rspons shapd y loal ontingnis of dominant in-
and ologial intra
fluntial individuals, prhaps' and to what tnt dos th ration flow from
(Rus 1996).Ovious
th logi of th values and assumptions within th ultu? Th answrs to
mnts of slim molds l
ths qustions ar ompl and involv dtaild aounts of th soial' ul-
orhstra who ar pl
tural, onomi, and politial history of ths soitisand ountris. For
th rsult of th intrp
mor dtaild aount, th radr should onsult th sustantial ody of lit-
.$7assr- th framwork of vo
ratur in this ara (fo xampl, u|| 1971Kohn and Kottlr 1.985;
mnt, kp in mind t}
sug and Ros 1984).
ioral, an onsidr
Hr w want to point to just on aspt' namly, how diffrnt soitis'
Natural sltion f;
rption of volutionary idas ontriutd to th frthr dvlopmnt f vo-
manag to inras t
lutionary thory. \Within Darwin's innr irl Thomas Llnry Huxly most
is dsrid y th t
visily mphasizd th importan of volutionary thory to promot a liral
sris th onsqun
and matrialisti agnda. To him, volution implid that thr is no intrinsi
that an rprodu 1
valu in heritag and pdigr, and tht all that should ount ar th ailitis
1,998).Th rpliator
of popl. As a onsqun, h promotd sin as a profssion rathr than
a voation and fought a onstant attl gainst rligion as th stalwart of r- its atually ontriu
variabl faturs ar r
ivd valus (Dsmond 1997)' Similarly, rnst Hakl fousd on th mat-
a (loal) optimum as
rialisti impliations of volutionary thory, whih h pandd into a whol
thrmor' th rpliat
systm of monisti philosophy. Nithr Huly nor Hakl thought highly of
ag to rprodu th
natural sltion as a mhanism of volutionary hange. Rathr, ah inor-
rprodution happns
poratd th prinipl of volution into his sintifi disiplin (mostly mor.
proprtis do not disir
phology) and his highly sussful tahing. In tahing thy oth intratd
potntial and atual l
with and shapd urrnt trnds within thir rsptiv soitis' whil thir
uls, gns' lls, l
sintifi ontriutions inflund svral gnrations of iologists.
groups' and idas, s
A similar pattrn an sn in Russia, whr ids and osrvations
1984; Sor and.Wils
aout symiosis and oopration also hallngd th dominan of natural
uolutio d Socit 7
ad limits of what kind of pfopr- oordinatd.How thsrgulationsom about and volv is on of th most
th fitnssf thir arrirs or thos fasinatingprolms of volutionary iology (Margulis 1981).
os, howvr, ar th sujt of a Ths rif thnial ommnts aout the asi assumptions of volution-
ir'now ask in what way a spifi ary thory also hlp us s th long-standing ontrovrsy ovr natur vrsus
:aturor a form of havior an nurtur in a diffrnt light. In th ontt of volutionary iology this dis-
:ssan additional qustion. Bfor tintion is on only of dgr and not of kind. Thr is no lar.ut spara-
nss'thy must first mrg within tion twn natur (or iology) and nurtur (or ultur and soity).
i produts of volution. In othr A multitud of fators, oth iologil nd ultural, ontriut to th dvl-
lmntal ontxt of organisms is th opmnt and formation of any human ing; indd, our dvlopmntalpro-
tur.only thos phnotypi vari- gram rquirs all ths diffrnt stimuli as ruial inpts.
: raw matrial for natural sltion' As w hav larnd from volutionary iology, ompard with our losst
of thir arrirs. Th fat that d- rlativs,w ar orn prmaturly (Portmann 1951). On onsqun of
nration of variation has rntly this fat is that in human infants th dvloping rain is stimulatd y a mul-
fif 1996 Hall 1998). titud of nvironmntal nd soial fators. In addition, w hav larnd that
lanationsof soitis' a diffrnt in our volutionary history th mrgn of soial skills, inluding th volu-
'f intrst.As w hav sn, all that tion of languag, was muh mor important than aumlatd gnti diffr.
'h availailityof auratly trans- ns(Diamond 19921Kupr 1994). Of ours, prisly thos haratristis
Lipis ommonly masurd by hri- also nal larning and th transmission of aquird knowldg. In onlu-
y dfindas th orrlation twn sion, all vidn thus far points to highly intrativ rlationships among
in many ass th rason for this diffrnt fators (iologial and ultural) in human volution. onsquntly,
.d to th ffts of gns (Falonr i n p l a n t i o n so f h u m n v o l u t i o n n d s o i t y W n n o t g i v p i o r i t y t o
ially importnt in th ontxt of any on of ths fators ovr all th othrs. This ralization shold also hlp
only an idas or mms intr- us avoid th naturalisti fallay; knowing th volutionary history of som
rf human' as wll as animal, soi- aspts of ur havior and soial organization dos not imply any valu
l to th nt gnration. From this judgmntaout ths proprtis. But ralizing th volutionary rasons for
ntradition twn th intriat thshaviors dos hlp us if w want to nfor soital ols, a fat that is
hangsand transformations of i n r s i n g l yr l i z d y a g o w i n g o m m u n i t y o f l g a l s h o l a r s . o n o m i s t s .
lf volutionry iology. Thr ar, and managmntonsultants.
]s gns that ontribut to human our rif analysis of th rlationship twn volutionary thory and so-
from th viwpoint of volution- ity notwithstanding, it is also tru that throughout th history of volution-
ns add many diffrnt lvls and ary iology th majoity of ttmpts to apply volutionary prinipls to
ithin th onfins of volutionarv xplanations of human soitis hav n grandios failurs (Hofstadtr
1959). Sn through historians' ys' ths ass ar prim xampls of th
]tiv dos not imply any form of mutual rlationship twn sin and soity disussd in th first stion;
utionary analysis of omple sys- at rtain tims speifi soital lifs and assumptions rsonat with l-
t w simultanouslyonsidr ffts mnts of the thn-urrnt stat of volutionary thory and om th foun-
J, in volutionary iology th most ..planations,'
dation of wid-ranging of natur and soity. Sn through
ions whr any simpl xplanation th ys of an voltionary biologist, ths ass tak on a diffrnt maning.
it twndiffrnt ntitis. Ths on th on hand, on might rgu that ths ass ar a rfltion of th
sribdy th lat John N4aynard thn-inomplt status of volutionary thory, at last whn ompard with
)ut somthing fundamntally nw, our prsnt undrstanding.Takn at fa valu, this viw is prolmati -
or th first human soitis (}ylay- aus it implis a rathr niv onption of sintifi progrss. Anothr o-
to surviv, ths nw ntitis nd srvationan volutionry iologist would mak, looking ak at prvious
:rst of th parts; in highr organ- assof volutionary planations of soity, is that all ths proposals tnd
,dutivintrsts' and in a omplx to mphasiz only on or a fw lmnts of wht rnst ayr nd othrs hav
'or, ths ativitis also nd to idntifid as th multipl elmnts of volutionary thory (Mayr 1982).
40 uolution nd Soiet
This obsrvation allows uS to omin historial ontxtualiztion of vo. tion' ruially dpn
lutionary planations of soity (what was th sintifi, soial, politial, hrditary matrial.
and onomi ontt of ths proposals?) with valuation in trms of thir Galton saw possiil
sintifi mrit (what wr th prolms with and wrong assumptions of volution and th fu
ths proposals? what was lft out' and what was aggratd?).This ap- th Vitorian progr
proah allows us to analyz ths various proposals oth in rlation to thir aus and ontrol it
historial ontxt nd with rgard to thir sintifi ondnt. Unfotunatly' rding with th n
W do not hav th spa to disuss ths idas in dtail, ut w want rifly In nd Super
to prsnt a fw eampls of how volutionary idas wr applid to human progrssiv intrpr
soitis in th past for onluding with a skth of how prsnt-day vo- that som ar natur
lutionary biology approahsth issu of human soitis. soity is urrntly
It will om as no surpris that as soon as thr was volutionary thory, it no informd rdin
was applid to plain aspts of human soity. rrt Spnr' a popular ftions now and als
philosophr and advoat of idas similar to thos of Darwin, oind th prmn.
trm sLlruiul of tbe fittest to dsri th onsquns of natural sltion. Indd, ugnis l
Dtahd from its stritly iologial maning' this trm ould (and was) Amrian visionari
applid to a varity of soial ontts. It ould takn to support a onsr- joind th all for go
vativ status quo whn it was omind with idas and osrvations that gradually did sint
volution favors onsrvation of xisting traits and hviors. In this way drgirdd th ugn
volutionary thory providd a naturalisti and matrialisti xplanation of ugnis was not as
soial diffrns y dmonstrating how some popl ar mor sussful . tors in rtrospt.S
aus thy ar fittr in Hrrt Spnr's sns. Thir variations ar ttr ugni programs a
adaptd to th omptitiv soial nvironmnt in whih thy liv, and thy ugni idas' tog
thrfor naturally nough ris to th top. raial hygin' ontr
Not surprisingly, it was th grat industrialist Andrw arngi who most nat ..unworthy lif,'
nthusiastially mrad this intrprtation of volutionary thory as - oalition formd to <
plaining soity. But th ida of natural sltionwas lso usd in support of logial dfinition of
radial transformation of soity, as th initial positiv rption of Darwin mnt on ra. This
y l\4ar, ngls, and othr lading soial dmorats attsts.Thy, of ous' antidisrimination pr
mphasizd th transformativ powrs of natural sltion rathr than th rathr than iologi
onsrvativ ons. onsistnt with stud
\/hatall ths initial appliations of volutionary thory to soity hav in nS among humans
ommon is that thy fous mostly on omptition' struggl' and a progrssiv nti diffrns in m
intrprtation of (volutionary) history, as wll as a hirarhial notion f populations hav ls
diffrnt soial lasss and ras. Furthrmor' thir notion of omptition is nalogial rlationsh
mor or lss dirt or hands-on. Struggl was oftn sen as a onsqun of our spis in ast A
diffrnt strngths of individuals, groups, stats' or ras. Indd, th arly sujt of gnti dif
yars of th twntith ntury rought svl attmpts to adapt Darwinian find a growing ons
logi to work as a guid to intrnational affairs and politis (Zieg|er 1918). of mankind an i
\/hat all ths appliations of volutionary thory to soity missd was n dvlopmnts in th 1
undrstanding of th intrnal onditions of organisms and th mods of ial tratmnt on th
transmission of hrditary matrial' whih wr' aftr all, ruial lmntsof Th two most imp
Darwin's thor. and human) havio
Th ntrality of hrdity to th thory of volution Was not missd y tion of smingly altr
Darwin's osin Franis Galton. Inhis l-Irditr Gnjus (l869) h ralizd tions, and hr sp
that th umulativ fft of natural sltion. *...,i.u for sradual volu- and othrs (Hamilton
uolution nd Soiet 41
istorial ontxtualization of vo- tion, ruially dpnddon oth th vailaility and th staility of adquat
as th sintifi,soial, politial, hrditarymatrial. Fousing on inhritnand stability aross genrations'
with valuation in terms of thir Galton saw possiilitisfor ontrolling populations and thry ontrolling
with and wrong assumptions of volutionand th futur of human populations. This was an ntiing ida to
,,hatwas xaggratd?).This ap_ th Vitorian progrssivist. Not only ould w hav progrss, ut w ould
lroposalsoth in rlation to thir aus and ontrol it through iology, Galton suggstdwith his all for good
sintifiontnt. Unfortunatly, brding with th ngativ.and positiv sltions of ugnis.
las in dtail, ut w want rifly In nd Suprm (190), Gorg Brnard Shaw also rfltda mor
rary idas wr applid to human progrssivintrprtation, though still vry muh groundd in assumptions
a skth of how prsnt-day vo- tht som ar naturally bttr than othrs. !(/hy not aknowldg that our
uman soitis. soity is urrntly wakr than it ould and should aus w hav
.!7
s thrwas volutionary thory, it no informd rding program' Shaw suggstd. an plain our imp-
lity.Hrrt Spnr,a popular ftionsnow and also sk to improv, gnrating a soity of mn and su-
.to ths of Darwin, oind th prmn.
:onsquns of natural sltion. Indd, ugnis rought hop for progrss spially to uropan and
ing, this trm ould (and was) Amrian visionaris in th arly twntith ntury. Lading gentiists
luld takn to support a onsr- joind th all for good rding and for ttr soity through ilogy. only
with idas and osrvations that gradually did sintists apt how littl solid sintifi knowldg rally un-
traits and haviors. In this way drgirddth ugni assumptions' so that in th 1910s and into th 1920s
: and matrialisti xplanation of ugnisWas not as ridiulously unsintifi as it has smd to ommnta-
m poplar more sussful- tors in rtrospt. Soial sltiv prssurs rinford thos who adoptd
sns.Thir variations ar ttr ugniprograms asd on biologial laims (Kvls 1985; Paul 1995).
nnt in whih thy liv, and they ugni idas, togthr with a iologial onpt of raial supriority and
raial hygin,ontriutd to th Nazi gnoids' th ampaign to trmi-
'ialist Andw arngi who most nat ..unworthy lif,'' and finally th Holoaust. As a onsqun' a growing
on of volutionary thory as x- oalition formd to ountrat what was privd as an inappropriat io-
tionwas also usd in support of logial dfinition of human ras, ulminating in th 1950 UNSO stat-
itial positiv rption of Darwin mnt on ra. This statment thn am th foundation for furthr
lmortsattsts.Thy, of ours' antidisriminationpoliis and laws. Ths dlarations mphasizd thni
natural sltion rathr than th rathrthan iologial diffrnstwnhuman populations, whih smd
onsistnt with studis that showd that th vast maiority of gnti diffr-
iutionary thory to soity hav in nsamong humans our within populations. Howvr' in rntyars g-
t i t i o n s, t r u g g | . n d a p r o g r s s i v nti diffrns in mitohondrial and Y-hromosomal DNA twn human
; wll as hirarhial notin of populations hav also n usd to illumint migrarion pattrns and g-
o r ,t h i n o t i o n o f o m p t i t i o n i s nalogial rlationships twn hman populations' inluding th origin of
,as oftn sn as a onsqun of our spisin ast Afria aout 130'000 yars ago. Today, vn though th
stats' or ras. Indd, th arly sujtof gnti diffrnstwn human populations is still touhy, w
ral attmptsto dapt Darwinian find a growing onsnsus that nithr th iologial nor th ultural history
'ffairsand politis (Zieg|er 1918). of mankind an ignord. This is spially tru as svral thnologial
y thory to soity missd Was an dvlopmntsin th postgnomiag inrasour ailitis to ustomiz md-
of organisms and th mods of ial tratmnt on th asis of th gnti onstitution of patints.
wr. aftr all. ruil lmnts of Th two most important thortial insights into th volution of (animal
and human) havior wr Hamilton's notion of kin sltionas an plna-
, of volution Was not missd y tion of smingly altruisti havior and th ppliation of gam-thorti no-
,ditr Geius (1869) h ralizd tions, and hr spillr th onpt of stratgis, y John aynard Smith
tion, nssaryfor gradual volu- and othrs (Hamilton 1|64;anard Smith |9s2). Ths idas rprsntda
42 uolutil d Soit
major hang in volutionary plnations of havior and st th stag for mpirial data and th
th dvlopmnt of rigorous mathmatial modls. mal and human ehav
Th thory of kin sltion nd of inlusiv fitnss fousd th dat on Th soioiolgy dt
th appropriat units of sltion in planations of havior. Rathr than lutionary xplnation
just looking at on individual's fat as th sol dtrminant of fitnss, th th- of th many dimnsio
ory of kin sltion hangs th wys in whih vo/utionary aounting is ism, supportd y his
.sithin ulation gntis at thil
don. th framwork of population gntis hangs, volution is rp-
rsntd as hangs in gn frqunis. Wht Hamilton ralizd Was that intrativ prsptiv
thr is mor than on way in whih a spifi gn in a spifi individual rs' and his ritis w
an mak it into th nt gnration. It an passd on to its offspring sintifi, nvironmn
dirtly or through th produtiv suss of Ios rlativs, who will shar molular iology). T
th sam gn with a rtain proaility (.5 for silings, .125 for ousins).In ns of that tim, wI
alulating th fitnss of a spifi gn' on thus has to fous on th inlusiv dats.
fitnss, whih is th sum of all th diffrnt mans y whih a spifi gn Today, mor than
an om to rprsntd in th nt gnration. tionary planations (
Prvious vagu notions, suh as th ida that a havior an volv - popular within iolo;
aus it is good for th spisor th group' wr rplad y onpts that asis for this rsarh
aordd with th prditions of population gntisand th thory of natu- ral dbat and or
ral sltion. Howevr, it is also important to not that ths notions Wr ogy. Yt som of todi
highly astrat, and whn thy wr first proposd, sintistsdid not rally soitis bttr rfl
know anything aout how ths proposd gneti rpliators atually ould any diffrnt filds
aus th rsptiv haviors. At this tim th argumnts wr lrgly tho- study of th volution
rtial: if a gn/rpliatorauss altruisti havior, it an favord by as for intrdisiplin
natural sltion as long as th ost to th individual ating slflsslyis kind of rsarh;as
smallr than th nfit to its ripints multiplid y th offiint of rlat- havior' it is in dan
dnss twn thos individuals. A similar approah guidd th many bat, vn if sintifi
gam-thortionpts, suh s th prisonr's dilmma and idas aout r. aout volution in g
patd gams and riproal altruism that wr usd to plain many asPts
of animalhavior.
Th appal of ths nw idas Was normous, and iology smdapal onlusions
of xplaining muh of soial havior. Th idas providd an important
oost to many disiplins, suh as havioral ology, ut also invitd far- volutionary thoy i
rahing gnralizations and in turn gnratd many' ftn polmial' ontro. th sins, iology
vrsis.Th most visil of th sandalsruptd with th puliation of . o. ogy and iomdiin
.lfilson's hildrn's, lot will im
Sobbiolog tn 7975.In this nylopdi tom./ilson' an prt
on ants and soial insts' olltd an normous amount of mpirial data aout th ngativ
on what Was thn known aout animal havior and organizd it in th futur of human soit
ontXt f th nW thortial framwork. In th final haptr h applid to at stak. Biolog;
ths idas to human havior and th volution of human soitis.Fur- srvativ iongings for
thrmor, h proposd that this nw synthsis would lad th soial sins us on th history of
to intgratd within th framwork of th iologial sins.Th ra- stag.
tion was vivid. Howvr' nsuing dat fousd mostly on th feard politi- It sems almost inl
al impliations-Wilson's most voal ritis long t th Lft-nd on th assoiatd transformz
onsquns of th rdutionism impliit in l/ilson's mphasis of gnti . triggr ration y ful
planations and in his modl of th rlations amog t\sins. Surprisingly ontxt of volution
littl attntion ws paid to th ulk of th ook. th normous amount of mnts highlights imp
uolution d Soiet 4
of haviornd st th stag for enrpirildat and th rigorous appliation of volutionary prinipls to ani-
odls. lnlnd humn hvio.
siv fitnss fousd th dat on Th soioiology dat rsnrld th many othr attmpts to pply vo-
nations of havior. Rathr than lutionary planations to human soitis in that it ovrmphasizd just on
'Wilson's
ol dtrminantof fitnss,th th- of th many dimnsions of volutionary thory. gnti rdution-
w h i v o I u t i o n a r y o u n t i n g i s isr-l-l,
supportd y his prin with soil ants' as wll as th stat of pop-
g n t i s h n g s ' v r l l u t i o ni s r p - ultion gntisat this tim, lft Iittl oom for othr fators or for th mor
Vhat Hamilton ralizd was tht intrativ prsptiv that has mrgd mor rntly. \Vilson' his support-
ifi gn in a spifi individual rs' and his ritis Wr all produts of thir soial and politial, as wll as
:an passd on to its offspring sintifi, nvironmnts (at that tim dominatd y th rnt susss of
s of los rlativs, r,r.howill shar molular iology). This ld to a rathr polarizd limat within th lif si-
i f o r s i l i n g s ., 1 2 5 f o r o u s i n s ) .I n ns of that tim, whih in prt plains th onfrontational ton of ths
l ths has to fous on th iniusiv dats.
tt mans y whih :r spifi gn Today, mor than 30 yars ftr th puliation of Sociobiology, volu-
sration. tionary xplanations of soity and disussions of ltural volution ar still
la that a havior irn volv - popular within iology. Howvr, the onptual, as wll as th mpirial,
p' Wrerplaed y onpts that sisfor this rsarh has om muh mor pluralisti. Thr is still onsid-
n gntisand th thory of natu- raldt and ontrovrsy,spially surrounding volutionary psyhol-
t t() not tht ths notions Wr ogy. Yt som of today's approahs to undrstanding human (nd animal)
rroposd,sintistsdid not rally sitis ttr rflt th insights of volutionary thory skthd arlir.
gntirpliators atually ould any diffrnt filds of volutionary iology ontriut' and indd th
l th argumntsWr largly tho- study of th volution of ompl soial systmshas om a paradigmati
i havior,it n favord y asfor intrdisiplinary rsarh.As suh, it fas all th hallngsof this
th individual ating slflssly is kind of rsarh,; as ah spilty tks on on spt of soity and human
rltiplid th offiintof rlat- bhavior, it is in dangr of ovrmphasizing that part. And th nsuing d-
rilar approah guidd th many at, vn if sintifially produtiv, is rivd y a soity that is unasy
n'sdilmma and ids out r- aout volution in gnral.
rusd to plain many aspts
th fundamntal prinipls of volutionary thory and in ploring implia- without ing undul
tions and limitations of ths sintifi insights for our urrnt soitis and ial organization and
for human slf-undrstanding (sth main ssays ..volution and Rligion'' prinipls of th rpli
y David N. Livingston and ..Amrian Antivolutinism: Rtrospt and produt of volution
Prospt'' y ugni C. Sott in this volum). l/ thus of{er a fw rmarks nomna (suh as th
on futur dirtions. studied within just or
volutionary analysis of human havior and soial organizations will r- solvd whn thy art
main an important rsarh ara within volutionary ilogy and will hav a urrnt xtnsion of
high potntial for insights, as wll as for onflit. volutionary mediin is al- ontrvrsil ontinu
rady transforming th ways w undrstand and trat rtain disass (s of ours, all this
th main ssay ..volutionary Biology of Disas and Darwinian diin'' tionary thory might
y ihal F. Antolin in this volum). volutionary psyhology offrs in- vid a justifiation. I
sights into th history of human havior and our potntial to modify and for thial thory, fl
ontrol havior, inluding possil transformations in our lgal systms and ..oug
annot lad to
pratis (sth main ssay ..SoialBhavior and Soioiology'' y Danil I. hav,xpt insofr
Runstin in this volum). Th mrging of onomi thory with volution- ory as good sini]
ary thory nfits oth filds and has givn volutionary thory W OnP-
tual and mathmatial tools, suh as gam thory and th notion of stratgis.
It has also introdud nw qustions y rplaing traditional idas of an ..in- B|LIoGRAPl-
visil hand'' with th havior of individual ators shapd y thir volu-
tionary history and onstraints. Appliations of volutionary thory vn Akrt, L. 2007. Th ,.
,l'885-1940.
transform rtain filds of nginring and omputr sin as th prinipls Jlur
A d a m s , . B . 1 9 9 4 .T |
of volutionary dsign and of gnti algorithms hav opnd nw vnus for -hought
d i Rt
solving omplx optimization prolms, suh as th dsign of airplan wings Bowlr,P, l, 1983.h
(Rhnbrg197i Ho|Iand 1995 ). i the Decdsr
Th rlationship twn volution and soity also provids an important Boyd, R., and P. J. Ril
as study for intrations twn sin and soity mor gnrally. Th Univrsityof hi
B r n d o n 'R . N . , n dR .
history of volutionary thory is among th st-studid aras in th history
otrouersislu
of iology and has rvald intrsting ways in whih sintifi rsults mrg Brown,J. 1995.hr
in a ompl intrplay twn sintifi invstigations and what an 203' hrls-
alld thir soial and ultural ontts. Although ths studis hav ld to Univrsityrss.
insights aout how sin happns and thus hav ontriutd to popular un. Dwkins, R. 1976,h
"1982.The t
drstandings of sin, thy also hallng philosophrs of sin to rthink .W.
. Frman.
som fundamntal assumptions aout th natur of sintifi vidn and Dglr,. 1992.I S
how historially ontingnt xpimntal and thortial pratis an lad Driism i Amt
to inrasingly ..aurat'' rprsntations of natural phnomna. Som vn Dsmond'^. J' 1997.I
go so far as to dsrib this ..soial'' pross of finding th ..truth'' in sin Rading'A: Ad
in volutionary trms' thus oming full irl. Dsmond,A. J., andJ.
Diamond, I. . 1992-
But th most important aspt of th ongoing intrations twn volu-
HumnAimI,N
tion and soity lis in th ways in whih volutionary thory transforms our ldrdg,., and S.J.
slf-undrstanding as oth iologial and soial/ultural ings. As propo- phyltigradualis
nnts of ultural volution suggst' thr is no inhrnt differn twn San F'raniso:Fr
th ways our odis and haviorsand our soitisand ulturs volvd.In- F a l o n rD' . S . 1 9 8 9 .1
G l t o n 'F . 1 8 6 9 . H r
dd, th rigorous appliation of volutionary thory to suh aras s th
London: Mamilla
volution of languag and of ultural transmissin.hasallowd ths propo-
Hall, B. K. 1998.uolt
nnts to formulat a thory of human soity and ultur that is matrialisti all.
uolutio nd Soiet 4''
'thr and in ploring implia.
without ing r"rrrdul gn ntrd and rdutionisti.Th insight that so-
ightsfor our urrnt soitis and il organization and ultual tanstnissiona govnd y the sam gnral
..volution
1 ssays and Rligion'' priniplsof th rpliator quirtion implis (1) that not vrythingthat is th
Antivolutionism:Rtrospt and produt of volution has to grrtiallydtrmind and (2) that many ph-
m). W thus offr a fw rmarks nomna (suh as th volution of oopration) that rmain puzzling whn
studidwithin iust on l/l of analysis (gntior soial/ultural)an r-
r and soial organizations will r- solvd whn thy ar approahd from oth dirtions. \/ithoutdout this
lutionarybiology and will hav a urrntxtnsion of volutionary theory to soity will an important and
nflit.volutiorrarymdiin is al- ontrovrsilontinuation of th long history of volution nd soity.
nd nd trat rtain disass (se .!
of ours, all this an takn too fa. must kp in mind that volu-
)isasand Darwinian Mdiin'' tionary thor might provid an planation for soity' ut it nnot pr-
volutionary psyhology offrs in. vid a justifiation. volution also annot provid an pistmi justifiation
and our potntial to modify and for thial thory, for ampl, dspit th numrous attmptsto do so. ..Is''
)rmations in our lgal systms and annot lad to ..otrght.''vlution annot tll us how soitis ought to -
ior nd Soioiology'' y Danil I. hav,pt insofar s it follows that soitismst apt voltrtionaryth-
[ ononrithory with volution- ory as good sinif thy gr to apt sinirt all.
r volutionary thory nw onp-
hory and th notion of stratgis.
llaing traditional idas of an ..in- BIBLIOGRAPHY
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Raft R. ^. 1996. The Shpe of Lif: Gs, Dulopmet, nd th uolutio of
il Form, lago: Univrsity of hiago Prss.
Rhnrg, I. 1973. uolutionsstrtegie: Optimirng tchisber Sstm nch
Prizipien dr biologish uoltil. Stuttgart-Bd annstatt: Frommann-
Holzoog.
Rihards, R. J. 2002. Th Romnti onption of Lif: Sien d Phiklsopb i
th Age of Goth. hiago: Univrsity of hiago Prss.
Rus, . 1979' Th Duii Reuolutil: Scic Rd i ooth d lu.l.
Chiago: Univrsity of hiago Prss.
uolution d Srliet 47
of Life: Si d Philosoph i
hiago Prss.
iRd i Tooth d l.
Givn the ompli
mntion Tlromas H
..dp sns
of rligi
(quotd in Brown 2
natur of Darwin's o
volution and Rligion at what point? Did
to do with it? Did
ntivolutiorristsali k
Duid N. Liuingston h ida that Drv
ontstd y thos l
to th nd'' (Moor
spiritual momnts lr;
ooks dating frorl tl
..Sopy Sm'' \trilrfor,th ishop William Paly's o
vr sin th mythi nountr twn
..Darwin,s sign; and th prson
of oford, and Thomas Hnry Huly, ulldog,'' at th 1860 mt.
trin'' of trnal rtri
ing of th Bitish Assoiation for th Advanmnt of Sin' thr has na
dath of his dlightf
widspad snsof an ongoirrg and invital lash twn volution and r-
thos who insist tlrtrt
ligion. Th lristorial narrtiv of tlris nOulltr' howvr, provs to th
mind, and motion (
lss lar-ut that suh irnprssionism miglrt suggSt.In th pags that follorv
athism or mtapy
w xamin th rligious viws of Darwin himslf and somthing of th histo-
ims\i tutnd arar
riography of th date for introduing atholi, Protstnt' Jwish, and
rusading sulrist
Islami rsponssto his thory. A numr of rurring thms 1rgfrom this
Sin irrslrtti
historial survy' and a rviw of thsissusorrltldsthis ss:ly.
iudgmnt of ths
ing to diagnos D:
hsitany ar mor
Darwin's Rligious volution wavring to whih -
fr xampl, h r,vr
Syms Covirrgton Was }lS Bgl's fiddl, ship's oy, and odd'jo man. For this is alwavs pair"
just ovr si yars h was also harls Darwin's srvnt.Bush was th athistially. But I r
evr-prslltassory to Darwin's gogrphi arrd sintifi travls, it is wish to do, vidn
hardly surprising that this virtually invisil Darwinian adjunt would othr hnd I anno
soonr or latr ath a novlist's imagintion. Th final fw sntnsof spially th natur
Rogr Donald's fitionirl aount of <lvirrgtontn r. Drtuin's Shootr brut for'' (Burkh
(1998' 410) rystalliz an irrrliguityat th hart of any aoDnt t>fth l- of vaillation sttrf
..H saw Darwirr on his kns,ad ..trukl''
tionship twnvolution and rligion: to puli
thr was no diffrntwn prayr and pulling a Wor from th grass. th origi oi S1li
As for r ovington, h prayd in th old-fashiond way.'' Th lision - ] 00 tims. Aor'rn
twn th transndntalrrd th asthti tlrat is apturd hr rsonats Josph Dalton Ho]
with Mihal Rus's onlusion to his rnt ilount in Drtuitl d Dsig. 1879 note tl J' For<
Fully irwirof Darwin's rpatd rsort tO th languag of uty and worrd tuats'' (quotl in
..of (|986, 27) onlus
in th ntural ordr, Rus (2003, 335) rn-lindshis raders th gnuin
lov and joy''-an ..ovrwhlming xprin'' touhing on th spiritual- to and flow..
that volutionists snsin thir nountrsr'vithth organi rvorld. Hr w mti athist; at hig
..Hosannah'' in intoirtddlisht asially agnosti-
ar alrtd tl th Drwin who brathd
at th sulinrity of th primval Bzilin rin frlrst. mit hirnslf o suh
48
uolutkln d Religio .t49
.vr'annot rstritd to fltu- prsidntof ornll Univrsity, A Histor of th Wrfr of Siee luith
flunof Paley's Ntrl Tholog holog i hristedom (\896). Th vry titls of works lik ths did
:onntionsPaly livd h ould muh to onfirm th imprssion of a monumntal dath struggl twn th
man agnyprovidd Drwin with forsof nlightnd sin and nightd rligion.
. Natur, Darwin judgd, was in- But th risp larity f this rivd imag sustituts monohrom ab-
athr lik th mannr in whih pi- stration for historial thniolor. Thr is a rihr narrativ to unov-
,,days',
mising variations. As h himslf rd.For a start' at lst sin ugustine, many ad read th ration
[y and hourlv srutinizing' thrugh. of th Gnsis rord symolially rathr than litrally. Various harmonizing
lightst;rjtingtht whih is ad, stratgisto maintain onord twn Gnsis and gology had long n
,od" (Darwin, 1959, 168-169). In laoratd,and th ida of a lngthy arth history was wll stalishd in th
lttriutsof Paly God. To this w minds of arly Vitorian hristian gntlmn.gologists. Natural thology
adaptationand th harmony of na- took a varity f forms' som lss susptil to Darwin's antitlologial
.as thy wr to Darwin's. Yt Dar- hallng' and of ours thr wr traditions, lik rtain strands of high
nnot limitd to arhittural Anglianism and strit unsntimntal alvinism, for whih th signifian of
nd onvitionsmrgd in dialogu th dsign argumnt was ngligil. And thr wr thos lik John Hrshl
hristian dotrin. His ritiqu of and William !hwll whos rligious outlook prdisposd thm to strss
, was umilially onntd to his ration y natural law rathr than y divin intrvntion' a mov that
nhuman and animl-how ould opndth door to th ida of volution as God's mthod of ration. Th
rind trustd? His allrgy to th ways in whih volution and religion ould mad to fit on anothr hav
>fPaly's uides of hristiit takn,and ontinu to tak, myriad forms.
lonaldFlming (1961, 231) spu- lfhat furthr ompliats mattrs is that whn onflit twn volution-
..yarning aftr a ttr God than ists arld rligious livrs did our, th ampaign was oftn wgd from
r nurturd y an inaility to r- th sid of sular zalots as muh as from thologial rationaris. Tak
: with lif's daily dtails (Brook Thomas nry uly, a ky mmr of that Darwinian gingr group, th
l Wr'ln on Way or anothr' thor- lu,who ddiatd thmslvs to driving out of powr Palyits and parsons
hought. alik' No soonr did h har of th atholi St. Gorg Mivart's advoay
of volution than h frrtd out a opy of Franiso Suarz's Sholasti th-
..omplt
ology to insist that volution was in and irronilal antago-
nism to that vigorous and onsistnt nmy of th highst intlltual,
moral,and soial lif of mankind-th atholi hurh'' (quotd in Brook
c hallngsof pi proportions to 1991, 308). Thn, in irritation at th stratgisof thos wh rad Gnsis
..prov that
1izdth osai narrativ of spial throughgologial sptals,h dlard his dtrmination to
..positivly
of using ilial gnalogisto dat rap,murdr 6( arson'' wr njoind'' in th old Tstmnt
zin dsign from natur y dmon- (quotdin Dsmond and Moor 1991, 472). Bsids,h was forvr talking
..hymns
l th ordinary, humdrum prosss aoutth ..sintifipristhood,'' prahing ..lay srmons,'' singing
lmns Wr' in som fundamntal to ration''' dnominating himslf a ..Bishop'' of th nw lsiology, or-
..th ..molular
l moral snsiilitynot in th human dainingfrinds to hurh sintifi,'' and pounding his
ut in th primitiv irpuls of a tlolog.''
righor to th Frnh matrialism At th sam tim advoats of onflit historis hav also had thir own so-
ough Lamarkian-nthusd mdial ial agndasin dlivring pugilisti ounts of sin and rligion. In th
Anglian-Tory stalishmnt. Not asof Drapr' who rmarkd that sin was th twin sistr of th Rfor-
'tl twn sin and rligion r- mation' th real ojt of opprorium Was th Romn atholi hurh; th
th-ntuytratissthat hartd' as dotrinof papal infalliility nuniated in 1 8 70 appalld him and ld him to
lhn Drapr, te Histor of tb on- prphsythat it would ring th atholi hurh into onflit with hithrto-
.!hit' .W.hit,
) or' in th words of Andrw frindly govrnmnts. As for th opposition that his nonstarian
52 uolutio d Rligil
ornll Univrsity had attratd from lrgymn strngthnd his solvto Frnh Dominian pri
kp sin at th forefront of th urriulum; ornll would an asylum o r g i q s( 1 8 8 7 ) ,a n d
fo sin liratd from th striturs of rligius dogma. It was manuvrs rol in th dvlopmnt
of this kind that promptd som historians to ronptualizth so-alld |ished uolution nd Dt
warfar twn sin and rligin in Vitorian soity as a soial struggl Both ths writrs su1
twn two ontnding intlltual lits-namly, th old-fashiond par- it within rtinlimi
son and th nwly professionalizdsintist (Turnr 1978|.In this snario foundations, was dirt
dbats aout sintifi knowidg in gnral and volution in partiular sim. thologial nmis of t]
ply am a furthr arna in whih tussls for ultural suprmay wre might th produt
played out. Sard ongrgation ol
. . v o l u t i o nt h o r yi
Howvr alluring grand narrativs may , storis of th nountr - tt
twn volution and rligion that trad in systmi onflit or, for that mat- human ody'' (quotd
tr' oopration srifi historial intriay t linar simpliity. In fat, ook was nsrdwas
rligious liversrspndd in vastly diffrnt Ways to vlution, and thir spdily rtratd in tht
stans do not follow any straightforward taxonomy of thologial orinta. find ways of rvisiting t
tion' dnominational affiliation, or dotrinl proupation. Bsids,volu. of th ook, ut it too :
tionary thory Was nvr nountrd in a ultural vauum' and dats It was muh th sam
aout it routinly am a mans of giving voi to a rang of othr ani- rognition for his S
tis. Somthing of th divrsity of ths ngagmnts my gland from tb oli Sietists (789
th following thumnail skth. st of 1 as th aU
nd Dogm was to d
..to admir'
muh in it
Th Viw from Rom xx). Likwis ground
ti gsisto justify a
atholi opinion on volution has long n dividd. From th arly days th intrnational suss,a
opposition of a group of Jsuit thinks rought togthr y Pop Pius I to of a fuming iuilt t
omat th fors of modrnity Was prssd through th pags of La lnt with rklss ass
iuiltd ttolic' an outlt that rtd vry onsidral influn among sumptions. Th ongr
ItalianJsuits. Thus in th arly 1860s th nssityof maintaining th fiity vry largly on th ma
of spisagainst Lamarkian idas of transformism was vigorously promul- lutionism was part of z
gatd by G. B. Piniani in its pags (Brundll2001). By ontrast' th fforts tion of ttul ritii
of th nglish omparativ anatomist St. Gorge Jakson ivart' a atholi lsiastial affairs. L
onvrt' to histiniz volution through a Lamarkian rading of transmu. voral rviw h r
tation in his 1871 volum the Gesis of Species was praisd in th pains, prSntlyam l
Catholi prss. Huxly might snip that Mivart ould not a sturdy sldir Various fators play
of sinand a loyal son of Rom, ut Pop Pius I wardd him th dgr i51''-1hg tndny or
of dotor of philosoph ln 1876, and the Blgian ishops prssdhim to tak indpndn and to :
up a hair t th Univrsity of Louvain in 1884, although h latr fll into suh for. This indr
disfavor on aount of rtain thologial writings. ntury atholis wr
Howvr voifrous th opponnts of volution undoubtdly wr, th of- with whih immigran
fiial ustodians of Catholi dogma did not tak any ation against Catholi rns rotatd round t
volutionists until th 1890s whn attituds gan to hardn as th aging arly trnds, notably t
Pop Lo III, hithrto a for for modration, lost ground to th inreasing iology wr thus ofl
influn of Roman Jsuits and th mahinations of th iuilt ttoli o- to stablishd ways o
tri. So whil ivart's fforts sapd at last for a tim th nsur of th authority and who h
Vatian' othr atholi volutionists did not far so favoraly' notaly th Thos who found v
t,ollttio d Rligio .].'.]
gymn strngthnd his solv to Frnh Dominian prist M.-D. Lroy, author of L'uolutio des esps
lum; ornll would an asylum orgiques(1887), and th Amrian Jhn Zahm, who playd ir signifiant
.ligiousdogma. It was rol in th dvlopmnt of th Univrsity lf Notr Dam in Indiana and pub-
manuvrs
Lsto ronptualizth so-alld |ised,uolutiod Dogm in 1896.
torin soityas a soial struggl Both thswritrs suppotd th th;logialaptaility of volution' al-
_namly, th old-fashiond par- it within rtain limits' Lroy's apologia, rtd foursquar on Mivart's
st (urnr 1978). In this snario foundations,was dirtd against both thisti rnditiorrsof volution and
al nd volution in pzrrtiularsim- thologialnmisof th thory. But aush hintd that th human ody
sls for ultural suprray wr might th produt of volutionary fors, th ardinal prft of th
SardCongrgation of th Indx of Prohiitd Books issud th judgmnt
' , storis of th nountr - ..volution thory
that is tmrarious and anti-hristian whn pplid to th
s,vstmionflit or, for that mat- human ody'' (quotd in Brundll 2001' 88). Although th finding that his
ily to linar simpliity. In fat, ook was nsurdWas onvyd to him privatly' Lroy was dvasttdand
]ntWys to volution' and thir spdilyrtratdin th pags of Le ond in arh 1895. Latr h trid to
taonomy of thologil orinta- find ways of rvisitingth whol issu y prduing a nw' orrtd vrsion
tl proupation.Bsids' volu- of th ook, ut it too fll foul of th uthoritis.
a ultural vauum' and dats It was muh th sam with Zahm, who had alrady ahivd widsprad
rg voi to a rang of othr anxi- rognition for his Sottd d si (1892) and tholi Scine d
ngagmntsmay glarrd from tholi Scietists(1893). Although h had arlir ondmnd Darwin's De-
sntof as th aus of soial inquity, his purpos in writing uolutio
d Dogm was to dlonstratits thologial aptailiry.Indd h found
..to admir' muh
muh in it that is nnoling and inspiring,' (Zah-l,|896,
x). Likwis groundd in ivart's sstm' Zahm engagd in dtild patris-
ti xgsisto justify a tlologial rndring of volution. Th ook was an
n dividd.From th arly days th intrnationalsuss'and an Italian translation rapidly appard,ut radrs
lught togthr y Pop Pius I to of a fuming iuilt ttoli wr assurd that it was all a tissu of lis' rdo-
'rssd through th pags of L lnt with rklss assrtions' and hoplssly ompromisd y duious as-
:ry onsidralinflun among sumptions.Th ongrgation of the Ind dnouned tt in 7897, fousing
nssityof maintaining th fiity vry largly on t mttr of human origins. To traditionalists' Zalrm's vo-
sformismwas vigorously promul- lutionismwas part of a pakg of radil proposals tlrat inludd th adop-
tll 2001). By ontrast' th fforts tion of txtual ritiism and th ir-rrasingsgrgation of dmi and
orgJakson Mivart, a atholi lsiastialaffairs. Lik Lroy, Zam withdrw his fforts dspit th fa-
r Lamarkian rading of transmu_ voral rviw h rivd from th nglish ishop John Hdly, who' for his
is of Spis was praisd in th pains, prsntly am undr th whiplas of iuilt ttoli tongus.
.,Amrian-
vat ould not a study soldir Various fators playd thir parts in ths intrigus. A far of
Pius I awardd him th dgr ism''-th tndny on th part of Amrian Catholis to display intlltual
:lgiarrishopsprssd him to tak indpndnnd to adapt atholiism to arr Amrian OntYt-was on
1884, although 1rltr fll into suh fr. This indits that Darwinian ontrovrsisanlong turn-of-th-
rritings. nturyatholis wr oftn lss aout sinthan aout issus of idntity
llution undoutdly wr' th of- with whih immigrant ommunitis wr grappling. Anothr suit of on-
tak any ation against atholi rnsrotatd around th routin onflation of Drwinism and modrn shol-
s egan to hardn as th ging arly trnds, notaly th nw ilial itiism. Rations to volutionary
i o n . I o s tg r o u n d t o t h i n s i n g iology Wr thus oftn all of a pi with rsponss to widr hallngs
rtiorrsof te iuilt ttolicct o- to stablishdways of thinking. ritil too Was th mattr of intlltual
astfor a tim th nsu of th uthorityand who hd th right to intptth tradition rivd anon.
ot far so favoraly, notaly th Thos who found vlution in th writings of Aquinas nd Augustin wr
54 t,oltio d Religio
pr-istntand living mattr'' was allowd' anv suggstionthat th humn th imag of a divin .
ra Was f plural origins was asolutly ond-rndausof its implia- ontraSt' Samul Wil
tions for th dotrin of original sin. A similar stipulation prtaind to th ally and philosophial
origin f th lruman sor.rl,whilr Pius insistdwas yond tlr rh of vlr'r- ].860 mting f th
..Th Franis Opn ois,
tionary transformisnr: atholi faith oligs us to hold that souls ar
immdiatly ratd y God'' (Pop Pius II 1950' [36). shir, who authord
or rntly Johrr Pul II addrssda plnry irssmlyof th Porrtifial Drinism (1869) and
Aadmy of Sinsin Otor 1,996r>nth sujt of volution and its sig. thought tht Darwinisr
nifian for undrstandingth human agnt. Ralling Pius II's nylial th Atlanti th lrgy
and noting that evolution had now gon yond th status of a mr hvpoth- gllags in Boston Tho
sis, h ondd th liklihood that th humn ody had originatd fronr hoi napsulatdin
prxistnt living mttr, ut h still rstatd th tditilnl rationist- ntion too should bt
as opposd to trduinist-dotrin of rlr rigin of th human soul irr ordr doyn of Amrian Prs
..th ontologial lap'' twn humanity and its Drtuiism?(l874) wit
to sustain what h alld
forars.By this mov mtaphysialmtrrs rotilting around morality, on. a judgmnt on Darwin'
sin,frdom' asthtippriation, and spiritual prinwr dl- ltion itslf.
gtd to phiiosopial anlysis rathr than to volutionary iologv aus Th swp of opinior
..momnt Th nglish Nononft
t of transition to th spiritual'' was simply not opn to mpirial
..osrvation''(Pop Thomas R. Birks, tlok
Joanns Pulus II 1996, II 6).
Not surprisingly, this statmnt has attrt]th ttntion of atholi Spnian form, in bo
ommntators. on th mor rationary sid, John Marthy osrvdthat uolutio pulishd in
th addrss had gnratd dismay and onfusion among th faithful. H ity's ultimat hoi wa
thrfr workd hrd to minimiz any opnnssto volution that th po- Seltio' His fllorv ot
nounmnt might foster y th introdution of a numr of srplrtinin- ously sought to Drwtn
trprtations and y rminding his radrs that mmrs of th Pontifial ( l88). Lss spulat
Aadmy of Sinswr hosn rgardlssof ligious rd. By ontrast' who tok up th ps
in 1998 th Vatian astronomr Gorg V. oyn Warnlly mrird th Univrsity) in 1868, t<
pope's announmnt of evolution's fatual status and wnt on to urg that Allian that instadof
..ontinuous ophrs would ttr
sin volution ws tlr mthod of divirr rtion' th ida of
ration'' Was th st way to oniv of th mrg of th human fllow Sotsmn, th th
spis (oyn 1998, |60). ah, and Hnry ldr
of thisti vlution. I
insistd in th first d:
Th Protstantosai lution was simply anot
This pathwork of l
Protstarrtrsponssto volution Wf no lss divrs.From th arlistdays around 1900 is math
thr wr thos who snsd in volution th mthod of God's modus Wer onntd with vl
oprandi in natur. Anglian pi1rson'novlist' and hristian soilist harls own mntor, th Angli
Kingsly, for arrrpl'Was nthusistiabout the rigi irnd wrot to Da- p r s s d a r l y o p p o s i t i
win tlling him that although th thory mant that h must giv up many of wik nd th Sotti
..just mathmatial physiis
th things h livd,it was as notal a onption of Dity, to -
liv tht H ratd priml forms apal of slf-dvlopmnt''(Brrrkhardt physial omputations
and Smith 1985_2005, 7: 380). Simllarly, Frdrik Tmpl, latr to om tim ndd for th o
arhishop of Cntrury, pssdhis support as arly as 1860, ontnd- And infamously th Pl1
ing that a world md to mak itsl{attrllhirndth rator's noilit. rspondnt of Darwiir
To suh figurs, a univrs govnd y lratural law was nror dignifying tha volution and rlid o
uolutio d Religion 57
ld' any suggstionthat th human th imag of divin onjurr who mgikd nw spis into xistn. By
.!7ilrfor
:ondmndausof its implia- ontrast, Samul thought that Darwin's thory was oth mpiri-
milr stipulation prtaind to th ally and philosophially unsound and famously pourd sorn on it at th
d was yond th rah of volu- 1860 mting of th British Assoiation for th Advanmnt of Sin.
L oligsus to hold that souls ar Franis Orpn orris' Vitorian ornithologist and Anglian rtor in York-
: I I1 9 5 0 ' I3 6 ) . shir' who authord a squn of pamphlts with titls lik Diffiulties of
plnry assmlyof th Pontifial Dariism (1'869) and The Demnds of Dr,iism o redulit Q890),
h sujtlf volutin and its sig. thought that Darwinism dsrvd only uttr ontmpt and drision. Aross
nt.Ralling Pius II's nylial th Atlanti th lrgyman Luthr Tray Townsnd, who taught ilial lan-
:yond th status of a mr hypoth- guagsin Boston Thologial Sminary, plad for his radrs th stark
human ody had originatd from hoi napsulatd in th titl of his trat F,uolutio or retio (1896).
atd th traditionlrationist- ntion too should mad of th Printon thologian Charls Hodg,
: origin of th human soul in ordr doyn of Amrian Prsytrins, who answrd his own qustion in.Wht ls
l iap'' twnhumanity and its Drtuiism? (1874| with th trs quip ..It is athism''' though this was mor
rs rotating around morality, on- a judgmnt on Darwin's antitlologial stan than on th prinipl of vo-
nd spiitual prinwr dl- lution itslf.
n to volutionary iology aus Th swp of opinion rprsntd hr ould rpatd many tims ovr.
' Th nglish Nononformist and ofoundr of th vanglial Allian,
was simply not opn to mpiril
)6,\ 6). Thomas R. Birks, took up th udgls against volution, partiularly in its
:tratdth attntion of tholi Spnrianform, in ooks lik odr Phsil Ftlism d tbe Doctrin of
id, John arthy osrvd that uolutio pulishd in \876. Th Sotsman.lfilliam Millr flt that human-
:onfusion among th faithful. H ity'sultimat hoi was' to us th titl of his 1897 volum' God, or Nturl
pnnssto volution that th pro- Seltio.His fllow ountryman Hnry Drummond' on th othr hand, vigor-
. i o n o f a n u m o f s r p n t i n i n - ously sought to Darwiniz thology inhis Nturl L, i the Spiritul World
rs that mmrs of th Pontifial (1883). Lss spulativ and mor traditionally alvinist, Jams osh,
ss of rligious rd. By ontrast, who took up th prsidny of th ollg of Nw Jrsy (latr Printon
V. oyn warmly mrd th Univrsity) in 1868, told th 1873 Nw York mting of th vanglial
al status and wnt on to urg that Allian that instad of dnouning the thory of volution, hristian philos-
..ontinuous ophrswould ttrmployd in pounding its rligious dimnsions. His
ration,th ida of
of th mrgn of th human fllow Sotsmn' th thologians Rort Rainy, Gorg athson, Jams lvr-
ah, and Hnry aldrwood, all addd thir vois in support of som form
of thisti volution. Latr th Amrian Baptist thologian A. . Strong
insistdin th first dad of th twntith entury that th prinipl of vo-
lution was simply anothr nam for hrist.
This pathwork of Protstant opinion from th thologians in th dads
lssdivrs.From th arlist days around 1900 is mathd y th pattrn of rspons mong sintists who
ion th mthod of God's modus Wronntd with various Protstant dnominations. In Britain Darwin's
list' and hristian soialist harls own mntor, th Anglian lrgyman.naturalistJohn Stvns nslow, x-
out th rigi and wrot to Dar- prssdarly opposition' as did th amridg gologist Rv. Adam Sdg-
lant that h must giv up many of wik and th Sottish natural philosophr David Brwstr. Th Irish
lal a onption of Dity, to b. mathematialphysiist at Glasgow, \/illiamThomson (Lord Klvin), whos
of slf-dvlopmnt''(Burkhardt physial omputations of th ag of th arth smd to dny Darwin th
Frdrik Tmpl, latr to om tim ndd for the oprations of natural sltion, rmaind antagonisti.
supportas arly as 1860, ontnd- And infamously th Plymouth Brthrn naturalist Philip Hnry Goss, a o-
lly nhandth Crator's noility. rspondntof Darwin and admirr of l/alla'was implaaly opposd to
tural lw Was mor dignifying than volutionand rlid on th ida h put forward tn mphIos (1857) (th
3.'8 uolutil d Rligil
Grk word f<>r ul) tht God had ratd a matur univrs with th sm- infiltration. To a on
lan of ag-prohronism, as h lld it. Aross th Atl:rnti th Swiss g- sin an trad
ologist and physial gographr at rsytriarrrintorr, Arnold Guyot, tist Gorg Mrd
and his anadian ollntrpart Jolrn Villiam Dawson, prsidnt of Gill Wekst Poit i the
Univrsity, also st thir fas against Darwinism. In th mid 1880s Guyot His flood gology
advoatd a kind of pr-Darwinian lvlopmntalismthat, in th fashion of apologist, lln \/hit
th Sottish stonmason Hugh illr, rad th gologial pohs into th litralist intrprtatio
dys of Gnsis, whil Dawson alkd at Darwinis]'s antitlologial alon, and h riv
stan. By ontrast th Llarvard otanist, Congrgationalist, and onfidant slf-styld rsarh s
of Darwin Asa Gray foLrnd oth odg's and Dawson's ojtions odd tor.William Bll Ril
and workd hard to fit volution into tlologial garmnts y arguing tht was also forthomin1
it was rnistakn t rstrit th id of dsign to th suddn flashing of a rati andidat for
Palyit wth into istn.Anothr ongrgationlist' th Orlin golo- paifist, whos fars
gist Gorg Frdrik Wright, also gav volution his ndorsmnt in th Grmani militarism
arly 1880s' vn if h wavrd ltr in lif, i.rndwnt so far as to draw a st th lrtd Sops
of rlling analogis twnCalvinism and Darwinism, onluding that Dar- ron traordinair or
winism was ..the alvinisti intrprtation of natur'' (\/right1882, 255). Darrow. Of ourse, tt
othr supplrtiv ommntaris. though with varying dgrs of nthusi- absnt from ths th
asl' wr fothoming from suh sintifi pratitionrs as th alifornia phlt titld Poisoig
gologist Josph Lont, th Yal minralogist Jams Dana' th Mihigan volutionism togth
.sinhll,
goIogistAlandr and th Printon lrgyman.iologistGorg volutionary transfor
aloski' Indd, figurs lik ths did a good dal to kp lins of om- and politial.In Brita
muniation opn btwn spialist sin and radrs of dnominational ship of Amros Flr
magzins. vrsity Collg Lond
A a t a l o g o f s u h p a r t i u l a r i t i s o u l d l a o r a t da d l i i t u m , u t a s a n allowd for th poss
ovrall grrrlization' most Protstant thinkrs livd that volution Indd, in Flming's t
ould mrad without aandoning dotrinal ssntials.Th majority ration of th ili
found ways of aommodating thir thology to mor or lss rvisd vr- al findings2 a mov t
siorrs of volutionary thory. As wll as hallnging widsprad assump- wrong way.
tions of systllihostilitis twrtvolutiol-land rligion, this trjtory Whatvr th sour
srvs to all attntion to a numr of sming inongruitis. Thus num- motion dlind fair
rd among th writrs of Th Fudmetls, a squneof 72 pamphlts though it promptd
i s s u d t w n 1 9 1 0 n d 1 9 1 5 t h a t g a v v o i t o r l . vF u r - r d a m n t a l i s m , Histor of No-eu
Wr svral writrs, inluding G. F. Wright and th Sottish thologian tnt and up.to-dt lil
Jams Orr, who advoatd volution in on form or anothr. There is also ayan until th 19.
.Warfild,
th s of th Printon thologian B. B. ustodian of th onfs- iblial thlogian Jo
sioral standards of trditional lvinism and rnodrn ahitt of th do- am th foundtio
trin of bilial inrany, a mainstay of th Amrian Fundmntalist Geesis Flood. |t |im
.!arfild
movmnt. Brought up on a Kntuky stok farm' knw firsthnd nd rassrtdrati
aout inhitd variation irr animal rding arrd latr musd that h was its various dtion
lrady a ..Darwinian of th purst watr'' bfor th oming of Jams lswhr in this volr
osh to Printon (\Wrfild \976, 652). In 1915 h limd that polmis and intrn
alvin's dotrin of rtion was in fat a ,.vy pur volutionary shm'' day rationism has d
( W a r f i l d1 9 1 5 , 2 0 9 \ . ally' In fat, th th
Dspit this plurlism aross th dnominational sptrum' rtain s- ontinud to dfn
tarian strands in Protstantism rraind staunhly rsistant to Darwinian dnominational rang
uolutio d Rligio 59
l a matur univrs with th sm- infiltration. T a onsidral dgr th mrgn of modfn ration
, Aross th Atlanti th Swiss g_ sinan trd to th nrgti ativitis of th Svnth-Dy Advn-
trin Prineton, Arnold Guyot, tist Gorg r:rdy Pri, author in ] 906 of Illogil Geolog: Tb
Lm Dawson, prsidrrtof MGill Wkst Point in the uoltio Theor and irr 192 of Th N Golog'
winism. In th nrid 1880s Guyot His flood glogy prssd th -lr4osaisin of th movmnt's hif
lmntalismthat, in th fashion of apologist, lln Vhit, whos dotrin of th Saath was groundd in a
d th gologial pohs into th litralist intrprtatin of th ration narrativ. Pri, of ours' ws not
at Darwinism's antitlologial alon, and h rivd noriragmntfrom th Prsyterian ministr and
ongrgationalist,and onfidant slf-styldrsarh sintist Harry Rimmr and th ]V1innsotaBaptist ps-
's and Dawson's oitions odd .William
tor Bell Rily. Support, though from a diffrnt idologial stal,
llogial garmnts y arguing that was also forthoming from William Jnnings Bryn' thr tims Dmo-
sign to th srrddn flaslring of a rati andidat for th prsidny of th Unitd Stats and ommittd
:lgrgationlist, th ()rlin golo- paifist' whos frs about th sming allian twn Darwinism and
:volutionhis ndorsmnt in th Grmni militarism nouragd him to tirk up th proseution's s in
, and wnt so far s to dfaw a st th lratdSops trial of |925.|t arnd for him th rputation of mo-
Darwinism, onluding that Dar- ron traordinair ourtsy of th polmis of his fisty opponnt' larn
t o f n a t u r ' '( w r i g h t 1 8 8 2 ' 2 5 5 ) . Darrow. of ours, this is not to say that mattrs of politial idology wre
with varying dgs of nthusi- asntfrom ths othr ontrovrsialists.Pri, for instan, issud a pam-
i pratitionrsas th alifornia phlt titld Poisoig Demor 1921) in whih h nttd soialism and
LlogistJams Dna' th ihigan volutionism togth as th mainspring of modrn-day ills. His loathing of
ton lrgyrnan-iolgistGorg volutionary trnsfortnisnr was thtls mr-rlridirrrrrsional-bilial,sintifi,
' good dal to kp lins of om- and politial. In Britain th volution Protst ovmnt' undr th ladr-
: and radrs of dnminational ship of Amros Flming, FRS, profssor of ltrial thnology at Uni-
vrsity ollg Lndon, lakd th vitriol of its Amrian ountrpart and
laoratdad liitum' but as an allowd for th possiility of som divinly guidd volutionary hang.
hinkrs livd that volution Indd,in Flming's own as h was vn prpard to rd aspts of th
l o t r i n l s s n t i a l sT. h m j o r i t y rtion of th ilial Adam in th light of various paloanthropologi-
logy to mor or lss rvisd vr- al findings, a mov that ertainly rued th anatomist Arthur Kith th
hallnging widsprd assum- wong Way.
r t i o n a n d l i g i o n ,t h i s t r j t o y !hatvr th sours of ths sntimnts' this flurry of rationist om-
minginongruitis. Thus num. motion dlind fairly dramatially in th yars aftr th Sops trial,
tls,a squnof 12 pamphlts though it prornptd th puliation of . T. Brwster's 1,927 rtio:
: voi to arly Fundamntalism, Histor of No-uoltior Thoris to alan th ..ampl and omp.
ight and th Sottish thologian tnt and up-to-dat litratur of volution.'' rationist agitation Wnt into
n form or anothr. Thr is also abyanuntil th 1950s whn th hydrauli ngineernry Morris and th
Warfild,ustodian of th onfs- bilial thologin John Whitom joind fors to produ in 196l what
and modrn arhitt of th do- amth foundational txt of th modrn ration sinmovmnt' l2e
lf th Arriirr-r}-undamntalist Gensis Flood. It limitd goiogial histoy to no mor thn 10,000 ys
k farm, Warfild knw firsthand and rassrtdration in six litral dvs. Th taitory of this movmnt'
ling and latr rusd that h was its various duational mahinations, nd its lgal Stratagmsar hartd
r'' for th oming of Jams lswhrin this volum. It suffis hr to add that dspit its ggrssiv
652). In 1915 h laimd that polmisand intrnational suss' it would mistakn to think that si-
..vrypur volutionary shm'' day rationismhas dominatd onsrvativProtstant thought mor gnr-
ally. In fat' th thory of volution, in on vrsion or nothr, has
rlinationaI sptrllm' rtain s- ontinudto dfndd y Protstant thologins and sintistsaross th
staunh[,vrsistant to Darwinian dnominationlrarrg.
60 uoltio nd Reliil
for rligius institutions and ustoms. \/hat was oftn at stak in thes d-
n
batswas th natur of Judaism itslf.
Somthing of th sam sptrum of rations is also disrnil in Islam,
r oftn ound up with roadr
though in th Islami as a rathr mor prvailing litralisti hrmnuti of
rnd faith ommunitis, not last
th Qur'n has mant tht advoats of volution ar lss ommon. Nvrth-
r g w i d r s o i t yo r h o w t o r t a i n
lss'Islami support for volution is not ntirly asnt. uhammad Iqal,
hallngs.Jwish dbats aout
for ampl, has laimd that th ida of volution originatd in mdival
a ar illustrativ, for ths wr
Islam, whil th Lans sholar ossin al-Jisr insists that th thory is n-
r th futur shap that Amrian
tirely ompatil with Qur'ani thology (Majid 2002).In th as of Ahmd
'ionalist,Modrat Rform, and
tfzaa| (7996), who likwis ontnds that ..th thory of volution sprouts
lw th Jwish ommunity should
from th Holy Qur'an itslf,'' a ruial distintion is prosutd twn Dar-
:mrdbausit ould mo-
winism and volution. Th formr h finds unaptal on aount of th
lsformation.Thus th radrs of
limitd appliability of ntural sltion, whras th prinipl of volution
ts during th 1870s and 1880s ..fitsprfetly in
th ovrall shm of God's ration as dsid in th Holy
'lal ralm and that Jwish ritual
Book.'' Indd' dploying th standard soul-ody dualism, fzaa| argus for
<pin stp with modrn sin.
an volutionary aount of th human spis that alloats physial dvlop-
lopmntonfirmd th prinipl
mnt to naturl prosss and th mrgn of slf.onsiousnss to divin
prvilingnvironmnt applid
ration. In this mov inrasingly humanoid forms-Homo hbilis, Homo
From th sam thologial stl
rtus,and so on-dvlopd to a point whr ..Almighty Allah (S!) s-
rit|eduolution d Judism and
ltda singl pair-a mal and a fmal-and ndowd thm with thir spir-
)n to th growth of rligion, th
itual souls.'' Not surprisingl, fzaa| found inspiration in th antimhanisti
l, th ida of God, and th dvl-
aountsof figurs lik Tilhard d hardin and Hnri Brgson.
Dspit ths irni fforts, vigorous opposition has n forthoming
d Darwinism or in som ass
from writrs lik T. H. Janai who laim that volution is without mpirial
ology.on th on hand, Samul
warrant and is promotd for soial and politial rasons. Similarly, Sha-
: Jetuish ssngr to launh at-
habuddin Nadvi issud rfutation of volution in his 1987 aount of th
La,a anadian immigrant' mar.
ration of Adm, whil Adm Tatli's uolutio: Bkrupt Theor 0990)
logist John !illiam Dawson to
inorporatd mattr originally prsntd to th Turkish govrnmnt. But pr-
,,oatof th viw that God had
..vo- haps th most influential Islami riti of Darwinism is Harun Yahya, rport-
radrsof his 1887 artil
dly an minnt Turkish sholar, who has n so prolifi in produing
3 was not a shrd f vidn for
antivolutionarytrats (\tkeWb Drluiism ls Inomptible ith th Qur')
n th othr hand' th traditional-
that his opponnts laim that th works ar atually th produt of a group
numrof whom wr traind in
of writrs. In any as, in te \999 ook Th uolutio Deit Yahya hit out
histi vrsion of volution along
at th ..dishonstphilosophy'' of ..matrialism''that ..sksto aolish th a.
>pald to th gradualism of vo-
si vlus on whih th stat and soity rst'' (Yahya 1999, 7). Dawinism,
ping ovrthrow of inhritd insti-
th trat urgs, providd a mythi foundation for matrialism's laims to si-
rntlssgnrations of traditional
ntifi status, thry plaining why arx rootd dialtil matrialism
.ish strugglfor survival against
in natural history. In anothr 2001 rharsal of ..th disastrs Darwinism
rought to humnity,'' Yahya has attriutd war, povrty' pain, and mas-
r rabini nountrs with volu-
sarto th malign influn of ..th selfish and pitilss world viw'' of Dar-
;ntimntsnvr dirtly mappd
winism on figurs lik Stalin' Trotsky, ao, Pol Pot, itlr' and Mussolini
s mans that th sintifi dimn-
(Yahya 2001). Translatd into mny languags' th output of th Harun
Jrssd;rsponsswr mdiatd
,nthat matrialism had on low- Yahya mind-st, partiularly in Turky, has n rportd as immns, with
's rationistooks oming mor influntial than standd ttooks in r-
impliatins for th natur of
tain plas (Konig 2001). To sur, ths movs hav n rsistd y a
s livd its aDtan ntaild
numbr of Turkish Darwinians, notaly th iologist Aykut Kn and th
62 uolution nd Rligion
mdil gntiist Isik Bksoy. But marshaling support for Darwinian vo- divin wisdom and powr
lution has provd diffiult in a ontt whr politial frs hav fostrd wrot inquiry int th ni
th tahing of rationism in shools and whr dirt support has n pro- vrs. In rspons' num
vidd y Amrian Protstant rationists(dis 1994). tlologially. In gnral,
sion had a mor diffiult
plans and th oprations
Ruring Thms thusiast for th way Argyl
divin wisdom mor in h.
Various historial modls hav n anvassd in th attmpt to gt a handl Th Duth alvinist, fun
on th rlations twn sin and rligion. Our travls thus far onfirm that o f - t h - n t u r yp i m m i n
any simpl portrayal of th rlations twnvolutionary thory and rligious zoologist Jan Lvr (195
lif is doomd to fall foul of th mssinssof history, ut a numr of prsis- whih onivd of natur
tnt thms hav rurrd, alit in diffrnt wys in diffrnt sttings. ours' this dos not ma
Qustions of sriptural hermeeutis hav routinly surfad in disussions tirly disappard. Th nt
of volution. Som adhr to a litralisti xegsis of ration narrativs, surfad in rnt tims st
whil thos who ar intrstd in onordism hav strssd th rol of mta- has n alld irrduil
phor and ultural ontxt in th intrprtation of partiular statmnts. an all-or-nothing varity l
Thos who ar willing to rlgat th osi narrativ to th ralm of prim- ations on whih natural s
itiv mythology hav n ngagd no lss in a hrmnuti undrtaking. i n i t s s t t i s t i a lw i z a r d r y '
ithr Way' qustions of gsis hav prsistntly otudd. Philip Hnry pliations of Darwin fr .
Goss's whol systm of prohronism was dsignd to prsrv th litral a- thr ontinus to b phil
uray of th Bil's timsal.By ontrast, th Orintal and ilial sholar th diffrnt positions ad<
\. H. Grn rlivd rtain strands of nsrvativ rotstantism of th tinga (199 l),who dfnds
urdn of Usshr-typ hronology (whih famously datd th world's origin rnan MMullin (1991I,
to 4004 B...) whn h dmonstratdin 1890 that old Tstamnt gnalo- ural law. Th fat that
gis ould not rlid on to dat primval tim on aount of thir stratgi s o u n d i n g v o a b u l a r yu r g i
omissions from th rords. That gav som thisti volutionists th y . t r i l a n d o . ' h ti t
.loo1
hrmnuti room thy ndd. Again, th atholi writr Dorlodot dvotd y snaking through
onsidral spa to luidating atholi thoris of inspiration in ordr to t i n s i n o r d t o i m p r o v i
allow for his onvition that th thought-forms of siptur wr aommo- whol issu (Grne 1989
datd to th ultural onditions of th tim. And th Islami volutionist Aross th rligious ran
Ahmd fzaa| urgd th lgitimay of xplaining ..th rlvant Qur'ani th sptr of mterilisn
vrss in trms of a mtaphor or paral'' and spok of ..th Qur'ani lgend that volution is rtd
of th .Fall''' (t{zaa| 1996). In on Way or anothr' volution thory has moral natr and has u
prsistntly ralld attntion to hrmnuti mattrs in th intrprtation of various strnds hav also l
anint sard tts. for instan, th Catholi
From its arlist days Darwin's thory onfrontd had-on th issu of tl' A Stud of Vitlism d
olog. As Darwin himslf put it at on point, ..Th old rgumnt from dsign outlook of th primn
in ntur, as givn by Paly whih frmrly smdto m so onlusiv' fails, th antirdutionism pr
now that th law of natural seltionhas n disovrd'' (Darwin 1888, 1: and P. G. Tait, whih first
309). Th onsqunsWr potntially monumntal, for natural thology John yndall's infamous -
had undrsord th fundamntally moral haratr of natur' nthusing - linations, Windl Was at
livrs, uring starian tnsions' end inspiring mn and womn to study mattr' and h rmaind t
.!0.hig, hanial univrs. By on
God's handiwork. !hthr from th pn of \/illiam Paly, a modrat
whos Nturl Thology (1802) was dvourd y Darwin, or in th multi- sondary uss-that G
volum Bridgeter Trtises (1833_1836), whih st out to dmonstrat f o r m o f m a t r i l i s m o n s i
uolutio nd Religil 6
ling support for Darwinran vo. divin wisdom and powr in ration, th natural thology ntrprisundr-
r politial fors hav fostrd wrot inquiry into th natural ordr. Darwin dstailizd this whol uni-
rrdirtsupport has n pro- vrse. In rspons' numrous writrs workd hard to intrprt volution
dts 1994). tlologially.In gnral,thos who wr still nthusiastiaout Paly's vr-
sion had a mor diffiult task than thos who found dsign in arhtypal
plans and th oprations of natural law. Thus Jams osh, alrdy an n-
thusiastfor th way Argyll's b Reig of I-tu rloatd divin dsign, found
divin wisdom mor in homologial struturs than in spifi adpttions.
Th Duth alvinist' foundr of th Fr Univrsity of Amstrdam, and turn-
:d in th attmpt to gt a handl
our travlsthus far onfirm that of-th-ntury prim ministr Arham Kuypr nd latr dvots lik th
zologist Jan Lvr (1958, 229) spoke of ..divin volutionisti ration'''
volutionarythory and rligious
lf history' ut a numr of prsis- whih onivd of natur unfolding aording to a prdtrmindplan. of
ours'this dos not man that advoats of Paly's mhanial vrsion n-
ways in diffrnt sttings.
r o u r i n l ys u r f a di n d i s u s s i o n s tirly disappard. Th nortionist Intllignt Dsign movmnt that has
surfadin rnt tims sks to idntify trnally drivd purpos in what
gsisof ration narativS'
has n alld irduilomplity, namly, rtain iologial systmsof
n hav strssdth rol f mta-
tation of partiular statmnts. an all-or-nothingvarity whose funtions annot rdud to simplr opr-
ations on whih natural sltion an work. In its nptual strutur, if not
narrativ to th ralm of prim.
in its statistialrwizardr,this rads lik Paly rdivivus. In ny as th im-
; in a hrmnuti undrtaking.
pliationsof Darwin for dsign hav maind a ky arna of date. That
sistntly otrudd. Philip nry
throntinusto philosophial milag in this whol mattr is lar from
signdto prsrv th litral a-
th diffrntpositions adoptd y suh minnt philosophrs as Alvin Plan.
th orintal and ilial sholar
lnsrvativrotstantism of th tinga(1991),whodfndsdivine intrvntionismin th history of natur, and
rnan Mullin (1991)' who inlins mo toward God ating through nat-
rmously datd th world's origin
ural law. Th fat that iologists hav frquntly tradd in tlologial.
]90 that old Tstamnt gnalo-
soundingvoaulary urging ..that volution is .opportunisti', that it prods
r i m o n a o u n to f t h i r s t r a t g i
y .trialand rror', that it rahs.dad nds' or aomplishs.rakthroughs'
som thisti volutionists th
]atholi writr Dorlodot dvotd by snakingthrough .loophols', or that natural sltion .an rmodl pro-
horisof inspiration in ordr to tinsin ordr to improv intrations' '' adds yt furthr ompliations to th
lrms of sriptur Wr aommo- w h o l i s s u ( G r n 1 9 8 9 ' 4 0 8 ) .
Aross th rligious rang opponnts of Darwinism hav n hauntd y
n. And th Islami volutionist
..th rlvant th sptr of mterilisz. Islami opposition springs in part from a sns
'plaining Qur'ani
..th that volution is rtdon a matrialist rd that donstrutshumanity's
nd spok of Qur'ani lgnd .Wstrn
moral natur and has ultivatd a dadnt ultur. hristians of
)r anothr' volution thory has
variousstrandshav also n trould y similar onrns.Brtram.!indl,
i mttrs in th intrprtation of
for instan' th Catholi advot of volution and author of Wbt ls Life?
Stud of Vitlism d Neo-uitlism (\908), prssd sympthy for th
rfrontd had-on th issu of tel-
..Th old argumnt from dsign outlook of th primntal physiist and spiritualist Olivr Lodg and for
t,
th antirdutionism prssd tn The Usee Uiuerse of Blfou Stwart
smdto m so onlusiv' fails,
and P. G. Tait, whih first appard anonymously in 1875 immdiatly aftr
: nd i s o v r d '(' D a r w i n 1 8 8 8 , 1 :
lonumntal, for natural thology John Tyndall's infamous Blfst addrss. An volutionist of Lamarkian in-
linations,Windl was attratd to th ida of a Vital Powr that anrmats
:haatrof natur, nthusing b-
piring mn and womn to study matter'and h rmaind hary of sintifi inlinations toward a purly m-
!illiam Ply, a modrat.Whig, hanial univrs. By ontrast, numrous writrs hav usd th dotrin of
sondaryauss-that God works through natural agnis-to argu for a
urd y Darwin, or in th multi. .lfarfild,
form of matrialism onsistnt with th Judo-hristian hritag. for
i). whih st out to dmonstrat
64 uoltio nd Rligion
xilmpl, insisd that,,a omplt systm of naturl ausation'' was ntirly told tlr radrs of his 1
..tlologialsystm implis a om- ..uslss
onsistnt with tlology and tlrat vry to pt ptlpl
plt .auso-mhanial' explanation as its instrrrmnt'' (quotd in Livingston trs of th Bil, if thy d
and Noll 2000,301). Diffrnt though all ths rsponssundoutdly ar, its first hapters'' (Pi1
thy ar witnss to matrialism as a rurring lous of dat. Th ralization that th
losly onntd with ths onrns is a st lf qustionstht ongrgat tIu"tl L'olutiontlI srip
'l
aror.rndth implizrtions of volutiorr for th ltLlrnd morlit of the a fr'rrthrpoint of issu.
hm get. Vaious stans hav n adoptd on th issu of whthr puliation of Darwin's (
human ings bu sods or ar souls. Som hav optd for th viw that th th Bil had to intr
soul is always th rsult of an at of divin ration. Thus th atholi volu- was widly doptd v t.
tionist rnst ssngr osrvd in his 1931 ount uolutio d Theol. prinipl of volutiona
..no hypotlrsis manating fr
og t,at Ctholi thologian or philosophr ould possibly allow that
th spiritual soul of man has volvd from a brrrtesoul, muh lss from inor- tion atd no stir wht
gani mattr. Th soul is immatrial, and an only om into istny a no among thosnthus
dirt at of ration'' (ssngr 19\, 14)' Othrs hav thought that th By th sam tokn, as wl
id of th soul mrging in som way from arlir forrsis ntilyom- applid to oth th instit
ptil with thir thology. As for th thial rprussionsof Darwinism, to th way in whih D:rr
som hav oild at th suggstionthat moral snsiilit has n shapd Jws. Suh uss of volut
y th imprativs of a struggl for survival. In suh a snario, as \William ing to many. Gorg Frd
..Grat in good part from his
Jnnings Bryn, th ommonr,'' mad lar, thr is no rason to
xpt humans dsndd from animals to hav any diffrntly from ani- mntay history. In S
rnals. Othrs' nthusd with th possiilitis of lsmi volution, hv sught, Williar Rortson Smith
lik Hnry Drummond, to us th languag of volution to mak sns f spookd onsrvativl
thial and spiitual dvlopmnt. Again, although diffrnt rligious om- hang smd thr tr
mntators hav takn diffrnt positions on th qustion, th issu of this attituds to Darwinism .
and volution has prsistntly manifstd itslf. thory's rlvan for ro
Howvr amiguous in Darwin's own thinking, th idas of progress and,
in rtain strands of rligious thor-rght,its thologial nxt-door nighor
shtolog hav frquntly ntrd th disssion. Som of ths assoiations Putting volution ar
wr idntifid y rnst Bnz in L-,uolutiod bristi Hop (1966), whih
assssdth rlationship twn Drwinism and a varity of futur-orintd Rlations btwn volu
idologis-arist and matrialist historis of salvation, Nitzsh's futur- Attituds, morovr, dl n
isti dotrin of th Urmnsh, zrnd spulations on volution and th fu- fssionirl traditions, or lo
tur of humanity in th writings of th Hindu Sri Auroindo. Within th light of this raliztionw
.Wstrn
Christian traditin h flusdon th volutionizd shatologyof t,ll'tion and rligio tr
Teilhard d hardin' who onivd of vlution as impelling all lif toward soial, or sintifi world
an ..Omga Point.'' Th writings of Jams Mosh, Hnry Drummond, and l l s i n d i f r n I o e t i
Gorg Frdrik Wright also atn within th ar of this analysis. Indd, politil, onomi, mora
thr is muh to said for olzrtingattituds to volution with shto. th rltionship twnt
logial stans. Postmillnrians, witlr thi oust onfidn in tlr trans- u n d l s o f i d a s .A l s st r
forming powr of hristian iviliztion, hav routinly n favoraly episds might thrfore1
disposd toward volutionary prlgrssivism,whil prmillnnialists,who - attak on th lrgy in hi
liv that th world will ontinu to dgnratuntil th millnnium is ush. Prsbytrians to rad v<
rd in, typially find volution rpugnant.Thus ir is not surprising that th with tlr thrats of ilil
modrn foundr of ration sin,Gorg rady Pri, flt that blifs Prsytriilns thr wr
aout th gir-rning and n]of tim wr umililly onntd,and that h Za|and rsponssw s
uolutio d Rligio 65
lf natural ausation'' Was ntirly told th radrs of his 1923 Sie d Religil i Nlltshll tat it was
..uslssto xpt popl
: | o l o g i asly s t mi m p | i s O m - to liv in th prditions givn in th last hap-
lstrumnt'.(qutdin Livingson trsof th Bil, if thy do not livin th rord of th vnts dsridin
thsrsponss undoutdly ar, its first hapts'' (Pri 1923, 13).
'inglous f dat. Th ralization that th prinipls of Darwinism might applid to th
st of qustions that ongrgat ttul uolutio lf sriptur and to the merg of religion itslf has n
te ttur nd morlit of the a furthr point of issu.Th ida of soial volution, of ours' prdatd th
rdoptd on th issu of whthr puliation of Darwin's origi, and th suggstion that th txtual history of
hav optd for th viw that th th Bile had to intrprtdin th light of soity'sstdial dvlopmnts
:ration.Thus th atholi volu- was widly adoptd y ttual ritis in th nintnthntury. Indd, th
31 aount uolutio nd bol- priniplof volutionary hang was so dply ingraind in th doumntary
sophr ould possibly allow that hypothsismanating from Grmn highr ritiism that Darwin's invn-
r brut soul, muh lss from inor- tion ratd no stir whatsovr in that Old Tstamnt sholarly ommunity
:an only om into eistn y a nor among thos nthusiastiaout the puliation of sss nd Ruies.
43). othrs hav thought that th By th sam tokn, as w hv sn' th aliztion that volution ould
r arlir forars is ntirly om- pplid to th th institutional and th ritual history of Isral was rlvant
.ial rprussionsof Darwinism, to th way in whih Darwin's hallng was rad among nintnth-ntury
moral snsiilityhas n shapd Jws.Suh uss of volutionary thought forms, of ours, hav bn distur-
.William .Wright's
al. In suh a snario, as ing to many. Gorg Frdrik ltr wvring on Darwinism sprang
mad lar, thr is no rason to in good part from his onrns aout its impliations for th Bil's dou-
' hav any diffrntly from ani- mntary history. In Sotland th promulgatin of ilial ritiism y
; of osmi volution, hav sought, william Rortson Smith with its dploymnt of volutionary anthropology
g of volution to mak sns of spookdonsrvativalvinists so muh that Dwin's own thoy of spis
l t h o u g hd i f f r n tr l i g i o u s o m - hangsmd rathr tam in omparison (Livingston 2004). On again,
n th qustion' th issu of this attitudsto Darwinism wr mdiatd through what was takn to th
tslf. thory'srlvanfor roadr ultural affairs.
rinking, th idas of progress and,
's thologial nt-door nighor
: u s s i o nS. o m o f r h s s s o i a r i o n s Puttingvolution and Rligion in Thir Pla
d hristin Hop (1966), whih
;m and a varity of futur-orintd Rlations twn volution and Iigion hav nvr n straightforward.
is of salvation, Nitzsh's futur- Attituds, morovr' do not map natly onto dnominational ontours' on-
:ulations on volution and th fu- fessionaltraditions, or loations on th onsrvativ.liralsptrum.In th
.!7ithin light of this ralization w might justifialy paus to wondr whthr th trms
{indu Sri Auroindo. th
r th volutionizd shatology of ulltioand religio trak any transndntal ssnin th thologial,
lution as impelling all lif toward soial, or sintifi worlds. At th vry last th wid-rnging uss of ths
osh, Hnry Drummond, and llsin diffrnt loations and thir dploymnt in a host of disourss-
n th ar f this analysis. Indd, politial,onomi' moral' and soial-alrt us to th fat that in spaking of
:titudsto volution with shato. thrlationshipbtwnvolution and rligion w a handling rathr triky
:ir roust onfidn in th trans- bundlsof idas.A lss transndntaland mor loal approah to partiular
L' hav routinly n favoraly pisodsmight thrfor prov nlightning. For ampl' whras Tyndall's
m, whil prmillnnialists'who - attak on th lrgy in his Blfast addrss of 1874 mad it diffiult for loal
nratuntil th millnnium is ush- Prsytriansto read volution thory sympathtially, th proupation
:. Thus it is not surprising that th with th thrats of ilial ritiism at th sam tim in Sotland mant that
1rady Pri, flt that lifs Prsytriansthr wr lss risd aout iologial mattrs. In Nw
umiliallyonntd, and that h Zea|andsponsswr shapd y th politis of Maori.sttlr rlations, and
66 uoltion d Rligion
in th Amrian South raial qustions prdominatd,though with diffrnt Brundll'B. 2001. tholi
outoms in ah as. h way in whih Amrian atholis rspondd to lourl for th Ilistor
soial Darwinism in th yars round 1900 ws shapd y th soial nih Burkhardt,F. ., and S' Sm
D r t u i . 1 5v o l s . a m
thy oupid: Amrian Catholiism Was vry largly a woking-lass rli.
yn' G. V. 1998.voluti
gion of nwomrs struggling to maintain thir idntity in nw nvironmnt.
T. Ptrs,d.,Scic
In all ths instans th partiulars of gographi loation and soial spa .Wstviw
Prss.
had hug rol tl play in what ould said out volution and, just as D r w i n , . 1 8 5 9 .o t h
important, what ould hard aout it. 1888.e Life d
From th momnt Darwin put forward his thory of volution y natural urray.
|959. h rigio
sltion, th dat was ngagd on its signifian for a wid rang of
N4.Pkham,d. Phil
rligious issus.Th fat that ontmporary volutilnaryiology, as a om' Dsmond,A.' andJ. R. ot
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uoltio d Reliio 69
lutionism: to guid thm and tlrLrswill hv no rason to hav proprly to tlrir fllow
itizns.Soity wuld suffer gratly, all usof ..lif''in volution. A
spt ommon rationist laim rlatd to havior is that if hildrn ar taught that
thy ar animals, thy will hav lik animals rathr than raturs mad in
th imag of God. ln ddition to ths pratial rasons for rjting volu-
tion, thr ar thologial rsons as wll. Gnsis is sn as th foundation
fr hristian lifs:if Adar and v Wr not ral flsh-nd-loodindividu-
als' spiaily rtd y God, whos sin of disodinausd thrn to
astout frm th Gzrrdenof dn, thn lrurnrrkindis not inhrntly sinful. If
War not inhrntly sinful aus of th Fall, thn Christ's sarifi rrd rs.
urrtion' th pivotal vnts for hristinity, wr unnssary.In th opinion
ring its sond ntury. Bgin- of onsrvativ hristians, th ook of Gnsis thus must e intrprtd -
rtinuing today, with no sign of atly as writtn, nd volution is rjtd baltsit is inompatil with mst
r tahingof volution to hil- formsof iblial litrlisrn.Ndlssto say' this is not th only Christian viw;
s waxd and wand in strngth atholisand mainstramProtstants(pisopalians,mmrs of th Unitd
n tht is taught in th lssroom hulrof hrist, most Mthodists, most Prsytrians,mmrsof th van.
trovrsywitlr long histry and glialLuthrn hr"rrhin Amria, arrdothrs)ar thistivolutionistswho
s toward rligiorr'duation, and livthat Gd rtdthrough volutiorr.
Th hyday of this fist priod was approximatly 1919 to 1927, when
attmptswr mad in svralstat lgislatursto an th tahing of volution.
Th bst-known prussionof thsfforts WaSth famous 1925 Sops trial.
Tnnsstahr John Sops was trid for raking a stat law that prohi-
itdth tahing of volr"rtionin th puli shools. Th trial of th ntury, so
alldausof th hithrto-unxpindoast-to-ollstradio and print nl-
on must undrstnd th history dia puliity thzrtit ngndrd,rsultd in th onvitin of Sops (ovrtund
l into thr tim prids (Sott on a thniality y th TnnssSuprm ut). Dspit th puli prp-
' volution from th puli shool tion of th trial as a dfat for Fundamntalisnr (as nouragd y th ply and
I th twntith ntury, volution ft|mInhrit th Wid), in its aftrmath am th gradual diminishmnt of
lks and thry into th urrlu- volutionin ttooks and thforin th urriulum of puli shools.
rnd in uran population growth Indd,aording to on ommntator, volrttion was fftivly removd
;m in th Unitd Stats (Larson from th urriulum y th 190s, and it did not rturn for svral dads
y an uran phnomnon' and th (Lason2003). volution rturnd to th lassoonr only in th lat 1950s
'nging onorrri lnditions that and arly l960s, whn pofssiorr:rIsintists wrking with th fdrally
farms into itis and y forign funddBiologil Sins urriulum Study (BSS) wrot trooks that
[' mor studnts wr attnding mattr.of-fatly irrluddvolution. ommrial ttooks susquntlym-
.ntithntury thn vr for. ulatdth ontnt <lfth popular BSS ooks, and y th lat 1960s and
;hool iology for ovr a dad' it ear||970s evolution was usually, though not amply, inludd in most of th
n Wrxposd to it. on that ttooksing sold in th Unitd Stats.Th rturn of volution to th ur-
;ity toward volution promptd riulumstimulatdtlr sond wav of antivolutionism in th Unitd Stats:
lum. thpiod of rctilt sitt..
72 mric ntieuolutkism
during th 1925 trial no laims vanglism ministris of Knt Hovind (until his rnt inarration on fd-
al ration. ral hargs of failing to pay payroll taxs [..Knt ovind sntndto 10
ion sin.Th plaintiffs argud ears',207|, lfaltr Brown, John aKay, and arl Baugh, among othrs.
tion sinwas not sin,and Spakrs from ths ministris and othrs ar onstantly rissrossing th
,ahingit in puli shool sin ountry' spking at hurhs' ollgs, and vn publi shools-anywhr
1 th stat promoting rligion, in thy an find an audin to listn to th antivolution gospl. Innumral
t'switnsss(th ration sid) individual wsits promot ration sin, and tlvanglists frquntly
dfnswas marrassdwhn invok lassi ration sin argumnts to attak volution. Th IR has a
lar vidnthat th Arkansas musumthat laims thousands of visitors annually, and in th spring of 207
r the purpos of singling out on AIG pnd a 90,000-squar-footpuli musum nar its headquartrs in
.!7ithin
jtsausit offndd th r- northrn Kntuky. th first svn months of its opning, AIG had al-
iv was struk down. So firm was rady dd its ptd annual attndan' drawing ovr 300,000 visi-
l to a highr out. tors in its first yar (..ration usum Unxptd Suss'' 2008). A
/asa vry similar ill passd in rationist usum of arth History, whih will fous on Flood gology, is
didd on summary judgmnt plannd to opn nr th popular Branson, Missouri, tourist dstination.
oth sids,without a trial) that th Thr is, thfor' plnty of opportunity for th Amrian puli to x.
lishmntlaus. Th stat also posd to ation sin through th movmnt's various puli outrah
;from th Suprm ourt' whih ativitis' puliations, lasss' and rvivals. As two spialists in volution
prm ourt agrd 7-2 (Justis duation wryly rmark' ..any studnts hav hd ampl formal and in-
:nting)with th lowr ourts that formal dutional oppotunitis to misundrstand volution'' (Altrs and
s position, and thus its tahing Nlson 2002\.
Th Suprmourt hos not to
:was sintifi, living that its
NoREAloNIsM
:omth lassroom. Th dision
Norationism rfrs to th post- drds rpakaging of ration sin
larglyto avoid its lgal prolms (Sott |997). It inluds proposing allgd
y advaningth rligious - ..altrnativ
sintifi planations'' to volution, suh as Intllignt Dsign
rankind.Th lgislativhistory
(ID), and/or proposing to ountr volution not with an altrnativ sin
n,'' as ontmplatd y th
but y allgd evidn against volution. Ths approahs wr nour-
:ahing.Th At's primary pur-
agd y th wording of te drds dision itslf, ut thy pristd in
:nurriulum to provid pr-
ationsin.
ious dotrin that rjts th
hus,th At is dsignd ithr ,,Itrntiue
sintifi tbories,': IntIligntDsign
: that modisa partiular r-
\n the urds dision Justi \/illiam Brnnan Wrot that rationism
lf a sintifi thory disfavord
ould not advoatd as an ltrnativ to volution aus of its rligious
r At violatsth First Amnd-
ontnt)ut that ..tahing a varity of sintifi thoris about th origins of
humankind to shoolhildrn might b validly don with th lar sular in-
s am to a halt at th foot of th tnt of nhaning th fftivnssof sin instrution.'' H also Wrot that
itslf did nt withr; instad' it tahrsould tah ..all sintifi thoris aout th origins of humankind''
sign of diminishing during th (drds u. gillrd, 59_594). In fat' thr ar no sitc altrnativ
,86ws introdud into (although thoristo volution, ut Brnnan's ommnt invitd rationists to propos
..puli
;islatur,proposing that thir xistne and providd a lgal foundation for arguing to prsnt thm in
l tratmntto th thory f sin- publi shols. rtion sinWas th original sintifiltrnativto vo-
:r.''lassi ration sin an lution; with its failr to onvin th sintifi ommunity or th ourts, it
I C R a n d i t s s i s t ro g n i z t i o n , rquird rpakaging. Th most important rpakaging of ration sin
t h i n d p n d n t r t i O n s i n into an ..ltrnativ sintifi thory'' is Intllignt Dsign.
376 meric Antieuolutionism
Te drds dision took a long tim to work its way through th ourts, ster was publishd,
ut vn tr th Suprm Court's dision ws issud in th summr of indst: origin of Lif l
7987, man antivolutionists had sn that th antivolution movemnt would ontnt in DNA'' in 198
hav to go yond ation sin and spially yond th young-arth nt sponsorship, follow
rationism (Y) of Hnry . Morris and th IR if it wr to xpand and Philosophial frn j
sussful.Thy sought form of antivolutionismtht oth would hav of Culrur''1 1996,.,er
mor sholarly rsptaility and would avoid th First Amndmnt Pro- Sientifi ntrprise.''
.!7ht
lms of ration sin. was Soon to e lld IntlligntDsign was Som of th fas at tl
nouragd y Jon Bull' ministr who dirtd (nd still dirts) a Txas who follows tlr antiv
nonprofit organization alld th Foundtion for T.houghtand this (FT). ationists' suh as Paul N
Bull nouragdfllow antivolutioniststo downplay ilial litralism and B i l - S i n A s s o i a t i o
Gnsis imagry and to invok a vagur form of spial ration to plain latr orgniztion g
highly omplx natural phnomna. In this apprlah God th Dsignr ationist studnts at th
would not rfrdto, and th inaility of volution to plain omplity morphd into an ID ad'
would strssd.1Bull notlragdth puliation of The ster lf Lif's sarh Ntwork. ost
origi (Thaton et a|' 1984), whih was writtr thr sintists assoi- a t d w i t h n o l d - a r t hv i
atd with FT. Although th ook onntrats only on allging wak- and dirts th llasons
nsss in thn-urnt thoris of th origin of lif, srupulr"rslyrlgating suh as mathmatiian/1
disussion of dsign to an pilogu' it is now rgrdd s a founding doumnt C. lvlyr, and thologra
of ID. fw of whih inludd
str ontndd that th origin of lif was an ampl of a lass of ph- movmnt to dvlop its
nomna that wr too omplx fo plantion hrough nurl auss.his strtgy for th otsid r
was atgorially so and was not th rsult of inor-npltknowldg. Th ori- vn in th ginning,
gin of lif was not only a diffiult sintifi prolm, it was an unsolval on lassroom (Sott and M.
if on rstritd onslf only to natural alrss. Narr-rralauss Wr qui1td ouragd th writing an<
with han, and suh omplity as that hiitd in th strutur of th first t o h i g h - s h o o l i o I o g yt
ll ould not xplaind y th random joining of llular struturs. Suh tion was that Bull soug
diffiultis ould ovrom' Thaton nd his oauthors laimd, on[y y tr would mak th blt
inluding th tions of n intlligntagnt in sintifiplanation. 1 9 8 9 ) .U n d t h a u s p i
str prsntdth possiility tht th agnt was matrial (suh as in. oauthord of Pds r
tllignt lins), ut it optd for a transndntagnt..yond th osmos'' was long on ritiisms o
(Thaxton et a|. 1984,200)-God. In th pilogu of th ook th athors pla it. FT attmptdt
spok frly of spial ration and th nd to xpand sin to inlud ook was widly rjt
suh ausation as a mans of otaining omplt viw of truth. |n 1997 a law profs
str had littl impat on th sholarly world. An Institut for Sintifi Phillip Johnson, publish
Information (ISI)itation ount notS only a fw rviws, largly in philos- Although h ster <
phy and rligion journals, and a total f only 28 itations twn 1984 and sarly any noti in thr
2008. In omparison' anothr popular ilount of th origin of lif, A. G. noti fom th gnral p
ains-Smitlt,sSue lus tl th origi of Lif, pulishd in 1985, had 95 iologist, lit in zr po;
itations during this sam tim priod. Although uninfluntialin th wold of baus of th novlty o1
..om-
sin' str is rvrd in rationist irls, whr adjtivslik ing an antivolution
plling'' and ..impotant'' ar irrvarialyusd to dsriit. only with privatorgn
dmi world; Johnson's i
SABLISFIING ID. on of th authors of str, historian and hmist guably put ID on th ma
harls Thaton, promotd th nw ID nrovmnt y organizing onfrns standard rationist argu
Th first ws hld in th sam yr that
of fllow hristin ntivolr.rtionists. rros.
meric ntieuolutionism 77
work its way through th ourts' str was pulish d, 1'984, and was titld ..Going yond th Naturalisti
on Was issud in th summr of Mindst: origin of Lif Studis.'' It was followd y ..Sours of Information
l antivolutionmovment would ontent in DNA'' in 1988. Othr grandly namd ID onfrns,undr diffr-
..Darwinism:
pially yond th young.arth nt sponsorship, followd 1992 rought Sintifi Infrn or
th IR if it wr to xpand and PhilosophialPrfrn?,';1995,..ThDath of Matrialism and th Rnwal
lutionism tht oth would hav of ultur''11996,,,ere ration''; and7997,..Naturalism, Thism, and th
.oid th First Amndmnt pro- Sintifi ntrpris.''
l l d l n t l l i g n D s i g n w a s Som of th fas at ths onfrns would hav n familiar to anyon
irtd(and still dirts) a Txas who follows th antivolution movmnt' inluding a fw young-arth r-
'n for Thought and this (FT). ationists'suh as Paul Nlson, who had workd with th innapolis-asd
downplay ilial litralism and Bil.SinAssoiation and latr with Studnts for Origin Rsarh. h
rm of spial rtion to plain lattr organization gan as a group of prdominantly young-arth r-
ris approah God th Dsignr ationist studnts at th Univrsity of alifornia at Snta Brara and latr
f volution to xplain omplxity morphd into an ID advoay organization with th odd nam Ass R-
liation of The str of Lif's sarh Ntwork. Most of th onfrn partiipants, howvr' wr assoi-
v r i t t n y r h r s i n t i s t s s s o i - atd with an old-arth viw, suh as Hugh Ross, an astronomr who foundd
]ntratsonly on allging wak_ and dirts th Rasons to Bliv ministry, and youngr antivoluionists,
n of lif, srupulously rlgating suh as mthmatiian/philosophr l/illiam Dmski' philosophr Stphen
rgarddas founding doumnt . Myr, and thologian and iologist Jonathan \/lls.Ths onfrns, a
fw of whih inludd dfndrs of volution' wr a mans for th young
Wasan xampl of a lass of phe. movmnt to dvlop its idas and, mor important, to hon its mssag and
tion through natural auss. This stratgyfr th outside world.
lf inomplt knowldg. Th ori vn in th ginning, ID advoats aimd thir fforts at th puli shool
lrolm,it was an unsolval on lassroom(Sott and atzk 2007). Bginning in th arly 1980s Bull n-
ss.Natural arrsswr quatd ouragd th writing and markting of a ttook intndd as a supplmnt
hiitd in th strutur of th first to hrgh-shool biology txtooks. Part of th rason for th dlay in pulia-
oining of llular struturs. Suh tion was that Bull sought a slar rathr than a rligious pulishr; th lat-
d his oauthors laimd, only by tr would make th ook far lss salal in th puli shool markt (Sott
in a sintifiplanation. 1989).Undr th auspisof FT, in 1'989Pival Davis and Dan Knyon
agntWas matrial (suh as in- oauthord f Pds d Pople, a rif ook that in typial ID fashion
..yondth osmos'' was long on ritiisms of volution and short on spifis of what might r-
dnt agnt
:piloguof th ook th authors pla it. FT attmptd to promot it to shool oards for adoption, ut th
d to pand sin to inlud ook was widly rjtdy oth tahrsand sintists.
lmpltviw of truth. In 1991 a law profssor from th Univrsity of aliforni at Brkly'
r world. An lnstitut for Sintifi hillip Johnson, pulishd th first widly rd ID ook, Dri o Tril.
a fw rviws,largly in philoso- Although The ster of Lif's rigi and f Pds d People attratd
ly 28 itations twn 1984 and sarlyany noti in th sintifi ommunity, Johnson's ook rivd wid
:ount of th origin of lif, A. G. noti from th gnral prss and Was vn rviwd y notd volutionary
f Lif, ftlrishd in 1985, had 95 iologist,alit in a popular sin journal (Gould 1992). This was likly
ough uninfluntialin th world of auseof th novlty of a pofssor from a major sular univrsity author-
..om-
rls.whr aditivs lik ing an antivolution ook. Antivolutionism had prviously n assoiatd
d to dsriit. only with privat organizations suh as IR that wr marginal to th aa-
dmiworld; Johnson's involvmnt ratd a flurry of mdia intrst and ar-
| str, historian and hmist gualyput ID on th map. Th ook itslf onsistd of a rworking of many
,vmenty organizing onfrns standardrationist argumnts and was roundly ritiizd y sintists for its
rst Was hld in th sam yar that rrors.
78 l ric tiuo luti o i sl
Th nxt major ID puliaion was Drxai's Blk Bo y Lhigh Univ- himslf as living irr
sity biohmist Mihl Behe (1996), pulishd y th Fr Prss. Dri,s prominnt lD thorists]
Blk Bo marks a high point in ID's effort to ast itslf as a sin. Th tion'' (Dmski1995,3
prmis of this ook was similar to that of ster: som phnomna in n- votlts' inluding in hi
tur a simply too ompl to plaind through ntural auss.Spili- (Bh2007).
ally, Bh pointd to moleular struturs suh rsth atrial flagllunr As PhillipJohnsonha
and prossssuh as th lood-lotting asad s ing too orplx ro rO thir hif onrrr
plainal through inrmntal natural sltion. Bh,s ook rivd tlrrough natural pross
som rviws in th sientifi litratur-whil Johnson's book did not- intrvntion is rquird
prhaps aus hr again Was th novlty of a professor from a seular uni- phnomna ar dsign
vrsity pulishing an ntivolutionism ook, this tim from a mainstram through natural pross
press' Drlui's Blck Box also rivd mor attntion from sholars than is tlrat it is possibl ttl
Johnson,s Dri o Tril us it attmptd to provid a sintifi argu- ompl phnomna of
mnt. Howvr' th rviws y sintists wr lmost uniformly ngtiv iologial phnomna,.
(oyn |9 9 Q D orit 1 997; Blaks tone 1997i vlir-Smit 19971 Thornhill y ration sin.rhr
and Ussry 2000). poaility and informz
B 7998, whn philosophr and mathmatiian \illiam Dmski pu- Dmski has ontn
lishd his first ook, Tbe Desig lferc, th asi strutur of ID had (..spifid''in advan
n stalishd.In be Desig Infere and othr puliations Dmski things thar hv hnil
prsnted a proaility-asd xplnation fo th furrdamntalida of ID: ( S I ) , i t i s l a i m d 'i d n
that som natural phnomna ar too ompl to plaind through nat- quir tl-lation of int
ural auss and thrfor rqui ..intllign''for thir xplanation. Invok- a n d o a u h o r s ' o l l t l l t
ing an ..xplanator hlter', (p. 7) of inrsing improbability, Dmski lif was too omplii1t
laimd to al to remov natural auss and han as potntial -pl- to idntify a lass f ph
nations of omplx phnomna, laving dsign as th rsidual planation. natural' that is, sintiI
Dmbski has had a diffiult tim onvining fllow philosophrs and proa- Mihal Bh's ida
ility thorists of his viw and has n quit unsussfulat prsuading i- is similarly usd to argu
ologists that his onpt of ompl speed informtil has any utiliry for Bo (7996)that rtin
undrstanding iologial phnomn (Fitlson et aL. |999; Shanks 2004; e p l a i n a l y n a t u r a ls
Wilkins nd lsrry 2001; Rosnhous2003; Pnnok 1,999,20\ lsbrry th tril flagllurn,-
and Shllit2003). thir many omponnts
ursor of an irrduil
TH ONTN oF INTLLIGNT DSIGN. In ontnt' Intllignt Dsign dif- natural sltion-thou1
frs fom ration sin primarily in its rlutan to mak spifi fat ats on On omponnt
.s(hras
laims' ration sin maks laims aout th ag of the eath a harm ralt.Bh i
(only a fw thousand yars old) and th origin of gographi faturs (ata- for irh addition of
strophi ation' suh as Grand anyon's ing formd y Noh's Flood), litlts hv tl b ssn
ID is mut aout th history of th univrse ausID intnds to a ig natural sltion arrn
tnt that will hous all antivolutionists'and ausinsid this ig tnt r misundrstandingof tht
popl with diddly inomptil idas (Sott 2001). ost of th ID advo- out a numr of ways t
ats ar old.arth rationists, ut thr ar young-arth rationists who pl struturs' vn s(
ar ativ supporters of ID. Similrly, as do th young-arthrationists,ID 1997; rr 1996).
advoatsallow for volution within ratdkinds, ut thr is no onsnSus
in th litratur of ithr an1pon what a ..kind'' is, th limits of gntivri- ID As SHOLARSHIP. B
ability tht rstrit volution outsid thsratd kinds, or what dtrmins ons' th is tlr isk tl
ths limits. Mihl Bh, a prominnt spoksprsonfor ID, on dsrid stutur that was disor
m r i c nt i euo lti oisnt 79
''sBlk Bo Lhiglr Univr- himslf as living in God-guidd volution (Bh 2000)' although othr
;hdy th Fr Press. Dri's prorinntID tlroistshav dlrd that ..ID is no find of thisti r,olu-
.t to ast itslf as a sin. Th tion'' (Dmski 1995, 3). In rany ways, Bhe stands apart from thr ID ad-
\ 4 s t r s:o m p h n o m n ai n n a - voats,inluding in his aptan of ommon anstry for living things
l through natural auss. Spifi- ( B h 2 0 0 7 ) .
i suh as th atrial f-lagllum As Philiip Jolrrrsonhas rommndd,ID advo:rtstry to lirrritthmslvs
iad as bing too orpl to to thir hif onrn, whilr is whth natur n xplaind solly
sltion.Bh's ook rivd through rraturalprosssor whthr intllignt-in prati, suprnatual-
whil Johnson's ook did not- intrvntionis rquird. This ida undrlis ID's or laim: som natural
of a profssorfrom a sular uni. phnomnaar dsignd y n intllignrathr tlrn havirrg om aout
lk, this tinrfrom a minstm throughnaturl prosss.Tlr sondary,though sintifiallyritial' laim
or attntin from sholars than is that it is possil to distinguish intlligntly dsignd phnomna from
rptd to provid a sintifi argu. ompl phnomna of ntur:rl origin. Typially, ut rot alwys, ths ar
w r l m r s tu n i f o r m l y n g a t i v iologialphnonrna,vr oftn th sm phnornsltdfor srurirry
,; avalir-Smit1997; Thornhill y rtion sine.2Th mthodology prposd involvs a onpt asd on
proailityand information thory dvlopd y Dmski.
mtiian William Dmski pu- Dmski hs ()ntnddtht low-proaility vnts of a partiular type
;e, th asi Strutur of ID had (..spifid'' in advan y sid information) ar assignalto th atgory of
and othr publiations Dmbski things that hav n intlligntly dsignd. omplx spifid information
' for th furrdamntalida of ID: (SI)'it is laimd, idntifisa nwly disovrdlass of phnomna that r.
pl to plaind though nat- quir th ation of intllignfor thir origins. Harking ak to Thton
n'' for thir xplanation. Invok- and oauthlrs'ontntioninTh strlt of Life,s origi that tl.rorigin of
rasing improaility, Dmski lifws too ompliatd to plaind y natural us' Dmski uss SI
;sand halr as potntial xpla- to idntify a lass of phnomna not only unxplaind ut unxplainal y
sign as th rsidual xplanation. naturl'that is, sintifi,prosss.
rg fllow philosophrs and proa. ihal Bh's ida of irrdui1orrrplity(I) of biologizrlstuturs
.ritunsussfulat prsuading i. is similarly usd to argu against volution. Bh limd in Drin's Blk
1dinformtio,asany utility for Bo (1996) that tain iohrnialprosssor llular strutursWer not
tlson t a|. 1'999; Shanks 24; xplinly ntr.rrlsltion.H ontn]dthat som strutllrs' sr-ras
)03;Pnnok 7999,2001;' lsrry th batrilflagllum,rquir th assrnlyin final Opratingordr of all of
thirmany omponnts for thy an b funtional. By dfinition' any Pr-
ursorof an irrduily ompl strutur is nonfuntional. Bh's viw of
[. In ontnt, IntlligrrtDsign dif- nturlsltiln-though not tht of volutionary iologists-is tht it or-
:s rlutan to mak spifi fat atson on omponnt of a strutrrrat tim, rathr lik adding harms to
laims about th ag of th arth a hamralt.Bh argusthat if natural slti<rn rquirs sltivvalu
ligin of gographi fturs (ata. for ah addition of ah omponnt of an I strutur' ut all t mpo_
's
ing formd y Nah's Flood), nntshv to ssrldtor thr is ny sltivvaiu, inrmntal
:rsbaus ID intnds to a big natural sltiorrannot xplain irrduily omplx struturs. This is a
rnd ausinsid this big tnt a misundrstanding of th pross lf natural sltion,and ritis hav pointd
(Sott2001). Most of th ID advo- out a numr of ways tht natural sltion ould oprat to produ om.
] ar young-arth rationists who pl strutrs'vn so-lld I ons (Thornhill and Ussrv 2000; Dorit
do th yong-arth rationists' ID 1997..rr 1996\.
td kinds, ut thr is no onsnsus
..kind'' is, th limits of gnti vari- IDAs SHOLARSHIP. BausID onfussr'lnsolvdprolmswith unsolval
g ratdkinds, or what dtrmins ons'thr is t[rrisk that ID might disprovd l,an irrduily ompl
poksprsonfor ID, on dsribd strutr
that was disovrdto hav a plar"rsil
naturl planatiorr.Howvr,
80 mri tiultilism
this dos not sm to an atual diffiulty, for ID proponnts an always al- ID artils in th si
low that thy w wrong aout th irrduil omplity of tht strutu doing sin y ID's
and om up with anothr. Ironially, thy sm to ignor strutus that ar not produ (som)n
irrduily omplx, suh s th mammalian middl ar or th avian wrist ural auss' th tho
ompl, ut whih in fat hav a wll-undrstood volutionary history (Gish- .!ha
atual rsarh.
tik 2004). From a sintifi standpoint, of ours, th inaility t rfut I y look lik othr than :
disproving laimd xampls is not a strngth ut a waknss. Sin pro- show, that natural pr
grsssy liminating planations that ar disprovd' not y proving plana- quat to aount for
tions in som solut sns. snario prsntd
Intllignt Dsign proponnts hav produd a larg numr of ooks thory hind ID dis<
sin th apparan of Th str of Life's rigi' th ulk of thm sin thr should no st
te 1996 puliation of Bh's Drlui's Blh Bo and Dmski's 1998 tur. Dmski hims
Th Desig Ifr. ost of ths ooks, howvr, must lassifid as th lak of progrssil
rligious or philosophial in natur rathr than sintifi;it is rvaling that ..d
mploying a vagu
most a pulishd y starian pulishs suh as IntrVarsity Prss, Bakr of thnologial inn<
Book Hous, nd Brazos Prss.his is urious' sin ID's laim to al al systms (Dmsk
to idntify intlligntly dsignd biologicl strutures would imply that its al phnomna will
adhrnts intnd lD to answr qstions that ar ing invstigatdin th will in som way sul
iologial sins.Th pla to do this would in sintifijournals, sp- out that iologial pI
ially in th filds of llular and molular iology, ut hr ID has n fatul.
asnt. vn Bh, an aadmi sintist t a sular univrsity (Lhigh) Although the sin
who has pulishd in mainstram sintifi journals on sintifi topis un- ration sin'thr
rltd to ID, dos not appar to pulishing or attmpting to pulish r- nals, primrily au
sarh that maks a positiv s for ID. Th most prominnt [D dvoats writtn for th gnr
appr to prfr to publish in popular ooks rathr than sintifi journals. tal xption. As a
Dmski has ommntd: ..I'v iust gottn kind of las6 aout sumitting Bh ar ook rviv
things to journals whr you oftn wait two yrs to gt things into print. philosophy or math
And I find I an atually gt th turnaround faster y writing a ook nd uss ID (.$ilkinsanr
gtting th ideas prssdth.y ooks sell wll. I gt a royalty. And th 2003)' Thr is a gro
matrial gts rad mor'' (MMurtri 2001' 8). ritiqu ID on sint
Normally, nw sintifi idas ar first psntd t aadmi onfrns Forrst and Gross 20
and, aftr profssional fdak and som gnral disussion, ar rfindto Young and dis 200,
th point whr artils ar sumittd to sintifi journals, aftr whih thy ussions among ID a
may or may not join th sintifi onsnsus.Aording to an artil in th vr (.g., www.pn
hroicl of Highr dutio, whn askd why h is not uilding his and Shallitt200;Wt
irrduil-lmplityida and th ID model at profssionalmtingsof ll For all th laims t
iologists and iohmists,..Mr. Bh rsponds that h prfrsothr vnus. ntrpris, vn a bri(
.I just don't
think that larg sintifi mtings ar fftiv forums fo pr- no on is applying tl
snting ths idas,' h says'' (MMurtri 2001, 8). stand or plain io
Again parallling ration sin,ID proponnts ontnd that thir viws 2004). Th urdn o
ar systmatially kpt out of sintifi journals austhy rjt th st- thir position is sin
tus quo. Yet unorthodoy is not routinly anishd from puliation (as th tionary biology app
historis of th rption of ndosyrnisisand puntutd quiliria wit- insights and undrst
nss),although unusual laims rquir mor than th usual amount f sup- nw fossil rmains r
port and tak long to stalish.A mor plausil rson for th darth o{ dvlopmntal biolog
meri ntieuolutioism 81
i writtn' ..lntlligntdsign is minimalist dfinition for virtually all sintists.But this miniml dfinition
lt invstigarsth ffts of in- nonthlssontains hratrististhat ar opposd by ID advoats.
t hallngsDrwinism nd its IntlligntDsign rationistslivthat th rstrition of sien to nat-
.ding divin ation'' (Dmski ural auss,known aS nthodologi1 naturalism, lads popl to philosoph-
logrmof ID has n unsu- ial naturalism.Philosophial naturlism is, of ous' th nmy ausit is
D movmnt is dirtd toward opposd to Christian thism' If a powrful ultural ntrpris lik sin
)mbski'ssond laus:th f- ould roadndto inlud God's intrvntion' and if mthodologial nat-
lviw,trrntmountto athisn-l- uralism ould aandond, thn philosophial naturalism would los what
th inadquay <lfits sintifi is privdas its strongstsupport. Th stalishmntof ..thistisin''
lsr th ID rovrntis in fat (orland 1994; Scott 1998), whih prmits th invotion of th hand of
'n and its onflit with mtrial. Gd (or ..intllign'')whn natural xplanations ar indquat for th
only nratrial(rnattrand n- jo, is a ky goal of ID ationism.
suprnaturalrality. Thisti sin is asd on th onpt of a dihotory twn origi sci-
, I D n l o v l n t t t a k s s i n e and opertion sitl, th lattr ing ID proponnts'trm for th ..nor-
nough,is not primarily through mal''' mthodologially matrialisti sin usd in th v:rst mjority of
and vn sttistialargtrmnts' sintifiplanations. origin sin is dfind s th sin usd to plain
'y philsophrsand proaility singular, unrpatabl vnts (th origin of lif, for xample)' hypothss
lts r2tsonrquis a short di- about whih supposdly ar untstal and thus r.rnfalsifiabl.Thrfore' at-
tidby sintiststody. triutionof ausality to God is aptalin origin sin,though not in op_
rgu aout tl-rdfinition of si- ration sin.a
.distinguishdfrom othr ways volution, thn, is a stalking hors for an attak upon sin, whih -
I p o r t n t o t h a t t | a l n t p i s aus of its matrialisti as (ID rationists do not distinguish twn
l tnd not tl think muh aout mthodologialand philosophil naturalism in any pratial sns)is sn
ivly asy to gt sintists (arrd as a thrat ro hristiarr thisnr. Th ID advoat who nraks ths links most
'on a oupl of basi lmnts larly is on of th most prolifi ID authors, Phillip Johnson, whos 1991
n r s d i d n o r [ u l l y d m a r a t s i - Drluilt o ril was th first ID puliation to attrat wid puli noti.
ffortto undrstandth natural ..wdg'''
Johnson promots a stratgyhe alls th whrin volution is at-
nrgy and thir intration. It takd as a mans to hallng naturalism, thus paving th way for th
hnomnonof religion: th pur- rstalishmntof histin snsiiiity in Amrian ultu and the hg-
y dfinition not limitd to th mony of thisti sin (Johnson 1997i Forrest and Gross 2004). H has
imitd ndvor:it is limitd to also n instrumntai in promoting a puii Prsnttionof ID that voids
ommitmntto any fatual laims, suh as a youn8 or old ag of th arth' as
; of sin is tsting plana- a way f avoiding the pitfalls of ration sin.Instad,as did th athos
t i o n a s d t r l v n i n s p i r a t i o n , of he ster of Life's rigi and f Pds nd Pople for him,
John-
not a sintifi planation, son onntratson th supposd inadquay of natural auss (speially
l that wy. only ftr onfirma. nturalsltion)to plain th origin arrd divrsity of living things:
:ntifixplanation. Th famous
/r th irular strutur of n- So th qustion is: ..How to win?'' hat's whn I bgan to dvlop what
til illustrtsthis wll: it took you now s full-fldgdin th ..wdge'' stratgy:..Stik with th most
t natural world-to stalish important thing''-th mhanism and th uilding up of information.
with th drm. It is also likly Gt th Bil nd th Book of Gnsis out of th date ausyou do
ponntof tsting is holding r- not Want to rais th so-alld Bil-sindihotomy. Phrs th argu.
th fft of otlrrs. of ous. mnt in sh a way that you an gt it hrd in th sular aadmy and
Ld do dat othr faturs that in a way that tnds to unify th rligious dissntrs.That mans onn-
vrious vrsions of flsifiation trating on, ..Do you nd rator to do th rating' or an natur do
.sntlhr would suffi as a
it on its own?'' and rfusing to gt sidtrakd onto othr issus,whih
84 mricn Antiuolutionisnl
popl ar always trying to do. Thy'll ask, ..W.hat do you think of of th univrs at one til
Noah's flood?,' or somthing lik that. Nvr it on suh qustions - ar ford to support tht
aus thy'll lead you into a rrklsswastland and you'll nv gt out idn gainst volutin
of it. (Kushinr 2000, 40) a frqurrtlyrworkd lil
suh as gps in th foss
Baus it dos not prsnt an altrnativemodl to volution, and eus law of rhrmodynamis
of its rlutan to mk fat laims, th ontnt of ID thus dvolvs to th imossiility of dvlop
ritration of argumntslong ago raisd y th young-arthrationists:gaps dom variation and natu
in th fossil rord, th inadquay of natural seltionto produ volution- living organisms'' (IR
..prolms'' of th amrian
ary hang, th plosion, and s on. As is th i d n a g i n s t v o l u t i o n
..<rntrivd
s with Y, ID appars to rting a dualism''' to quot who ar puzzled that un
L u. rkss, in whih if volution an shown to untru, [D is apparan of amrian
thus stblishd. sltiv mhanisms i
thoy o rsons to rj
uidence gist euolution Tahing th ..vidn
Aftr th deurds dision struk down qual-tinr lws, Wndll Bird, who mnt laus prolm gl
had argud Louisiana's position fore th Suprm ourt, gamly advisd stratgy is thrfor fav
his fllow rationists that ration sin might still lgally taught if it who ar oprating in th(
wr taught with a sular purpos (IR 1987). His prsptiv was not for xampl, has om tt
shard y judgs in distrit and appals ourts who intrpretd d,rdsas ID ut rathr to tah th
not distinguishing twn rtion sin and rationtsl:.drds as
n itd y fdral distrit nd appals ourts in disionsthat hav struk
down attmpts by tahrs in lllinois and alifornia to bring ration sin Th Futur of Antit
into th lassroom.
A dissent to do,rds,writtn y Justi Salia and signd y Justi Rhn- Sott and Branh hav t
quist, suggstdan altrnat approah that was quit ompatil with xtant th priods of Amri
..Th popl of Louisiana, inluding thos who ar hist- ontrovrsy today as thr
ration sin:
ian fundamntalists,ar quit ntitld' as a sulr mttr' to hav whatvr 282). t any givn tim
sintifi vidn thr may against vlutin prsntdin thir shools, o f t h r a t i o n i s tp s i t i o
iust as r. Sops ws ntitled to prsnt whatvr sintifi evidnth 1. volution is wak o
was for it,, (dturds v' guillrd,64). Th ICR sizd upon this immdi- aandoning in inr
atly aftr te drds dision was pulishd. hoing Salia, it nour- 2. volution is inhr
..In th mantim' ..
agd tahrs to tah th vidn against volution: 3. It is only fair to
shool boards and tahs should strongly nouragd at last to strss o r l n t l I i g n tD s i g
th sintifi vidns and argumnts against voltion in thir lasss (not good pdagogy to ..
iust ar8umnts agains som proPosd volutionay mlranism,ut against ..waknsss''or ..
volution pr s), vn if thy don't wish to rogniz ths ils vidnsand did.''
argumnts for ration (not nssarilyas arglllnts for a partiular dat of
ation' ut for ration pr s)'' (IR 1987). Ths thr thms w
Th link twnation sinand ..vidnagainst volution'' is asy trovrsy in th futu. I
to find. Proponnts of rtion sin ontnd that thr ar only two possi- studis of antivolutiln
ilitis: rationism' onsisting of thir spifi vrsion of hristian spial stongsr motivation for
ration, and volution, whih also inluds all th otlrr origin storis from 1980s nd arly |990s,
othr rligions and ultus.Thrfor, to thm it is logial to infr that vi- rtionism in favor of t
dn gainst volution is vidnfor rtionism. Pratiill onsidtions ral puIi and, spi
ar aiso involvd: thr r no rdil data that suPport th spial ation Both ration sin
Amri Atieuolutionism 85
sk, "'S7hat do you think of of th univrs at on tim, in its prsnt form, so ration sin advoats
ar ford to support thir viw with a ngativ argmnt' that of allgd v-
r it on suh qustions -
land and yu'll nvr gt out idngainst volution. Th ontnt of ration sin primarily onsists of
a frquntly rworkd litany of argumnts prsntd as disproving volution,
suh as gaps in th fossil rord, th supposd inompatiility of th sond
rodl to vlution, and aus law of thrmodynamis with a gradually inrasing omplity of lif' th
tnt f ID thus dvolvs to th impossiility of dvloping phylum-lvl ody-plan diffrns through ran-
young.athrationists:gaps dom variation and natual sltion, and th ..vast information ontnt of
sltionto produ volution- living organisms'' (IR 1'987).It prhaps gos without saying that suh ..v-
xplosion' and so on. As is th idnagainst volution,' gneratslank looks from volutionary ilogists,
..ontrivd dualism,'' to quot
who ar puzz|d that unsolvd aras in sin suh as th origin of lif, th
rn shown to untru, ID is apparanof amrian ody plans, or th rlativ rol of sltivand non-
sltivmhanisms in volution ar onsidrd waknsss in volutionary
theory or rasons to rjt th infrn of ommon anstry.
Tahing th ..vidn against volution'' avoids th ovious stalish-
mnt laus prolm gnratd y ID, whih is ..\/hois th dsignr?'' Th
rl-tim laws,.tJ7ndllBird, who
iuprm ourt, gamly advisd stratgyis thrfor favord among th mor sophistiatd ntivolutionists
who ar oprating in th arly twnty-first ntury. The Disovry Institut,
night still lgally taught if it
for ampl, has om to urg shool oards not to rquir tahrs to tah
987). is prsptiv was not
ID ut rathr to tah th ..strnsthsnd waknsss',of volution.
ts who intprtd drds as
and rationism; drds has
rts in disions that hav struk
i f o r n i t o r i n g r t i o n s i n
Th Futur of Antivolutionism
alia and signd y Justi Rhn- Sott and Branh hav rfrrd to thr thms that hav run throughout th
rasquit ompatil with xtant thr priods of Amrian antivolutionism and ontin to illustrat th
inluding thos who ar Christ- ontrovrsytoday as th ..pillars of rationism'' (Sott and Branh 200'
282). t any givn tim in this history, ths thr argumnts hav n part
ular mattr, to hav whatvr
of th rationist position:
tion prsntdin thir shools,
'hatvrsintifi vidn thr
1. volution is wk or unsupportd sin that sintists ar
IR sizd upon this immdi. aandoning in inrasingnumrs.
;hd. hoing Salia' it nour- 2. volution is inhrntly antirligious.
..In th mantim'
t volution: 3. It is only fair to ..alan''th tahing of volution with rationism
gly nouragdat last to Strss or IntlligntDsign. A vaiant of th ..fairnss''argumnt is that it is
st volution in thir lasss (not good pdagogy to ..alan'' th tahing of volution with
..waknsss'' ..vidn
l t i o n a r ym h a n i s m , u t a g a i n s t or against'' volution ..and lt th hildrn
rogniz ths as vidns and did.''
'gumntsfor a partiular dat of
7\. Ths thr thms will ontinu to shap th rationism,/volutionon-
.W
idn against volution'' is asy trovrsyin th futur. an illustrat thm y xamining som rnt as
nd that thr ar only two possi- studis of antivlutionism. Although rligion ovrwhlms all ls as th
stfongstmotivation for antivolutionism' aus of lgal disions of th
;ifi vrsion of Christian spial
1980sand arly 1990s, rationists hav downplayd th sond pillar of
all th othr origin storis from
lm it is logial to infr that vi. rationismin favor of th first and third, at last whn addrssing th gn-
ral puli and, spially, puli offiials suh as shool-oard mmrs.
tionism.Pratial onsidrations
Both ration sin and ID proponnts hav hammrd away at th ida of
that support th spial ration
86 A m eri t iuolutioism
volution as wak sin and hav strssdth importan of fairnss' sp- Indd, onsrvativso1
ially in trms of tahing ..vidn against volution.'' that thy did not look k
Anothr trnd has n th ..morphing'' of Y to ID to ..vidnagainst t h s t a n d a r d s( u n n i n g
volution.,' In a numr of ommunitis, popl who ar young-arth r- D u r i n g h s p r i n ga n
ationists bgan to idntify thmslvs as ID supportrs' thus avoiding th trs to th ditors argur
stigma of ration sin, whih is at st onsidrd a minority viw, if not Inlusion of volution .
a fring prsptiv'in oth rligion and sin.IntlligntDsign, on th othr vlop qustions for th s
hand, is lss wll known to th gnral publi, and aus of th vaguenss in thos xams' it ws
with whih it is prsntd (making no laims aut th ag of th arth, fr rooms. Tahers and s
xampl), it appars lss marginal than ration sin.Latr' as a ommu- hav a prominnt pla
nity that is dating volution duation oms fanriliar with ID and its standards tht ws sub
sintifi strility and duious onstitutionality, th antivolutionists oftn th National Sind
rtrat to a fallak position of ..vidnagainst volulion,'' usually por- Aadmy of Sins,ar
traying it as a ompromis that should aptal to ll. Morphing of Sin (AAAS) Bnh
this sort ourrd during th wll-puliizd Kansas stat sin standards bn influntial in shap
ontrovrsy of 1999. NSS and th Bnhma
inlud it throughout th
In midsummer 1999
As sUDY: KANSAS
standards ommitt'sd
As a rsult of duation rform movmnts that bgan in th 1980s' most l r g | y n w r i t t n b y
Amrian stats hav stalished duational standards for history, math- ation for Mid.Amria,
matis, sin, and other aadmi sujts. Aftr th fdral No hild Lft of St. Louis. Arams's st
Bhind At of 2001 mandatd priodi high-staks tsts for studnts, stat or apabl th wor
standards took on vn mor importan. Tst topis would asd on th ludd muh aout how
standards; if volution wr inludd in th standards, thn it would in- puli harings oftn ml
ludd in th tsts. Stats that did not alrady hav standards or framworks draft of th standards t
gan to dvlop thm in th 1990s; th vast majority of thm inlud volu- ompomisdraft thtit
tion, though in varying dgrs of omptny (Lrnr 2000). any stats draft ompromisd btl
that had ignord volution prviously now rquir that th topi taught. thos who wntd the ,A
A good dal of th antivolution ativity of th past fw yars has n di- is suh s th ig ang
rtd t th stalishmnt of stat sin duation standards: rationists standards and adding s
oppos th inlusion of volution in th standards,or try to gt ID addd, or of sien.'
try to gt ..vidn against volution'' inludd. Muh as th rturn of vo. A loud hu and ry r
lution to th urriulum in th mid-1960s gnratda rationist aklash' paprs' tlvision, and r
so also has th standards-asd rquirmnt that volution taught gnr- tlvision talk shows, ;
atd its own aklash. Kanss Wizrd of z ir
Kansas undrtook to writ sin duation standards in 1999. 27- Kansas suffrd at th l
prson ommitt of mastr tahrs and sintists was appointd y th th nt shool-oard l
Kansas Dpartmnt of duation and th Kansas Board of duation to draft at majority swiftly abar
standards that thraftr wnt out for puli ommnt' wr rvisd, and adoptd th final draft 1
wr ultimatly sumittd to th oard of duation for approval. This pro- th nw oard took offi<
dur was prolongd, ontntious, and ittrly fought. A onsrvativ lo Th Kansas ontrov
of fiv shool-oard mmers had om to powr in th 1996 state shool- rationism, th lim tt
oard ltion and lashd with th fiv modrat shool-board mmrs Proaly baus th
ovr many issus. Whn th sin duation standards am up for onsid- arth rtionists' and t
ration. it was an asy ordition that volution wld a ontntiousissu. ludd in th sinsta
mri Atiuolutionism 87
and shool oards hang om- pulishroffings,Prsntd on-siddtratmntof volution nd did not
ldratsprvaild, in 2002 th provid ny ..sintifi altrntivs.'' Aordirrg t<l a nwspapr aount'
5 ratio of nsrvativsto mod- aldwll first proposd tht ID e inludd in th urriulum nd in th tt-
: llt alon during that trm, ut books (Rosn 2003a). Rathr quikly, howvr, aldwll harrgd his pro-
. ftr th 2004 ltions mant posal to promot tahing ..idasthat ountr volution'' (Rosn 2003). H
< pla in 2005 was highly on. rommndd a vidotap promotd y th Disovry Institut asd on
l mmrs supportd standards Jonathn!ells'sook los of uolutio (22)plus anothr lD-proroting
rtionsinor ID, but for th video, Ulokig th ster oi Life (2002). Arrlthr itizn sumittd a
ln. Th standardswr passd in Y diatri y Jonathan Sarfati at Answrs in Grrsis cal|ed Rfutig uo-
rrds Adoptd in Kansas'' 2005) Iution(1999). A uiquitous ration sinook pulishd y th Jhovah's
; and sintistsand national si_ Witnsss, Life: Ho Did It Get Hr? B uolutio lr retioi (Wth-
ing t AAAS and th National twr Bil nd Tat Sorty 1985) was also propos1.
lf th Kansas SinStandards'' Instrutionalrnatrialsproposd for lassrootn us in Rosvill, :rsin lros
j t i o n y a r ' n d t w o o n s r v t i v ommunitis'lllst rviwd y a ommitt of tahrs and drninistra-
lum Swingsin Kansas" 2006). In tors. T giv thir opinion mor rdiility, th tahrs askd sintists
:t shool oard rsindd th fom loal univrsitis to rviw aldwll,s suggstdmatrials. T.h om-
nithrdnigratdvolution nor mittrportd to th shool oard that th matrialswr unsuital for th
:ion Rturns to Knss'' 2007). Rosvill urriulunr and, rfltingtlr sintists' valuations, sustandard
g of young-rth r t i o n i s t si n t o sin.In Sptrrrr2003 th ord, stymid y th unitd fronr prsntd
ts who ativly opposd th 1'999 by th valuation ommitt, votd to lt th individual shools did
ID supportrs.John alvrt is a whthror not t use th matrials.Thrsonsidrdthis an nd run that
Dsign Ntwork (IDNI), an or- might possily rsult in th use of th ooks in th lassroom' vn though
rnspromoting ID in Kansas thyhad n rjtdy th valuirtionommitt'
rr iruit ridr for lD, travling rtionistson th oard lso rommndd tlrt shlol lir:rrisst up
i n i e ,a n d o n t n a t o p r o m o t i t . evolutionrsour ntrsthat would hav ooks, vidos, and othr matri-
to disouagth promotion of als ritiizing volution. Th ord mmrs apparntly rasond that if
vidnagainst volution.'' Th tahrsould not omplld to tah ..vidnagainstvolution,'' at last
ah sin the 202 ohio sin studntsould go to th lirary to gt antivolution information. Lirarians
onttious2005 rvision rfth protstdtht th hd littl nough spa in thir liris fr matrials tht
spportth r.rrriulum-th linritd purpos of a shool lirary; thy did not
wnt to wast mony or spa on what thy onsidrd a suprf-lttous issu
unrlatdto th nds of tahrs for urriular support. Th volution r-
)RNIA
sourntrsWr nvr stalishd.
,rsis;rndantivolutior-rismlgis- aldwll nt attllptdto hav th oard pass il poliy h draftd,whih
:,th majority of antivoltionism halld Quality Sindr-ration(Qs). It would hav rquird th th-
lvl. A ontrovrsy in Rosvill, ing of ..strngthsnd waknsss.'of volution- hoi of wording similar
lf rationism,as wll aS th mor- to poliisthat hv appard in som othr omnrunitis.ontovrsy ovr
volution. QS ontinud ovr th wintr and ulminatd in a ontntious mting on
ly 92,000 lotd 20 mils north. May 10, 204, in whih mmunity mmrs datd th poliy for ovr
o n i . A s u s t a n t i l o n s r v a t i v two hours. A ptition signd y 28 of te 2 sin thrs in th distit
shool-oardmms who shar opposingth poliy was suritrdto th oard. In addition, tlr rrrajorityof
r-war''ontrovrsisin Rosvill poplwho tstifidat th shool-oard mting opposd QS.
duationas wll as volution. Th In Jun 2004 th shool oard votd 3_2 against QS, nd th issu of
n th summr of 2003 whn high- rationismin Rosvill smd to hav n didd in th ngtiv. Pr-
wd for adoption. A loal itizn, haps ausof onrplaints from Parnts tht th ord ws Wasting tim
:tooks,all standard ommrial arguirrgovr rtionism whrr ilportant issttsof funding, ovrrowding,
9 0 A m ri ti uo luti li sm
and othr rad-and-Llttronrns wr rrot ing addrssd,rh tssuws that this is whr th ratit
not brought r.rpagin, and in th N<lvmber2-004ltion mor modrat th mo obvious prsslo
andidats prvaild, putting an nd to th urrnt Wav of antivolutionism sign and toward th lss l
in Rosvill. Som tahrs' howvr, rport that it is mrly a mattr of against volution.''
..vidnagainst voltion'' riss in
tir until th issu of rationism nd
Rosvill irgin,at last as long irs rligiotrslyonsvtivshool.oard rnm- PRED||oNs FoR
rs kp gtting eltd. In January 2005 Larry aldwll sud th distrit
and som offiials on th grounds that his onstitutionl rights wr irrfringd Prdition is unrtain,sp
during th ontrovrsyin Rosvill.Dfnding thmslvs,th distitlaird to hv said, ut som ft
on th ontrary that aldwll's proposals had rivda full hring, and, in ism,/volution ontrovrsy
fat' an undu amount of tim irnd nrg Was spnt on aldwll.s issus. distrit-ourt judg larn
A judg sidd with thrn in Nor,mr 2007, and ldwll lost his sLlit trit struk down txtbo(
(Rosnhall 2007). y th Co ounty, G<
. . T h i s t x t o o k o t i n sn
fat, rgarding th origin ol
oRPHlNG oF RA|oNlsMs
with n opn mind, studi
Kansas and Rosvill illustat tlr morphing phnomnon tht has n tak- P a r n t si n t h o m m u n i 1
ing pla for svral yars. ommunity mrnswho ar idntifid with or 2002, ut th as took a l
hold Y positions astthmslvsas ID advots.In truth, rhis is nor a th shool oard hd a rl
lngthy jotrny, aus all young-arth rationists r supp()rtrsof th Christianity) in rquiring tl
dsign viw, although not all ID supportrs ar young-arth rationists.In furthrmore, undr Gorgi
Rosvill, for xampl, oth Y and ID rsorrrswr offrd to th shool tional aus it rquidt
distrit. Th lgal advlntagof ing idntifid wit positior-r'lD, that has Th judg struk down th
not alrady n struk down y th ourts is lar. Yt ID is not without its i n g r I i g i o n .I n t h d i s i
waknsss:th phras itlligetdsig arris th ilpliation of a dsigner, sholarly rtils' as wll a
.,onol
an agnt, and it is no srt tlrirttl-risagnt is th hristian God. Prhps for fat'' langug was
this rason, th Disovry Institutativists' fom approimatly th last half m p l o y dh y a n t i v o l u r i o n
of 2001, hav n arguing not fr th inlLlsionof ID in th urriulum but gling out volution from
f o r t h i n l t l s i o n o f . . v i d n g a i n s t v o l u t i o n ' ' ( o r . . s t r n g t h s : r n dw k . would hv th fft of pr<
..ritial ..tah is rationism. Th distrit
nsss of volution''' analysis of volutiorr,'' or simpl th
ontrovrsy''). Tahing ID is mor likly to run afoul of th stalishmnt pals vaatd th ruling an<
l:1Lls than thing ..vidnag:rinstvolution,'' altl-roughth nt fft of sideration or rtrial. Th gr
nouragilrgstudntsto apt spial-rationtholog is onrmon to itlrr not onrn th judg'srulil
approah. i n g v i d n .I n d i s u s s i o
Parallling th IR's ditum that ..vidnagainst volution is vidn agrd to rtry th as.It h
..limittions plaintiffs' lgal tal riv
for ration sin.''Wiltim Drski hs writtn that on volv-
aility y ratrial rlhanisms onstitut vidn fo dsigl-t'' (Dmski prt witnsss' whih strn
2002). Talring studnts that volution is wak or unsupportd sirrthat ruld in thir favor. Th pll
annot xplin th divrsity of lif nourgsth aptanof rationisnr: tion of som of th attorn
if volution dt>snot plin living things, y dfault,rationismdos. shool ord poliy rqui
..vidnagainst nation of fators was suffi
Suh morphing of Y to ID to volution'' has ourrd
in ir numr f ratitlrrism/volution ontrrvrsisovr th pst fw yars, The sttlmnt was ver fa
inluding ontrovrsis ovr th writing of sit-tduation standads in agring nvr again to dis
Kansas, Ohio, ./stVirginia, Nw io, and Minnsota and in shool- orally, or in any othr for
distrit ontrovrsisin Dary. ontana. d Dovr. Pnnsylvania.It is lar gia stat standards, whih
Amricn Atieuolutionist 391
t ingaddrssd,th issu was that this is whr th rationism/volution ontrovrsy is hadd: away from
r 2004 ition mor modrat th mor ovious xprssions of rligion or ration sin or Intllignt D-
l r r n tw a v o f n r i v o l u t i o n i s m sign and towrd th lss lgally vulnral stratgy of tahing ..vidn
t that it is mrly a mattr of against volution.''
n againstvolution'' riss in
:onsrvativsho1-boardmm- PRDI|oNs FoR (NAR) FUURE
-arry aldwll sud th distrit
lstitutiOna1rights wr infringd Prdition is unrtain, spially aout th futur, as Yogi Brra is supposd
g thmslvs, th distrit laimd to hav said, ut som rnt vnts may rshap th futur of th ration-
d rivda full haring, and, in ism/volution ontrovrsy in th Unitd Stats. In January 2005 fdral
Was spnt on aldwll's issus. distrit-ourtjudg larn oopr tn Slm u. obb out School Dis-
l07, and aldwll lost his suit trict struk down a txtook dislaimr ordrd insrtd into iology ooks
y th o onty, Gorgia, Board of dution. Th dislaimr rad:
..This ttook ontins matrial on volution. volution is a thory, not a
fat, rgarding th origin of living things. This matrial should approahd
with an opn mind, studid arfully, and ritially onsidrd.''
arntsin th ommunity had sud th distrit and th oard of duation in
; phnonrnonthat has n tak-
nrswho ar idntifid with or 2002' ut th as took long tim to om to trial. Th plaintiffs argud that
d v o t sI.n t u t h , t h i s i s n o t th shool oard had a rligious purpos and fft (of promoting starian
.ationistsar suppotrs of th hristianity) in rquiring that th dislaimr plad in txtooks, and that
r young-arthrationists. In furthrmor' undr Gorgia law' th shool oard's ation was unonstitu-
oursWr offrd to th shool tional aus it rquird th xpnditur of stat funds to promot rligion.
[d with a position, ID, that has Th judg struk down th prati ausit indd had th fft of promot-
is lar.Yt ID is not without its ing rligion. In th dision th judg took notie of law.rviw and othr
risth impliation of a dsigner, sholarly artils' as wll as tstimony that pointd out that th ..thory' not
s th hristian God. Prhaps for {at''languag was ..on of th latst stratgisto dilut voltion instrtion
from appoimatlyth lst half mploydy ntivolutionists with rligious motivations'' (Slm, 1308). Sin-
rsion of ID in th urriulum ut gling out volution from all sintifi thoris for spial ngativ tratmnt
..strngthsand wak- would hav th fft of promoting th favord altrnativ to volution, whih
;tion'' (or
..tah th is rationism. Th distrit appald th as. Th Fifth iruit ourt of Ap-
:volution,''or simply
o run afoui of th stalishmnt palsvaatd th ruling and snt th s ak to th Distrit judg for ron-
ution,'' although th nt fft of sidrationor rtrial. Th grounds for rturning th ase to th lowr ourt did
ion thologyis ommon to ithr not onrn th judg's ruling pr s, ut rathr a mssy ourt rord and miss-
ing vidn.In disussions among th judg and th two lgal tams, it was
n against vlution is vidn agdto rtry th as. It gan to look unpromising for th dfenswhn th
..limitations on volv- plaintiffs' lgal tam rivd prmission to ropn disovry and bring in x-
lrittnthat
vidnfor dsign'' (Dmski prt witnsss,whih strngthndthir as for a judg who alrady had
ak or unsupportd sin that ruld in thir favor. Th plaintiffs' lgal tam was also uttrssdy th addi-
tion of som of th attornys and witnsss who had sussfully dfatd a
] st h p r n o f r a t i o n i s m :
y dfault,rationism dos. shool oard poliy rquiring ID in Dovr' Pnnsylvania; prhaps th omi-
againstvolution'' has ourrd nation of fators was suffiint to nourag th distrit to sttle out of ourt.
ovrsisovr th past fW yars' Th sttlmntwas vry favoral to th plaintiffs, howver, with th distrit
,f sinduation standards in agringnvr again to dislaim vlution, whthr with txtbook stikrs, or
, and Minnsota and in shool- orally,or in any othr form. Th distrit also was rquird to follow th Gor.
nd Dovr, Pnnslvania. It is lar gia stat standards, whih urrntly rquir th tahing of volution, and to
92 Amri Antiuolutioism
rfrain from ising or rdating matrials On vollltion from instrtional rligious ida. Th pl
rnatrils.This lattronsrraintwas stimulatdy an arlirpratiin th dis- dfinitionalhartr
trit in whih th pages on volution in ttl.looks routinly wr t out to allss nd thus tsta
prvnt studnts from rading thm. posd that aus ID
Had th o ounty distrit lost on appal' othr ..thory'not fat'' poli- thws no pdagogi
is and dislaimrs would hav n nouagd, and I prdit w would At th nd of a long t
hav sn an aundan.Dislaimrs ar popular with shool oards aus did not mrly strik d
thy provid th apparan that th shool ord is ..doing somthing'' qustion of whthr ID
aout th tahing of volution without rquiring urriulum hange or th that th shool oard u
othr and pns of rviw and purhas of nw instrutional matrials. ligious viw' ut .|udg
Thy satisfy a onsrvativ hristian onstituny without spnding mu mnts for and agair-rst t
..in th h
tim or mony' that h did so
In Dmbr 204, a month aftr th Slm as Was trid' a lawsuit was and othr rsourswh
fild y parnts of hildrn in Dovr, Pennsylvani' protsting a poliy of its ing th pris qustion
oard of duation that would hav rquird th tahing of ID and a variant Although ID suppor
of ..vidn against volution.,' Th Dovr rsolution, whih latr was in- ritiizd te Kitzmille,
srtd into th urriulr-rm,rad: ..Studnts will mad awar of gaps/pro- rtly) dlaingID was
lms in Darwin's Thory and of othr thoris of volution inluding, ut not quird th judg to ult
limitd to ID. Th origins of Lif is not taught.'' In Novmr 2004' amid ould onstitutio
ommunity rumors of a possil lawsuit' th Dovr Board of duation is- iudg's dision on wh
sud a ..larifiation'' that statd that th ID ttook of Pds d Pople witnsss for oth sid
would mad availal to srudntsas a rfrnfor th topi of ID. on- widly usd by sintt
fusingly, th larifiation also statd that tahrs wr not to tah ration- Was that th dfnswi
ism or IntlligntDsign. ut th judg disagr
In thir omplaint plaintiffs argud that ID is a rligious onpt indistin- shool lassroom.
guishal from rationism' and that its tahing would violat th stalish. Th witnsss for th
mnt laus. Th tril took pla ovr a si-wk priod in th fall of 2005. p r s u d r h j u d g o f r
Th Disovry Institut pulily distand itslf frorn th lwsuit, stating that irduib1 omplxit1
..although of th llod lottirrg
w think disussion of intlligntdsign should not pohiitd,
w don't think intllignt dsign should rquird in puli shools'' thy wntd and mort
(..Lading Intllignt Dsign Think Tank'' 2004). Although originally a num- Shool Board disior
r of Fllows of th Disovry Institut agrd to tstify as prt witnsss sin-whih will
on half of th dfnse, all ut two withdrw from th as, ut not until it of ID.
Was too lat to add nw prt witnsss. But th two who rmaind, If th Dovr Ara S
Mihal Bh and Sott Minnih' wr th ID proponnts with th st si. th ountry would har
ntifi rdntials, and thus th most qualifid to dfnd th sintifi rdi_ gardlss of th hor
bility of ID. tahing of [D y st
Th ky lgal issu in Kitziller u. Dour Ar S'hool District 'vlas viwd as zr fornr of
whthr thr was ir valid sul rason to th ID. Th ovious rligious ouag th ringing
onnotations of lD would not hav n suffiint groDnds to strik down diffrnt nam. Howr
th poliy had thr n a valid sulr purpos and fft for tahing it. p a l d l l d t h r f o i s
Th dfns,thn, had to dmonstrat that ID was a valid sin,and that ID proponnts might
tahing it as an altrnativ to volution would provid studnts with an ity of an ID poliy' wi
llnt ritial thinking xris. vr, b:tttsof th l
Th plaintiffs had to onvin th judg that ID was ot valid sin,and matrial irr th trial r<
thrfor th only rason to tah it was unonstitlltionallv to Dromot a stantially diffrnt out
mri Atieuolutioism 9
on volution from instrutional rligious id' Tlr plaintiffs olltndd that ID did not mt important
c y an ariir prati in th dis- dfinitional hrirtristisof sin (suh as ing rstritd to natural
books routinly wr ut out to usSand tlrus tstal)and tlr:rtits fat lairns wr inorrt. Thy pro-
posd that as ID was not :l sin and its fat laims wr rfutd,
..thory,not fat'' poli- thr was no pdagogil rason to tah it in a sin lass.
al, othr
rragd, and I prdit w would At th nd of a long trial' th plaintiffs wr rwardd with a dision that
'ular with shool ords aus did not mrly strik down th Dov poliy, ut that also wighd in on th
..doing somthing'' qustion of wlrthr ID was sintifi.Thr ws plnty of vidn showing
ol oard is
uiring urriulum hang or th that th shool board unonstitutionally intndd to promot a strian r-
' of nw instrutional matrials. ligious viw, ut Judg John . Jons III additionally onsidrd th argu-
ituny without spnding muh mnts for and against th lim that ID was valid sin.Judg Jons wrot
..ir-r
that h did so th hop that it may prvnt th ovious wast of judiial
lJtiC?Sewas rrid, lawsuit ws rrdotlrr rsourswhih would oasiond y a susqunttril involv-
,lvania' protstinga poliy of its ing th pris qustion whih is for us,, (Kitzmiller,75).
r h t a h i n go f l D n d a v a r i a n t Although ID supportrs from th Disovry Institut (D.!7olft al. 2006)
. r s o l u t i o nw
' hih lt was in- ritiizd the Kitziller dision as unnssarily (and in thir viw, inor-
vill mad awar of gaps/prob- rtly)dlaring ID was not a valid sin,th lgal thoris of oth sids r-
s of volutioninluding, ut not quird th judg to ul on prisly this pint: if ID was valid sin,thr
ught.''[n Novmr 2004' amid ould b a onstitutionally admissil sular rason for tahing it. Th
l Dovr Board of F-duationis- judg'sdision on whthr ID was sin ws rlativly straightforward:
' ttook of Pds d People witnsss for oth sids tstifid that y th standard dfinition of sin as
frnfor th topi of ID. on. widly usd y sintiststoday, ID did not qualify as sin.Th diffrn
h r sw n o t o t h r a t i o n - was that th dfnswantd to hrngth dfinition of sinto inlud ID,
ut th jirdg disgrd thr sul.ra dision should b md in th high_
D is a rligious onpt indistin. shool lassroom.
:hingwould violt th stalish- Th witnsss for th dfns did not far any ttr in thir ttmpt to
.wk priod in th fall of 2005. prsuadth judg of th validity of th laims of ID (suh as th supposd
slf from th lawsuit, stating that irrduil omplity of th atrial flagllum and th unvolvaility
dsignshould not prohiitd, of th lood lotting mhnism). In th nd, th plaintiffs got vything
rquird in puli shools'' thy wantd and mor: a dision not only striking down th Dvr Ara
004). Although originally a num- Shool Board dision, but a dision valuating ID as not qr-ralifyingas
.edro tstifyas xprt witnsss sin-whih will itd in any futur attmpt to rquir th tahing
w fom th as, ut not until it of ID.
:s. But th two who rmaind, If th Dovr Ara Shool Board lrad Won th as' shool distrits around
ID proponntswith th st si- th ountry wold hav n rrouragdto impos similar ID poliis, re.
id to dfnd th sintifi rdi- gdlssof th hortations of th Disovry Institut not to rquir th
tahing of ID by statut. Int[lignt Dsign is widly-and auratly-
|ouer r Shool Distrit was viwd as a form of rtionism, and a lgally sussfulpoliy would n-
l tah ID. Th ovious rligious ourag th ringing of rationism into th lassroom, alit undr a
uffiintgrorrnds to strik down diffrnt nam. owvr strong a dision, te Kitzliller ruling Was not ap-
urposand fft for tahing it. paldand tlrrfois prdntonly in th iddl Distrit of Pnnsylvania.
: ID was a valid sin, and that ID proponnts might attmpt to find anothr vnu in whih to tst th lgal-
would provid studnts with an ity of an ID poliy, with a diffrnt st of fats and diffrnt judg. How-
vr, aus of th rdth of th Kitzmillr dision nd th walth of
,hatID was not valid sin, and matrialin th tril rord' it is doutful that anoth trial would hav a su-
unonstitutionallyto promot a stantialiydiffrnt outo1.
94 mricntiuolutilism
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1. B2.
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"itiqu of Itlligt Dsig Thlr.
y rirsning.lntrtkl J ou'rnl
l Did lt Gt Hr? 8y
7athtowrBibl and Trat Soity
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m Hdoisfs.Downrs Grov, IL:
40
gssiz
BlBLIoGRAPY
Agassiz,JanLuis Rdlphe$8a7_t873\
Born in Switzrland in 1807, l,ouis Agassiz as a oy Was a kn naturalist.H o lssifitio (|8,
studid iology at svralGrman univrsitis,arning an advand dgrin aross mryoni for
..On !
narural history \n 1829. H soon arnd a rputatioll as a brilliant rsarhr' th istn of
thanks to ooks dsriing spis of fislr' living and fossil' and to his arful blans for h a
.silliam
studisof Alpin glairs(Agassiz 1833_1843' 1840) (sfigur).Bginningin Paly, whos
1837 h ssmldvidn for surprisirrgnw thory aout an I Ag, makr or nginr Wh
whn a thik lyr of i had rvrdnorthrn urop. Aftr moving to th pupos. Agassiz.sGod
Unitd Statesin 1846,e amon of Amria's lading sintists,founding ral variations on a fv
th usurn of omparativ Zoology at Harvard in 1859. Agassiz rgudthat
A harismati lturr' Agassiz hamd th puli as wll as his studnts ation and nnot har
with dsriptions of rnt disovris in mryllogy' palontology, and jtd th iblial sto
anatomy. Lik Rihrd Own in Lndn, Agassiz mphasizdth pattrn of hallngd whn h n
rsmlansalld hornolgv. Althouglr it had long n known that tlr dos him no rdit'
strutur of an animal's organs is latdto thir funtion, not until th nin- an opinion wlomd
tnth ntury did hiologists onfront th fat that divrs raturssm to Darwin's o th '
b modifitions of ommon luprint or typ. Vrtratsar on sllh prtation of narr.rr. Pt
typ' austh sam layout of ons and oth organs an trad in a win argud that hom
fish, a frog, a at, and a humn. Agassiz imprssdhis audinswith unfa- ommon ansto,wh
miliar ampls of tlr sam prinipl:tht s,spidrs,and lostrsar 2rlso n t r a l s l t i o n .B f
honrologous to otl anothr' s ar otopuss,snails, and lams, or stirrfish, a P r d v o l u r i o n ,i f
jllyfish, sa urhins, and orl polyps. F.ollwing Gorgs uvir, h alld t o B r a z i l i n 1 8 6 5a n d r
thsfour goups vrtrats,ilrtiLllats,mollusks, and radits.In his ssay his mind. H did in 1
Agssiz 40''
; sulrr simply not wll takn.
lrs,a strippd-down.pu Dr-
i v ,i t i s s i n t i f i a I lsyu r i o r -
ommunitv, whih has spnt 1
d ooks rfuting ky laims of
,t$5st
haviour7I.lourl l| Thortil
t7-1873)
as a oy Was a kn naturalist. o lssifction (1859) Agassiz argud that homologis, whih tnd
is'arningan advand dgr in aross mryoni forms and ak thrugh th fossil rord, ar vidne of
putationas a rilliant rsarhe' th istn of ..one Suprm Itllign'' who hd onivd ths rsm_
iving and fssil, and to his arful blnsfor h ratd thm. This agumnt is quit distint from tht of
l 3 , l 8 4 0 ) ( s f i g u ) .B g i n n i r l gi n William Paly, whos Ntrl Tholog (1s02) omard God to Wth-
n8 nW thory out n I Ag' makr or nginr who uilds mahins that ar lvrly adaptd to thi
hrn urop. Aft moving to th purpos.Agassiz's God is lik musiin or arhittwho njoys playing
sv_
rria'sladingsintists,founding ral variations on a fw asi thms.
a r v a r di n 1 8 5 9 . Agassiz argud that ah spis was th produt of a listint at of
r-
th puli as wll as his studrrts ation and annot hang, ut h ndrstood gologial tim and firmly r-
mbryllogy'palorrtologyJ nd jtdth ilial story of ration. His onpt
of spis was svrly
\gassizmphasizedtlr pattrn of hallngdwhn h nountrd popl of Afrin ds.,'t, and his rspons
it had lorrg n known that th doshim no rdit. insistd that human ras ar atually distint spis,
thi ftrntion'not Llntil th nin- an opinion wlomd y slav ownrs'
.atthat divrsraturs sm Darwin's on tbe rigin of Spcies was a srious thrat to Agassiz's intr-
to
) typ. Vtrtsr on suh prtationof natur. Pointing to th sam phnomna itd y Agassiz,
Dar-
d othr organs an b trad in a win argud that homology and iogography are vidn of dsnt from
a
prssdhis audins with unfa. ommon anstor, whil th adaptdnss of form to funtion is viden for
s,spidrs,and lostrs ar also naturai sltion. Bfr long Agassiz's students and rost of his ollagus
ss' snails' and lirms, or starfish, aptdvolution, if not natural sition, ut Agassiz nvr did. Travling
lowing Gorgs uvi, h alld to Brazil in 1855 and to th Galjpagos Islands in 1872 did nothing to hang
rllusks,nd rdiats.In his ssa1' hls mind. H did in 787.
106 lndr
Th prolm of ltruism is dividd into two sparatut rltd issus:i. (1998) argu that' und(
ologial altruism and psyhologil altruism. In iologial altruism, an or- ltion. If th avrag1
ganism prforms n altruisti at whn th at nfits anothr organism at a thn th avrag fitnss
ost to itslf. Bnfits and osts ar masurd in trms of rprodutiv fitnss. fitnss onsquns of
Suh altruism is wll doumntd in many spis. Vrvt monkys nd t a i n p d o m i n a n l '
prairi dogs hav sntris at th outskirts of olonis that alrt mmrs of suvrsionfrom wirhi
th olony of approahing pdators,oftn at th ost of oming th tar- twn groups' Althou1
gts of thos prdators. Psyhologial altruism, whih is th olloquial sns workd out and th pll
of th trm, is intntionally prforming an ation at on's own ost in ordr studis of flour tls
to nfit anothr. Th nfits and osts for psyhologially altruisti ats, sltion modl ar m
howvr, nd not hav anything to do with rprodutiv fitnss. An xampl ratly xplain naturl p
of a psyhologilly altruisti at wold to plant a slow-growing tr, not xplin how altru
knowing full wll that you will not nfitfrom its shad or fruit' mutation in bhaviora
Although iologial and psyhologial altruism ar sparatprolms, th- for thy om nur
orists hav offrd iologial undrpinnings for psyhologial altrism. Th A sond rson fo
prolm of altruism, oth iologial and psyhologial, is at th ntr of pat of lWilliam Hamilt
grounding a thory of morality within iology. I will disussth two kinds of has om to known.
altruism in turn. havior is gntlallya
Altuism sms to a prlm for volutionary thory whn looking at s- tivs, with whor thy
ltion from an individual prsptiv (i..,sltion that ous on th lvl of altruisti gns througl
organisms). If altruisti organisms liv among slfish organisms' th slfish r- \/ith this xplanation,
ganisms will always mor fit than thir altruisti ountrparts. ovr tim, Frm gn's-yviw
altruisti individuals will sltd out in favor of slfish ons. For ampl, fi hrslf for th livs
sntry monkys lowr thir fitnss y aling othrs to th prsn of prd- of hr gns.But th
ator. ovr tim, slfish monkys, ons that do not tak thir turn on wath' fo th ost of slf-sar
would produ mor offspring and vntually liminat any altruisti gns. Kin sltion appar
harls Darwin' tnTh Descet of n (7871)' notid this prolm and m o t h r s d f n d i n g h i
..although high
hintd at a solution: a standard of moality givs ut a slight planation is that it r
or no advantag to h individual man . . . an inras in th numr of ut largr familial rla
wll_ndowd mn and advanmnt in th standard of morality will r- than distant ons' Whi]
tainly giv an immns advantag to th group,, (p. 1'66).Darwilr ontinus sual or olfatory us' tJ
by suggsting thet groups that ontain mor altruisti individuals will out. fftiv prois for rl
ompt groups omprisd mainly of slfishindividLrals.From this pssag,it that r los y'' will
appars that Drwin Was suggstingthat sltion an our at th lvl of Suh aggrgationn o(
groups' a radial dpartur from his or afgumnt aout th importn of if disprsal from th ho
sltionamong individuals. But how an w pli
volutionary iologists hav vaillatd in thir aptan of this so-alld rlatd if group sltio
group sltion argumnt. Darwin's influn gav it grat rdn for d- tion was artiulatd by
ads. It fll from favor in th 1960s undr th influn of Gorg C. iprol altruism. A<
IWilliams's writings (e.g., 1966| aout sltion and adaptation aus
toward othr organism
group sltion for any attribut am to onsiderd a iologial rarity. This planation dos
on of th main prolms with the group sltion argumnt is that omp- stats in animals. A sta
tition within a group will always favor th slfish individuals, so that vn if havior of vampir a
groups of altruists ar mor fit as a group than groups of slfish individuals, out fding; howvr, v
th groups of altruists will suff suvrsion from within. Rntly, lrowvr, individual nds. Vam
.Wilson
group sltion nrodls hav rivd a spiritd dfns.So arrd bats who hav rntly
Altruism 409
o sparatut rlatd issus:i- (|998) argu that' und rtain onditions, altuism an promotd y s-
r. In iologial altruisnr, an or- ltion. If th avrg fitnss of groups prdominatd y altruists is gratr
:t nfitsnothr oganisrn at a than th avrag fitnss of groups prdominatd y slfish individuals, th nt
in trms of rprodutiv fitnss. fitnssonsquns of ing an altruist may positiv. Groups an main-
y spis.Vrvt monkys and tain a prdominantly altruisti population and dfnd thmselvs ginst
olonisthat alrt mmrs of subvrsion from within if thr ar priodi migrations of individuals -
at th ost of oming th tar- twn groups. Althorrgh th mathmatis for group sltion hav n
lm' whih is th olloqtrial sns workd out and th plausiility of th thory onfirmd with primntal
tion at on'S own ost in ordr studis of florrr tls, it is lss ovious that th rquirmnts of th group
lr psyhologiallyaltruisti ats, sltion modl ar mt oftn in natur and thus that th modl will au-
produtivfitnss. An ampl ratly xplain natural phnomna. Furthrmor' grouP sltion modls an-
to plant slow-growing tr' not plain how altruists, whn initially rar (as would happn through
'om its shad or fruit. mutation in havioral traits), avoid ing liminatd by slfish individuals
'uism sparatprolms, th- fothy om nutnrous nough to at a group sltiv advntag.
for psyhologialaltruism' Th A sond rason for th lips of group sltion xplanations is th im-
'syhologial,is at th ntr of pat of William amilton's famous pprs (|964a,1964) dsribing what
4y.l will disussth two kinds of has om to known as kin sltion. amilton argud that if altruisti -
havior is gntially asd, thn organisms that wr altruisti toward rla-
'onary thory whn looking at s. tivs,with whom thy shr gnti matrial, would promot th sprad of
:ltionthat ours on th lvl of altruisti gns through th nfits to th rlativs that rry thos gns.
g slfishorganisms, th selfish or- With this xplanation, group sltion was not ndd to plain altruism.
ltruistiountrparts.ov tim, From a gn's-yviw it would b sltivly nutral for a mothr to sri-
avor of slfish ons. For ampl, fihrslf for th livs of two hildrn aus ah hild possssson-half
] othrs to th prsn of a prd- of hr gns.But th nfit of saving thr offspring mor than ompnsats
do not tak thir turn on wath' for th ost of slf-sarifi.
l y l i m i n a t a n y a l t r u i s t i g n S . Kin sltion apPars ommonly in th biologial world (just think of
( 1 8 7 1 ) ,n o t i dt h i s p r o l m a n d mothrsdfnding thir young). On potntial prolm for th kin sltion
lard of morality givs ut a slight planationis that it rquirs organisms to idntify not only family mmrs
. . an inras in th numr of but lgr fmilial rlationships, eus losr lativs ar mor valual
r standard of morlity will r- than distant ons. \ihil many organisms n idntify thi kin through vi-
oup'' (p. l66). Drwin lntinus sual or olfatory us, th kin sltion argumnt works vn if othr us ar
'r altruisti individuals will out- fftivproxis for rlatdnss. A huristi rul lik ..shar with organisms
i n d i v i d u a l s F. r o m t h i s p s s a g ,i t that ar los y'' will work whn family mmrs tnd to aggrgat in spa.
ltion an our at th lvl of Suhaggrgation an ou vn in animals without advand soial systms
rgumntaout th impotan of if disprsal from th hom sit is limitd.
But how an w plain altruisti havior twnorganisms that ar not
t thir aptanof this so-alld rlatdif group sltionis not th answr? A proposd solution to this qus-
1 gav it grat rdn for d- tion was artiulatd y Rort Trivrs (|97|) and has om to known as
:rdr th influn of Gorg . riproal altruism. Aording to Trivrs, an organism havs altruistially
ltion and adaptation aus toward othr organisms in th ptation that othrs will hlp it in rturn.
onsidrd a iologial rarity. This xplanation dos not prSuppos a onsious dision o vn mntal
sltion argumnt is that omp- statsin animals. A standard ampl of iproal altruism is found in th
slfishindividuals, so that vn if haviorof vampi ats. Vampir ats an surviv only a day or two with-
than goups of slfish individuls, out fding;howvr, vampir ats that find pry an fed in xss of thir
n from within. Rntly, howvr, individual nds. Vampir ats will oftn shar mor, and mor oftn, with
.Wilson
riritd dfns. Sor and batswho hav rntly shard with thm.
410 ltruism
tim. Having stalishd th two vnts and thir synhrony, thy laoratd
th ologial impliations-that a larg impat hd ausd a dust loud to
nirl th arth, utting off photosynthsis and making it old and dark for
a suffiintlv long piod so that virtuallv all land animals ov 50 pounds
did' along with som 7 5" of marin organisms-and drw th infrn
that th impat had ausd th tintion of a larg fration of spis 65
million years ago' inluding th tintion of dinosaurs.
Alvarz's framing of th issus providd a road asis for analysis in di-
vrs filds. His original artil has n itd thousands of tims in hundrds
of sintifi journals from iology and gology to mdiin and nular
physis. Th original hypothsis has bn strngthnd y furthr disovris,
most dramatially by dirt vidn of th impat providd y th Chixu.
lub ratr and its uniqu hmial signatur. Furthr, th thory has apturd
th popular imgination, inrasing intrst in sintifi invstigation and
promoting undrstanding of th unity of sin. 1arezhas himslf writtn
a popular aount of th disovry of th tratrrstrial impat and its im-
pliations for lif on arth in . Re.ld the rter of Doom (1,997).
B|LloGRAPHY
Th astrnspadfo
Alvarz, L..v.'.!(/.|varz,F. Asaro, and H. V. Mihi. 1980. xtratrrstrialaus notid inhaitantof t
for th rtaous-Trtiary tintion'Si208: 1095_1108. plosivly in th spri
Alvarz, \/. 1997. T. R d th rtr of Doon. Printon'NJ: Printn grow rapidly and mt
UnivrsityPrss. fd on insts nd ott
Alvarz, !.' . G. Kauffmann' F. Surlyk, L. !. Alvrz, F. Asaro, and H. V. ihl. intl sandy soil ttl wa
1984. Impatthoryof msstintionsand th invrtratfossil rord. is part of an old lina
Siee 22:,l',l' 5 -1'1'4 1'. -P.NI.P,
Stats,whlstadpol
dsrtin s littl as 10
Amphiians
Limd vrtbrats aros mor tan 37 5 million yars ago in th Dvonian Most salmandrs,
priod and thy ar gnrally alld amphiians. This was a vry divrs as- Although amphiian
smlag of aquati and trrstrial animals, ranging from th siz of a sala- ntr watr.
mandr to largr than an alligator. Som of thm vn ame limlss. But Frogs (Anura), wit
by th nd of th Palozoi priod and th grat Prmian tintion, aout tail, and wll dvlop
300 million yars ago' most of th linags Wr tint. A fw thrivd in th sudivisions. Many 1
arly sozoi priod for thy, too, disappard. Rnt amphiians and adults, thy ath pr
th amniots rprsnt th living dsndantsof this arly radiation. Thr ar toris, ut th most g
thr vry diffrnt kinds of living amphiians. Prhaps th most familia and in ponds in th spri
rtainly th most numrous and widsprad ar th frogs, whih ar found qualitis of th mal
on vry ontinnt pt Antartia and vn on a fw oani islands (s whih ar gnrallyl
figur). Th salamandrs ar wll known to inhaitants of th north rmprat tiv tadpol, a larva
zon, ut only on linage has ahivd any suss in th tropis. In ontrast' sason' during whih
th rlativly unfamiliar and srtiv ailians ar rstritd to tropial r- alga or vgttion.I
glons. Tadpols may liv m.
Rnt amphiians diffr dramatially fom ah othr in strutur and way any frogs hav no
of lif. All hv moist skins, howvr, and rspiration is largly utanous. that dvlop dirtly
mphibins 41
thirsynhrony,thy laoratd
pat had ausd a dst loud to
and making it old and dark for
'll land animals ovr 50 pounds
rnisms_and drew th infrn
lf a larg fration of spis 65
f dinosaurs.
a road asis fr analysis in di-
l thousandsof tims in hundrds
ology t mdiin and nular
ngthndy furthr disovis,
impat povidd y th hiu-
Furthr, th thory has apturd
;t in sintifi invstigation and
n.Alvaz has himslf writtn
tratrrstialimpat and its im-
l rtr of Doom (|997).
rillion yars ago in th Dvonian ost slmandrs, a fw fogs, and on ilin lak lungs ntiely.
ians. This was a vry divrs as- Although amphiians ar assoiatdwith moisr haitats' many spisnvr
nrwatr.
' ranging from th siz of a sala-
f thm vn am limlss. But Frogs (Anura), with mo than 5,500 spis,hav vry short odis. no
) grat Prmian tintion, aout tail, nlwll dvllpdlims, spiallylong hind lims tlrt hav for main
Wrxtint. A fw thrivd in th sudivisions.ay frogs ar apal of prodigious laps. arnivorous as
rppard.Rnt amphiians and adults,thy ath pry with projtil tongus. Fogs display divrs life his-
:sof this arly radiation. Thr ar toris'ut th most gnral nd familiar is on in wlrih rh ssongrgat
rns.Prhaps th most familia and in porrds in th spring, witlr fmals slting mals as mts asd on th
rd r th frogs, whih ar found qualitis of th ml all. Mating involvs xtrnal frtilization of ggs,
:vnon a fw oani islands (s whih ar gnrallv Iaid in lustrs lr strings. ggs dvlop into th distin_
inhaitantsof th nrth tmprat tiv tadpol, a lrval forrrr uniqu to rogs.Tdpols typially liv for on
sussin th tropis. In ontrast, sson'during whih tim thy onsum primary produtivity in th form of
ilians ar rstritdto tropial r- algaor vgtation'Howvr, thr r many varitions on this lif lristory.
adpolsmay liv more than on yar for nrtamorphosinginto froglts.
lffIaChothr in strutur and wa ny frogs hav nl larval stag at all ut ly larg ggs in small numrs
l rspiration is largly utanous. that dvlop dirtly into miniaturs of th adult. othr frogs hav tadpols
414 Amphibis
that may roodd in spial ompartmnts tht form in th skin of th of th skin and lads to dt
bak, in spializd pouhs on th ak, in th voal sas of mals, or in l i n s i n s p i sl i v i n g i n 1
divrs ologial settingssuh s aroral romliads.A fw frogs ar vivip. California, whr pidm
arous' with ggs dvloping in th fml rprodutiv trat' whr thy r- idly, and thr is no know
ivnourishmnt. pliatd in th dlins
Salamandrs (audata), with mor than 560 spis'rsmlanint am- fators inlud ffts of
phiians mor losly than do frogs or ailians. Thy ar gnralizd in tion from pstiids and
strutur' with a ody of modrat lngth, a wll-dvlopd tail, and two thse fators and othrs
pairs f lims, with th hind lims ing only a littl longr than th for- many parts of th plant
lims. ads of salamandrs ar rlativly smllr than thos of frogs, ut zlmnt ovr th dlins.
oth groups hav wll-dvlopdys and llnt vision. Salamandrsof- Today's living amphiia
tn gathr in ponds to rd ut mls hav no alls. Larva ar arnivorous jor group an trad a
rathr than hrivorous) and mtamorphosis is far lss dramati than in to dliat, thy wr
frogs. Som spis rmain in prmanntly larval or smilarval stat and trouling that amphibran
som lrva om vry larg. ost lads f salamandrshav som vari- should faing xtintlo
tion of th asi lif histoy, ut th most sussfulnd numrorrssalaman-
drs, in th family Plthodontida, hav dirt dvlopmnt' and it is only |BLIoGRAPHY
ths that hav sussfully invadd th tropis (rstritd to tropial Amr- Dullman,.W.., and L. Tru
ia). A fw spisof salamndrsar viviprous. Hill.
C a i l i a n s ( G y m n o p h i o n a ) a r l i m l s sa n d x t r m l y l o n g a t ,w i t h i - Huthins, N4.'!. . Duilm
lopdi.2nd d.V
thr a vry short tail or no tail. Th had is aout th sam diametr as th
Lnnoo, M. 2005.mphibi
strongly sgmntd trunk, and th ys ar inonspiutlus or invisil. A Spis.Brkly:Univ
uniqu snsory organ, th tntl, is formd from omponnts of th nos Stuart,S. N., J. S. hanson,J
.!7
and y and provids nvironmntal information to ths mainly urrow- Fishman,nd R. !(.
ing animals. Som ailins hav aquati larva ut m()st hav ithr di. xtintionsworldwid.j
.Wlls,
r t d V l o p m n to r a r v i v i p a r o u s .A m o n g t h m n y v i v i p o u ss p i s K. D. 207.be col
of hiagoPrss.
a r f o r m s t h a t a r l u s i v l y q u a t i . T h r a r a p p r o i m t l y 1 7 0 a -
ilian spis.
Amphibians play imprtant ut gnrally undrappriatd rols in
Andrson, dgar (18
osystms.Frogs ar important prdators as adults, ut thir larva an b
important onsumrs in equati systms.Rmoval of tadpls from strams dgar Andrson is rgard
an quikly ld to ovrgrowths f alga and forration of larg algal mats. tionary iology during th
Although salamandrs ar rypti and lss ovious than frogs, in som intlltual traditions, Anr
osystms thy an trmly numrous. As arnivors, thy ontriut in gntis and ytology t
to rgulation of trrstrialfood ws. Mmrs of th soil ommunity' a- systmatis.H is stkn
ilians ar major onsumrs of arthworms, ut thir ologial rol is lss natural sttings and for
wll undrstood than that of frogs and salamandrs. hyridization, a pross;
Amphiians hav rivd attntion ausof onrns aout vidnt d- hyridization and ak
lins and tintions lf populations as wll as spis throughout th l949 with th puliatio
world. A rnt assssmntof th sttus of amphiian spis aross th tio. Te ook was widl
glo found that a highr proportion (mor than on-third) Wr at risk of ontriution to plant gn
tintion than for any of th othr vrtrat taa. spially trouling volution.
Was th finding that a larg prntagof th dlins Wr from unknown Andrson, orn in For
.Whil
auss. most amphiian dlins doutlss ar rlatd dirtly to d m i n i s t t o ra n d a p i a n i
habitat dstrution nd modifiation y humans, th ar also othr implr- th }IihiganAgriultural
tant fators. On inftious disas,a hytrid fungus that ttaks th kratin Andrson was thr. Fro
Aderso 415
s that forrn in th skin of th of th skin and lads to dhydration, is rsponsilfor wll-doumnted
d-
th voai sas of mals, or in linsin spisliving in prottd ars in ntral Amia,
Australia, and
l m l i d sA. f w f r o g s a r v i v i p - alifonia, wh pidmis hav n rordd. Th disas
sprads rap-
'rodutivtrat, whr thy r- idly, and thr is no known Way to stop it. Gloal limat .h".'g.
lso is im-
pliatd in th dlins of many frogs in tropial montan
forsts. Othr
0 s p i sr' s m l a n i n t a t n - fators inlud ffets of introdud spis (.g., for
sport fishing), pollu-
:ilians.Thy ar gnralizd in tion from pstiids and frtilizrs, and synrgisti intrations
of som of
wll.dvlopdtail, and two ths fatos and othrs suh as ultraviolt adiation (UVB).
Howvr, in
ly rr littl lorrg than th for- many parts of th plant amphiians rmain aundant, adding
to the puz-
mallr than thos of frogs, but zlmntvr th d1rns'
tllntvision. Salmandrs of. Tday's living amphiians ar rmnants of a vry anint linag;
ah ma-
l t l i l s ' L v r r n i v < l r o u s jor group an trad ak out 200 million
yars. Altho,.gh ,.,y appar
;is is far lss dramti thn in to dliat, thy wr ruggd survivos of many *tin.tio.,
vnts. lt is
y larvirl r smilarval stat and troulingthat amphiians that lived happily with th long-tint
dinosaurs
l [ s a l r n d rhse v s o m v r i - should b faing xtintion on or wath'
;ssflnd nnroussalatnan-
t dvlopnt' and it is only BlLloGRAPHY
lis (rstritdto tropial Amr- .W.
Dullman, ., and L. Tru. 1986.Biolog of Amphibin's.Nw
York: Grw.
fous. Hill.
,nd xtrmly lorrgat,with i- Huthins,M., ). . Dullman,and N. Shlagr.20.Grzimek's
niml Li|
clopedi.2nd ed.o|. 6, phibins. Farmlngtonills, I;
a b o u tt h s a m d i a m t r a s t h Gl Group.
Lannoo,. 200.'. mphibi Dclines:Th onseruti-l Sttusof (Jitd Stts
inlrrspiuousor invisil. A Spis.Brkly:Univsityof alifonia Prss.
:d fror omponllts of th nos Stuart,S. N.' J. s. hanson,N. A. o' B. . Young,A. s. L. Rodrigus,
D. L.
ation to ths mainly urrow. Fishman'nd R. W. Y/allr.2004. Statusand1rndsof amphiin
dlinsand
larva but most hav ithr di- xtintionsworldwid.Sice6 1783-1786.
Wlls,K. D.207. h olog d Behuiorof mphibin.s'hiago:
n g ,r h n l r r yv i v i p a r o u s s p i s Unrvrsity
of hiagoPss.
r ar approximatly 170 a- D.B.w.
:ally Lrndrappritdrls in
dgar(ts97-|969)
Andrson,
s adults, ut thir larva an
movl of tadpols from strams dgar Andrson is rgardd as on of th lading sintists
of plant volu-
rd formtin of lag algal mats. tionry iology during th twntith ntury. A produt of
numr of divrs
ss ovious rharr frogs. in sonl intlltuaitraditions, Andson ws al to synthsiz nwr
dvlopmnts
Ls.As arnivors' thy ontriut in gntisand ytology to hlp a.ddrssmor traditional prolms
in plant
r r s [ t h s i l o m m u n i t y , a - systematis. H is st known for his invntiv studismasuring variation
-i't.og..,,'u.in
;, ut thir ologil rol is lss naturl settings and fo his artiulation of th onpt
of
mandrs. hyridizatin' a pross y whih nw gnti matrial is introdud
through
usof onrnsaout vidnt d- hybridization and akrossing. Muh of this work
was formalizd in
wll as spis throughout th 1949 with the publiation of smll volum tit\ed Itrogrssiu
bridiz-
of mphiian spis aross th tion. The ook was widly rd and immdiatly .og.'i,.d
as an original
than n-third) wr at risk of ontrition to plant gntis that ould shd light o' , g..'","l
thory of
:rattaa' spially toubling volution.
:hdiinsw fronr unknown Andson, on in Forstvill, Nw York, ws th son of
an duational
loutlssr rlatd dirtly to administrator and a pianist. His fathr am a prfssor
of dairy sin at
n n s .t h r r I s o o t h r i m p o r - thihigan Agriultual ollg and movd his family to
ast Lansing whn
id fungusthat ttirks tlr krtin Andson was thre. Fronr n arly g, Andsn ws fasintd
y plnts.
416 ndrson
dsrisas olltions of populations that display variation in many hara- in volving tolranto I
trs aross th spis'gographialrang. ain disussshow this variation t i a l r p < l d u r i vis o | t i
is promotd and maintaind in som ass y diffrnt sltion prssurs in mtal-ladn soil (Anton
diffrnt loations and in othrs y gnti drift. H argus that th uiquity was a striking dmnstr
of intraspifi variation of this typ oviats th lassial dfinitions of gn flow twn popt
speis; in fat, he argus that no singl dfinition of spis is adquat fo dnts prformd lgant
th normous natural divrsity of organisms. Cain disusss fiv typs of gntially divrs offsp
spies that rflt fiv diffrnt spis onpts' from th iologial spis in a spatially varial an
onpt hampiond by rnst ar (1942) to what ain alls agamospis, 7984). Whil thoris f<
olltions of similar individuals that rprodu asually. H also disusss suh an advantag,thr
mods of spis formation, offring on of th first attmpts to link diffrnt isly whih ologial l
typs of spis with diffrnt mods of spiation. Th finl haptr of th third, far.rahing pro;
ook is a prsint disussion of sympatri spiation, spiation without a ours of an inftious
gographial sparation twn forms. Th ovrall argumnt is that a attl of pathogn and h
spis is nithr a olltion of idntial individuals nor a fixd ntity' ut pathogns' hosts, and th
rathr a dynami st of populatins that ar wapoints on a sris of volu- tions aross th landsp
tionary rrajtoris. w o t h m s m g f
If th ook's mssag is obvious to th modrn radr, it is aus th strong for in natur t
book mad so prsuasiva as. ain's influn would tnd from oiogial physiologial strssof h
gntiists to systmatists. Svrl topis raisd in th ook, suh as th adaptatiln-mutations 1
adquay of th iologial spis onept or th inadquay of rtain mth- mon mong plnt sp
ods in systmati infrn for apturing th rlationships among sts of orhr agnts of slti
volving spis' would rmrg rpatdly ovr th nsuing yars. Although plants to prdatls and
ain's onis tratmnts of ths topis is oviously not informd y th molds diversity, on mus
walth of data now at our ommand' his tratmnts rmain among th most of sltion an vary and
luid and ritial disussions of many important qustions onfronting volu- mov radily. Th intr
tionary biologists who think aout what a spisis and how spisar r- mrgs in spa and, l
latd to on anothr. Ths thmes ar mr
tionary iology ar intr
B|LIoGRAPY In addition to ths r
thord a numr of high
ain, A. I. 1954. iml Spcisd Thir uollttio.Londn: Huthinson's
UnivrsityLibrary. portant topi, posd n
ayr, . 1942. Sstemticsd th rigi of Spis.Nw York: olumia dirtions. Ths paprs
UnivrsityPrss. -I,. (Antonovis 1976), te d
(Lokhart et aL. 7996),
among intrating spl
Antonovis, Janis (. 1942) ology has inflund rnnt
Whil th study of plnts has shapd many fats of volutionary iology. influn is visil throug
fw sintists hav usd plants to illuminat so mny important issus as has
Janis Antonovis. Antonovis has ld a sris of rsarh programs in natural BlLloGRAPHY
plant populations that hav unovrd som of th myriad ways in whih
^ I , ' . ' ^ - | ^, . l t 7 ,'- l r
natural seltion shaps th faturs of organisms nd the divrsity of thir ! r r . r l r . r
dynmis lf nthr-st
populations.
uirll. llurl of
Thr ontributions stand out among many. His arly work, prformd Alrtnovis' J. 1976. h n
.!0'als,
with his dotoral advisor at th Univrsity Collg of North Anthony B ltil Grd 6: 2'
Bradshaw, and ollagus,showd how populations of plants on min tailings, 2003. Towrd t-l
tonouis 419
isplayvariation in man hara- in volvingtolranto havy mtal dpositsin th soil, lrad also volvd par-
s ow this vrition
i n d i s u s s h til rprodutivisoltion frol losl adjantpopultions tht wr not on
y diffrntsltionprssurs in mtal-ladnsoil (Antonovis and Bradshaw l970; Antonovis 2006). This
rift' I-Iargus that th uiquity was a striking dmonstrationof inipint isolation in th fa of onsidra1
gn flow twn populations' In arrothrprojt, Antonovis and his stu.
lts th lassial dfinitions of
nitiorr of spis is adquat for dnts priormd lgant primnts to show that individuals that produd
rs. ain disusssfiv typs of gntilly divrs offspring through sual rprodution had gratr fitnss
pts' from th biologial spis in a spatiall,vvarial and ot-rrptitivnvironmnt (Antonovis and llstand
o w h a t a i n | l sa g a m o s p i s ' 1984). Whil thois for th origin of sxul rprodr.rtionhad postulatd
Luasxually.H also disusss suh an dvntag' thr was littl' if any, mpirial work that addrssd pr-
h first attmpts to link diffrnt islywhih ologial faturs of th nvironmnt wor"lldfailitat it. In a
iation. Th final haptr of th third, far-rahing projt' Antonovis and his studnts hav shown that th
spiation,spiation without a ours of an inftious dis:rsis dtrmind y a alan twn th loal
.h attlof pthognr-rdhost in an individual population and th movmnt of
ovrall 2lrgumnt is that a
pathogns' hosts, nd th gns for viuln nd rsistan among popula-
dividuls nor a fixd ntity, but
: waypoints on a sris of volu- tions arossth landsap('g.'Alxandr and Antonovis 1988).
Two th-lsmrg from Antonovis's work. First, rrtural sltion is a
odrn radr, it is baus th strongfor in natur that mrgsfrom a vrity f ologial ftors. h
n would xtnd from ologial physiolgialstrssof havy mtals in th soil may sm an ovious agnt of
:aisdin th ook, suh s th adptatioll-mutations that onfr tolrn of havy rtals r not om-
r th inadquayof rtain mth- mon among plant spis-ut Antonovis and his ollagus hav found
t h l t i o n s h i p s m o n g s t s o f othr agnts of sltin, from soil minral profil to rowding of othr
ovr th nsuingyars. Although plants to prdatos and pathogns. Sond, to undstnd how sltion
moldsdivrsity,on must undrstarrdth spatial sal n whih th prssurs
; oviously not informd y th
atmntsrmin among th most of sltionan vay and th spatial sl at whih individuals (or thi gns)
mov radily. Th intrplay f ths two sals dtrmins how divrsity
tant qustionsonfronting volu-
spisis and how spis ar - mrgsin spa nd, ultimatly, th as With whih spei:ltion ours.
Ths thms ar mrad y Antonovis's thsis that logy and volu-
tionaryiology ar intrtwind disiplins.
In additiln to ths rs:rrhprograms' Arrtonovis hs auth<lrdor oau-
thord a numr of highly influntial ssays' ah of whih xamind an im-
|tion, London: Huthinson's portant topi, posd nw qustions aout it, and inspird nw rsarh
dirtions.Ths paprs irrludd disussions of adptiv gnti varirtion
leies. Nw York: olumia
-1.r. (Antonovis |976I, th dynamis of inftious disassin natural populations
(Lokhart t al. ] 996), and th potntial for riproal gnti influns
mongintratirrgspis (Antonovis 200). is appro:rhto vlltionarybi-
ology has inflund innumral othr sintists for ovr forty yars and that
ry ftsof volutionary iology, irrflunis visil tlrroughout th disiplin.
so many important issus as has
s of sarhprograms in natural |BLIoGRAPHY
m of th myiad ways in whih
Alarrl,
. N,t.,ndJ. Arltlnovis.
198B.Dis:rssprdnd popul:rtion
ganismsand th divrsity of thir dynamisf nthr-smutinftionof Sillb ausdy th furrgusUstilgo
uioltl.
JourIof rllog76:9|-14'
nany. is arly work, prformd Antonovis,
J.1976. Th ntuf limitst ntualsltion.
ls of th issouri
.Wals, BotilGrd6: 224_247.
ollg of North Anthony
-. 2003.Towrdmmunitygnoris? olog84: 598-601.
ulations of plarrts on min tailings,
120 Arhaoptryx
Arcbeoptr
rhopteryr (anint ird, or Grman Uruogl) is th gnri naln givn
to 10 skltal spimns (as rvll as:r sondry flight fathr disovrdin
1860) of th arlist known ird' from ltst Jurassi lithogrphi limston
dsits kn.,vn as th Slnhofn Limston, loatd lust to th nrtl.i of th
ity of unih in southrn Grmny. Dating to approxinratly 150 rnillion
yars old, th dposits Wr in a hyprslin lilg()on that was llsdoff to
opn sa y orl rfs.Althlugh th disovryof th skltonsspnndsom
h most famous fossi
145 yars, th two most famous spimns ar th first two disovrd, th
disovrdarly in th
L o n d o r r n d B r l i n s p i r n sd
, i s o l ' r di n 1 8 6 l a n d 7 8 7 7 , r s p t i v i y . spinrn,''distlvrdi
Th last rovrd speimns ar th Solnhofn (th largst at 1096 largr
t h a n t h L o n d o n ) , d i s o v r di n a o l l t i o n i n 1 9 8 7 , a n d t h f i n a l s p i -
mn) th Thrmopolis spimn' dsribd in 2005. Althtrgh onsidral
vaition xists, th first svn spilr1nsar grlrallv rfrred tO as a sll.l-
g1 spis' Arhopter1, lithogrphic, w1|eth 199 ampl (Aktin- rPtilian fingrs with
Vrin) is distintiv in many faturs nd is rfrrd to as a nW spis' of modrn trunk-lim
Arhoptr bt,ri.[t is th onl1's1-limn tl show n ossifidstrnum' beoptrl ma,v hv
and th proportions of th legs ar qllit diffrnt from th othrs,lrly in. rchoptry also p
diating aroral haits. arly rtaous mir
Th lbrat]Brlin spimn (s figur) has n lld vrital to rvrsd as a lrallu
Rostta ston of volution aus it is omplt ard utifull1,prsrvd it w;rs lss t hom or
with outstrthd wings. It illustrats n almllst prft intrmdiat - bra had a pir of t
twn two major lasss lf vrtrats:irds and rptils. rhlptr sarum ar quit rpti
fulfills th Dwirrin xptation of :r missing link twn mjor vrt- ag l-riitd by rrro
brt groups. Its lliptial wings in profil ar similar to tl-rosof modrn irds. h skull lf r
woodland ird, with modrn flight fathrsvirtully idntil to thos of liv- tilian tth, ut with
ing irds' vn dlwn to hir tirostrutur' as shown y ltron mi- innr r wr qr.ritsi
rosopy. Fth sttltur' on ill'u'ntd,rrrrind r'rrlrngd in any ability.
major way for 150 million yars. Tl-rprimary and sondary flight fathrs Although rblp
ar nhord to th wing s in rnodrn irds, nd th flight fthrs hiit fossil irds hav n
asymnrtrivans that prodr"rdindivilual airfoils' indiating arldynami t o t h 1 9 7 0 s .M o s t o
fur-rtion.Mst palontologists liv t,atrheopterj' was a pal al- rsnt various typs
though primitiv powrd flir, and th sapula and orid ar primitiv boloris, whih sport
ut join at a 90" angl' as in mldrn irds. Th avian wings nd with thr muh lik tht o[ th l
Arhaoptry 121
ion in loslyadjrrtplant
ndar.Hrdit 25 49_62.
m n t x |\ 1 t l L l i os[ t h v o l u t i n r y
of th frquny-dpendnt
1996.Suallytrnsmittd disass
itins. BitiogiI Ruil'l,s lf th
'1. -rT
207:Qu
007.rhoptr1'
ARlsoL,s PHlLosoPHY oF sIN
h hindlimbfathrsin
Aristotl was al to aomplish what h did aus h had givn a grat
32 417_41.
l|-prsrvdArhtl!rspimn dal of thought to th natur of sintifi inquiry. ow dos on pogrss
,-1486. from th suprfiial and unorganizd stat of vryday prin towrd or-
disputsand
of irds:Disovris, ganlzedsintifi undrstanding? To answr this qustion' you nd onpt
issshft9\: 45547 \ . -A.F .
of th goal to hivd'and Aistotl dvlopdsuh a onpt in his Prior
d Postrior rzltis. Th goal of irrquiry, h argud, was systm of on-
ptsand propositions organizd hirarhially, ultimatly rsting on knowl_
dgof th ssntial naturs of th ojts of study and rtain othr nssary
riginator of th sintifi study of firstprinipls. Ths dfinitions and prinipls form the asis of ausal pla-
th thortialstudy of natur. His nationsof all th othr univsaltruths abrrtth sujtof study.
rs of imIs, and th Gr- Th sond ook of t Posterior Aaltis disusss how to ahiv this
;pializdstudis) provid a tho. gal. Plato had fornrr.rlatda famous paradox of inquiry in his dialogu
rr biologial invstigation and th o: either you know th oit f y<luinquiry, in whih as inquiry is
r Was nothing of si.rilarsop and unnssay' o you don't knw th ojt of your inquiry' in whih as in-
uy. In |837 t grat antomist quiryis impossil.Aristotl rminds us of this prdo in th first haptr of
rl Sinspran.from his [Aristo- thePostrior Anltics, ut his full ply is in ook II.
linrva from th Had of Jov, in a Thr h argus tht prptual xprin givs us a grasp of th targt of
)wn l 992.9l ). Bforxmining inquiry that dos not ount as knowldg, ut dos provid suffiint infor-
ls aut its rator ar in ordr. mation to dirt furthr inquiry. His xampls in th Postrior ltis are
familiarphnomna lik thundr' tipss,and sasonal shdding of lavs,
ut w an s his nrthod at work in his s'stmatiinquiris into th
hrn Agano;lst il1 84 ... His anatomyand dvlopmntof animls Portrd i Histor of Aimls (isto-
.ing Amyntas of Madon, and his ri imliur)' on the Prts of Aimtls (De prtibs imliun), and n
Lisland of uoa. H was snt at th Grtiott lf imls (De gertion imlium)' From hr on w
d in Plato's Aadmy for 20 yars, will rfrto ths y th shorthand H, P, and G.
had dvlopd his own distintiv
Lfor th study of natur. H joind a BloLoG|AL INoUlRY AND BIoLoG|AL KNoWLDG
t of Asia Minor' ut soon movd to
t Thophrastus' a yolrng rnan with For Aristotl, systmatis-th organization of irrformation aout animals-
n th trvo of thrrr thy originatd srrvsth goal of ausal planation, nd xplanation y rfrnto goals
o u t a s y s t m t i i n v s t i g a t i o no f a n - andfuntionsis fundamntal.Th following rmark, intoduing his system.
plants. ati prsntationof information aout :rnimals' strsssth first point: ..In
;totl to tutor his son Alxandr' but thfirst instan w should grasp th attriuts and th diffrns prsnt in
.With
w undr Alandr's ontrol. all animals. Aftr w hav dn this w should attmpt to disovr thir
)Wn as th Lyum' hadd th auss.This is th rratural way to arry out th invstigation, ginning with
lath in 323. With anti-Madonian inquiry (histori) out ah thing; for it oms apparnt from suh in-
l rtirdto his mothr's irthpla. quirisaut whih things and from whih tliings thr ought to dmon-
s t r a t i o n '(' | ' 6 . 4 9 | a 7 _ 1 4 \ ,
424 ristotl
.W. .!
gin with a study of th similaritis and diffrnsthat w osrv in hav spokn, th
the animal kingdom; only ftr this informrion is systmtiallygathrdand how many thy ar, .
organizd ar W in a position to pursu ausl dmlnstrations.H gins y anothr. Nt w nt
outlining kinds of similarity (in form' in kind, y analogy) that on firrds first off in loodd r
among animals and thn prsnts te for ctgorisof differen h will us: loodlss that th gr
in parts, ativitis, ways of lif, and hartr. This disussion pr<lvidshis Now all that ar f
framwork: ooks I-IV disuss similaritis and diffrns among parts' windpip, positiond
V-VIII among tivitis and ways of lif, nd I among trits of hratr. too in all of th fu.
Among parts, Aristotl distinguishs twn uniform and nonuniform parts they [eah of th krn
(tissusand organs, as w would say) and twn trnal and intrnal parts, parts. Put gnrally,:
.,gratkinds.''
moving systmatially through thsatgorisin nin groups of xhal hav a lung, v
Th ..loodd kinds'' (orrsponding to our vrtrats)ar irds, fish, sophagus and wind1
taans' four-lggd liv-aring animals, and flur-lggd gg-laying ani- as of th lung nit
mals; th ..loodlss''ar th hrd-shlld(tstaousmlllusks),soft-shlld larly. . . .
(rustaans),soft-odid(phalopods),and instdanimals (insts).Aris- Not all th lood
..division and orrlation'' t-lstalish rltionships of
totl uss a pross of do any othr animal
..kinds''
similarity and diffrn at various lvls of gnrality,using th aov 56a12)
as an id in this prss.H nots that many animls (inludinghuman ings)
..gratkinds.'' Taxonlmi tidinss is not his primary goal;
do not fall into his In his rviw of wht l
ths atgoris help in idntifying lli of systmati tlrrlations, and ths t r ad diffe r entit i l
will aid in th disovry of th ausl rlationships among animal diffrntia. bn disussd.H is no
Primary among ausal rlationships ar thos that xplain a part's diffrntil first lists th kinds tht l
attriuts y rfrnto th funtionl dmands <lfan animl's way of lif. izs to a oxtnsiv or
.!7hy
th Prts of AimIs presentsAristotl's attmpt to provid suh ausal organs and lung.
undrstanding.It gins with a dfnsof th propr mthods for iologial at that lvl; this o
study. It argus for th priority of tlologialxplanatin (planatilnsthat baring animals' and fot
idntify what a fatur is for) ovr matrial nd ffiintausal planation aus of th fish, whil
(what th featur is md from and how it is prtldtld)and for onditional ntwork of organs dos
cessit, whry matrials and prosss ar plaind y ing shown to This disussion is r
nssaryfor a goal. (Noti this rinforsth pririty of tlology.)This ity. In disussing th snl
viw of nssityand plantion prsuppossthat animals irr unifid om- ar th sam [in snaks
posits of bodily parts with funtional apaitis,whih Aristotl rfrsto as a n d n o w n s st h v i s
..Form'' in iologial ontts rfrsto th
thir matrial nd formal natus. pip is vry llng nd th
asi apaitisof nutrition, rprodution' l<ltlmltion, and ognition, whih (whals and dolphins),t
A r i s t o t l r e f r st o a s t h n i m l ' ss o u l . m a l s ( 5 0 5 3 0 )a n d d i s
This first ook also dfnds a mthod of multifatoril division, whry of animls ar in fous
many univrsal diffrntia r dividd in pralll in ordr to haratriz gnralizationswithout
orrlatd variations within kinds, and it also dfnds a mthod for stab- It is qually imprt
lishing th kinds that ar prsupplsdy this mthod. P II-IV attmpt oth usI pltions on
to stalish auss and to provid ausl plantins tht paralll th first ar givn fo thm. ri
four ooks of H. Th third of ths lrg wlrks is G, whih provids ar nssryand says n
ausl planations for th sujtmattr ognizdirr H V_vII. ths parts. The lgtl
essec' sl ctrl to r
N o w o n t r s tt h d i s
E MEHoD A WoRK
Th following quotation from initiatsAristotl's disussion of th intr- Not all animals hav u
nal organs of loodd animals: ruhicb tlte nk is nt
ristotle 425
:sAristotl'sdisussion of th intr- Nt all animals hav nk, ut only thos with th partsfor th sk of
hih the nk is turll prsnt-ths ar tlr windpip and th part
426 Aristotl
pfsntb ture for the sk al to know thir auss and ar y natur philosophrs. Surly it
animals draw in and xpl air would b unrasonal,vn asurd, for us to njoy studing liknsss
thos without a lung hav no of animals-ln grounds that w re at th sam tim studying th art,
sophagusis th part through suh as painting or sulptur, that mad thm-whil not prizing vn
o tht animals without nks mor th study of things onstitutd y natur' t last whn w n b-
it is ot |Issrto hav th hold thir auss.For this rason w should not hildishly disgustd
)notsnothing. And furthr, it at th amination of th lss valual nimals. For in ll ntural things
(t to th position of th mouth, thr is somthing mrvlous. (P I. 5, 645a7 =1'7)
r ndsfist to b somthing
lin two and through whih th on Frury 22, 1'882,driddn aid nar dath, harls Darwin rd
th lung m best aomplish thswords in nw translation snt to him as a gift y William ogl a
rgan onntd with rathing ..From quotations
month arlir.In a thank-you not to Ogl, Darwin wrot:
:ssr|or thr to b an soph- I had snI had a high ntion of Aristotl's mrits, ut I had not th most r-
Th sophgusis flshy, with a mot notion what a wondfr.rlman h was. l,innaus and uvir hav ben
lilat whn food is ingstd, yt my two gods, though in vry diffrnt ways, ut thy wr mr shool-boys
-rotdmagdwhn it is srapd ..Thank
to old Aristotl.'' Ogl rplid just thr days for Darwin did:
1-35;mphassaddd) .th parts of animals'. It gav m
you for your kind and ulogisti lttr r
muh plasur.I am glad also to hav addd a third prson to your gods and
rssity,pssiility (and impos-
jr to ontrast it with th disus- ompltdth Tinity.'' As a studnt of oth of thsgrat naturalists,ir givs
m grat satisfation to know that Aristotl affordd Darwin, during a p.
ur H passag.Hr th goal is
riod of grat suffring)som momnts of plasur.
haratrthy do pimarily -
y th oganism'sfuntional r-
at so imprssdth grat Frnh |BLIoGRAPHY
only systmatialldisussing th nglishtranslations an fundin Harvard
of all of Aristotl'sbilgialtratiss
i s a l s s h o w i n gt h a m a z i n g w y UnivrsityPrss'sL lassilLirar.In addition,for translations I
itutan organl sstem. rommnd:
Aristotl.1949.Prbr d Postriorltis..$7. D. Ross,d. Oford: larndon
Prss.
AimIiumI and De Grtionimlium
1992.Aristotl'sD l,rti|:ls
I. D. . Balm.trar.rs.Oford:(]lrndonPrss.
rolutionarynatur of what h was 2001'. th Prtslf imls. Books l-IV. J. G. Lnno, trirns.oford:
rviwing th rrors of pvious larndonPrss.
..h mlst asi rason
failurs: Forfurthrdisussionof Aristotl,siologilworks and thirsignifin' s:
ssdrspirationproprly is a lak A., and J. G. Lnrro,ds.1987.Philosophillssusi ristotl's
Gtthlf,
c a failur tO grasp what natur Bioklg,mridg:mridgUnivrsityPrss.
.W.,
p r s n ti n n - Kullmnn, Biologi.Stuttgart:Franz
and S. Follingr,eds.1'997.ristltlish
; w h t r s p i r a t i o ni s
Stinr'
sptingth parts suh as gills and
Lnno,J.G.2001. ristrltl'sPhilosophlf l|illog:Stttdisith rigis of Lf
quikly'' (47\24_29). Si.amridg:ambridgUnivrsityPrss.
ns without doing disstion and own,R. 1'992.h Hutri Ltrsi omprtiuett m. P. R. Sloan,d.
rking about funtion' is doomd to hiago:Universityof hiaglrss.
at on sk th funtional qustion P. 1986. ristotl'slssifitilof imls: tsiologd th lcptl
Pllgrin,
I.Jitlf tb Aristltliorps.Anthonv Prus'trans.Brkly:Univrsityf
k an vry unplasant. Nar th
liforniaPss.
rh study of lif, P I, Aristotl ad-
disussionof Drwin'sthoughtson Aristotlin:
Finally'thris a fasinating
Gotthlf, ^. |999. Darwin on Aristotl.Jourllf th Histor of tsiolog32:
ll to prption'th natur that 3_0. -1.G.r,.
linary plasursto thos who ar
428 Th rkstts rtil Tril
for th nt dade'
Th Arkansas ration Trial
Amrian sin-r
Th mdrn rationist movmnt dats from th puliation in 1 961 of The
|LloGRAPH
Genesis Flood, oauthod y ilial sholar John . lil/hitom and hy.
draulis nginring profssor Hnry M. Morris (Numrs 2006). This Gilky, L B. 1985.re
..young rth'' aount Minnapolis:Wins
of ration attratd supPort and grw in strngth
Gish, D. |973. ulluti
during th 1970s, thirnks to th poslttzingwork y formr iohmistDu- N u m r s 'R . L . 2 0 0 6 ' 7
an T. Gish, author of uoltio: Tbe Fossi/s Sy No/ (|97|' Sin th Desig.pnddr
Sops trir[,tlr lgal situation hd harrgd.No longr was it possil to x. Pnnok,R., nd N{.RL
lud volutiorrfrom sh<lols.Th prolm now Was to gt th Bil in. Nt- Qltestili th r,
urally, th rrionist supportrs wantd to gt thi idas taught in th Pronrtlrus.
Rus, N4.,d. \988,Bul
iology lasss of stat.supportd shools, so thy dvisd a variant-
retio/t,'oluti
ration sin-that supposdly mad no rfrnto th Bil and was .ihitomb, ., nd H
J.
asd on purly sientifividn,ut just so happned to support a young- d Its Sitln
ag arth' si days Of ration, with humns last, and somtim thraftra Pulishingompa
massiv worldwid flood.
In 1981 th stat of Arkansas passd a law mndating th baland trat.
Artifiial Lif pro
mnt in shool iology l:rsssof volutionary idas :rdrirtionsin'Im-
mdiatly, th Amrian ivil Lirtis Union (ALU) supportd a suit volution is tlr dfin
laiming tlrat th l:rw was unonstitutiorralirusit rahdth sparation whrvr it riry ot
of hurh nl st.lt.Th trial was hld in Littl Rok in Dm 1981, with ird pollinting a fl
ALU xprt witt-tsss inluding philosophr ihl Rus, palontologist uts of volr.rtion.Li
Stphn Jay Gould, and population gntiist Franiso J. Ayala. Th as and is bginning to 1
.W.alla
Was mad that ration sinis far from ing traditional hristianity, ut unravld thr
is rathr an idiosyllrati prvrsion of nintnth-nturyAmrian Prots- dtails. A rlativly n
tantism; that it is rlot gnuin sinut rligion drssdas sin;that th instans of voltrtio
mpirial fts nd thortial hypothssspak against it; and that it is n- \/hilmillions of li
tirly inppropriat tat iology lasssshould prsnt indiffrntlydiffrnt thy all tra ak to
positions simply usvrious mmrs of soit-vhav strongly held dif. only a singl t of li
frnt pt-lsitions. iology and our p
Aftr a tw<l-wktrial, Judg William R. Ovrton ruld firmly that th [w truly ompartiv i
was indd unonstitutiona1and should disardd.Th stt didd not rquir knowldg of
to appal th dision. In oming to his onlusion, Ovrton argud that si- tion. It is livdth
n follows rtin fixd ruls of mthodology-it appals to unrokn law, ut' unfortunatly, th
it opns itslf to omparison aginst th findings from th mpirial world In its ssn,th p
(falsifiation),ani it is always tntativin its onlusions-and that ration gneti variation and
sinfails on all of ths sors.Aftr th trial, sonrphilosophrs ojtd atristis of tlr popu
strnuously that rro ition of dmaration an proply sparat sin arth is th produt r
from rligiln. Altlrouglr tlris oasiond xitd disussion in pofssional howvr, th poss
irls, gnrirlthinking in th sineotntrlltnity'not to mntion tlr world Thr hirs n s
lf journalisrn :rnd puli opinion, sidd lvith ovrton and this smdtO may th produt
th nd of mttrs. irth to nw univrs
rationists thmslvsargud that th right way to pursu thir nds was vary among unlvrs
not through laws that ar oviously opn to attk' ut at th grassroots Darwinian fitnss,lt
lvl, prssuring shool oards and individul tahrs.This st th pattrn physis, whih govr
rtifiil Lif Progrts nd uolution 429
rat volution an inhait th yl an rpat svral tims ut may progrss to th volution of ..hypr-
t tlrt volutilnis th dfining pasits''that atually ttak parasits,stalingthir nrgy.
t n st l f . l i g i r e l v o l u t i < l nm a y Somtims,whn on kind of rtur ompltly d<lminatsth mmory'
amatillyalin lif. If w had suh that ll th raturs ar losly rltd, thy will volv kind of so-
rts,it likly wor"rld rborr- iality in whih irrdividul ratr'trsliving in isolation ar not bl <lrpro-
ommon with lif on arth. Ar- du, ut ratrsliving in groups ar bl to rprodu. Th raturs in
h volr'ltitlry pross itslf in this kind of digital world ar typially sxul in th sns tat thy do not
in than lrf on othr plants. mat to produ young ut rprodu individully, opying only thir own
ph,vsial/lrmial mdiunl; it is gnti matrial into thir offspring. Howvr, it was disovd that volu_
n w i n s . t t r . 'si v o l u r i o n a r tion ontinud vn whn mutirtions wr turnd off. This was eausth
n i v o l u t i o n( . g . 't h l a w s o f raturshad inventd a kind of primitiv suality in whih offspring wr
r ',htamounts t() anoth uni_ produd ontaining mitus of gnti matrial from mor than l par_
h logi of th omputr. Thy nt. It involvs a kd of s with th dad, in whih offspring inlud th g-
1omputis onstutel;thy nti mtrial from thir parnt, mid with som gns from raturs that
1 oprtingsystrn.Ths ruls havdid.
ir havior.Thus thy liv in a !hil th ologial ovolutionary dynami dsrid abov is th main
' l r i n t o t h t l i g i r a ln i d i t r m g i v s driving for for evlution' thr is also prptual sltion for ffiiny.
s and rvhat it dos. on form of ffiiny involvs rduing th amount f tim it taks to pro-
r r a t i n 8t,l i g i t l v o l u t i o n i s t > du an offspring. This is usually aomplishd y rduing th siz of th
..digital or-
rih hv nalld 8nomthat dsris th lrganism, thry rduing th tim that it takS to
lm On a Omputr with a Dar- mak a opy. owvr' som optimizations hav n ahivd through th
ing systrrrmanags a populatiorr volutionof mor ompl omputr ld.
a that w nr"Iatlrrs ar Th volution of mr omple od is a tantalizing ampl of th holy
nory inhaitd th progams. grail of Atifiial Lif. volution transformd simpl moluls into th om-
ldttsmutation y flipping bits pl nd bautiful lif foms tht w 6nd on arth today. It is th aility to
. h antlommutations
r i u r s gnratmplity that is th sour of volution's powr. It rmains an
itlr th rsult thertsom individLr_ unralizdgoal of Artifiil Lif to produ an tifiiirlsystm that hiits
wlrih lds to a natural pross opn-nddvolution' lading to vr mr omple artifiial lif forms, suh
ll rnditrm. as ourrdon ath.
l i n i n s t a n t l f v o l u t i o n , i t i s Lif appard on arth rughly 3.5 illion yars ago' ut rmaind in th
3anivolLrtionon arth. Prhaps form of singl-lldorganisms until aout 600 million yars ago. At tht
l o g n i s rs v o l v a d a p t t i o n st o point in tim, lif mad n arupt transformarion from simpl mirosopi,
:hir nvironmnt.A digital ol. singl-lldforms laking nrvous systms to l;rrg and omplx multilld
:ng flrof voltrtitlrr. forms with ompl anatomis, physiollgis, ologis, and nrvous systms
s l f - r p l i a t i ndgi g i a l o r g a n i s m , apal of oordinating sophistiatd havior. This transfomation o-
mmory lf th omputr with a urrdso auptly tht vlutirnaryiologists rfr to it s th ..amrian
thn a vry promillnt fatur of plosionof diversity.''
t sour of sltiv fors. Para- Somhav put forth a vision of a digital ntllr' a kind of iodivrsity r.
;irlly volv, arrd tlry st off an svfor digital organisms, prlrps distriud aross th Intrnt. Popl
to an ongoing sris of volution- ould ontriut som of thir mmory and CPU yls to th digital natur
prsrv'and digital orgnisms ould liv in t spa' migrating from om.
ron ots targt for xploita- putrto omputr' fding on unusd Inmory and CPU yls. Th ida is
Omommon' thy driv othr thatif a larg and omple rgiol of yerspa ould st asid for digital
rrio would gin with th volu- natur'thn a digital amrian plosion of divrsity and omplity might
-rof immunity y thir hosts. This ourthr.
42 Artificil Life Progrms d uoltil
Humans hav n managing th volution of othr spisfor tns f thou- Although th vast in
sands of yars through th domstiation of plants and animals. It forms th out of rah, mor mo
.W
asis of th agriultur that undrpins our ivilizations. manag volution approah has n to f
through rding' th ppliation of artifiial sltion to aptiv populatins. prforming omputatio
Similar approahs hav n dvlopd for working with volution in th plx omputations. Thr
digital domain. It forms th asis of th filds of gnti lgorithms and gn' algorithms and has
ti programming. Howvr, aus digital volution has not yt pssd to produ what appa
through its vrsion of th Camrian xplosion, thr xists th possiility to Studying organi v
..managing'' digital volution. .!nd
us a radially diffrnt approah to ours ovr vast sals
not limit ourslvs to using volution to produ superior versiRs of.qisting proah to th study of
softwar appliations. Rathr, w should llow volution to find th n ap- srv th pross of v
pliations for us. To s this pross mor larly' onsidr how w manag modst omplity. Als.
appliations through organi volution. allowing us to know tht
Som of th appliations providd y organi volution ar ri' orn'
what, arrots' f attl' dairy attl' pigs, hikns, dogs, ats' guppis, ot- lLIoGRAPHY
ton, mahogany' toao, mink' shp, silk moths, yast, and pniillin mold. Adami, . 1998.Itrrldu
If w had nvr nountrd any on of ths organisms' W would nvr hav G o l d b r gD ' . . 1 9 8 9 .
.W
thought of thm ithr. hav mad thm into appliations ausw r- Lrnig. Rading,N
ognizd th potntial in som organism that was spontanously gnratd Koza, J. R. 1992.Ctt
s of Nturl S
within an osystm of organisms volving frly y natural sltion.
l - n g t o nC' . G ' . d . l 9 q 5 .
any diffrnt kinds of things our within volution. Brding rlts to Lnski,R. ., . Ofri,R.
..ttr,''
volution within th spis, produing nw and differnt, possily of ompl fturs.]
forms of isting spis. Howvr, volution is also apal of gnrating Lvy, S. 1'992'Artifiil Li
spis. vn mor signifiantly, volutin is apal of ausing an plosiv Books.
inras in th omplxity of rpliators, through many ordrs of magnitud R a y , T . s . 1 9 9 1 .A n p p r
of omplity. Th amrin xplosion may hav gnatda mplity in. J. D. Farmr,and S. F
Worksh'ptl r!ifii
ras of ight ordrs of magnitud in a span of 40 million yars.Harnssing .!sly.
ths normously mor rativ proprtis of volution rquirs a ompltly 1994a.volutin
diffrnt approah. 239_26.
.W |994. An volu
know how to apply artifiial sltion to onvrt polr-quality wild
r. r' .L "d tr ir .r .r a t r| ri rL". r 4r '. t. i, f/ ,r L. li 6 1
orn into high-yild orn. Howvr' w do not know how to rd alga into
d.,rtifiil Life:
orn. Thr ar two ass to this inaility: (1) if all w know is alga, w Smolin, L. 1997. Th Lif
ould not nvision orn; and (2) vn if w know all aout orn' W do not
know how to guid th volution of alga alng th rout to orn. our p-
rin with managing volution onsists of guiding volution of spis
through vriations on isting thms. It dos not onsist of managing th
gnration of th thms thmslvs. Ayala, FranisoJ.
An altrnativ is to lt natural sltion do most of th work of dirting Franiso J. Ayala was l
volution and produing omplx softwar.This softwawill ..wild,'' liv. guishd mmr of th
ing fr in th digital iodivrsity rsrv. In ordr to rap th rwards and knowldgd for his stu
rat usful applitions, w will nd to domstiat som of th wild digi- sin.H has bn D
tal organisms, muh as our anstors gan domstiatingth anstors of vrsity of alifornia at
dogs and orn thousands of yars ago. U n i v r s i yo f l i f o r n i
This vision of digital natur is urrntly unattainal, and may always r- Ayala's arr has ot
main so. Still, mny sarhrsin th fild of Artifiil Lif ar still working xpansion of molular
toward an opn-nddvolution in th digital mdium. has n a ladr in otl
l 4
tlrrspisfor tns of thor-r- Although th vst inrassin omplxity nvisiond for digital nture ar
rnts and animals. It forms th out of rtrh'mo[ modst inrassin on,rplxityhav n ahivd. on
.$
zations. manag volution approah has n to fd numrs to digitl organisms nd rward thm for
,tionto ptiv popLrltions. prfrming omputations on thm; gratr rwards ar givn for mor om-
working with volution in th pl omputations. This pproalr has ld to the volution of quit ompl
f gntialgorithms and gn- algorithms ar-rdhas n usd as a dtnonstrtionof th aility of volutirl
voltrtionhas not yt passd ..irrduily''ompl stuturs.
t produ wht appar to b
, thrxists th possibility to Studying organi volution an frustrating aus it is a proess that
.W
rg'' digital volution. nd ours ovr vast sals of tim and sp. Artifiial Lif is an xiting ap-
: supriorvrsionsof xisting poah to th stlldy volution aus it provids an opportunity to o-
l volution to find th nw ap- srv th pross of evo'lution in ation, gnrating novlty, ologis, and
rrly, onsidr how w manag modstonrplity.Also,.it roadns our prsptivon lif and volution y
allowing us to know th volutionary pross in a nonorgani mdium.
ani volution ar ri, orn,
. k n sd. o g s , r s .g u p p i s , o t - BIBLIoGRAPY
thsoyast,nd pniillin mold. Adami, . 1998,ltrodutioto rtifiil Llle. Nw Yok: Springr-Vrlag'
rganisms,w would nvr hav Goldrg,D. . 1989. GtiIgorithmsi Serh,optimiztio, nd hine
o appliationsausw r- Lerittg.Radirrg,A: Addison-Wsly.
Kza' J. R' 1992. GetiPrr-lgrmmig: th Progrlnmigof omputrsb
was spontanouslygnratd
s of Nturl Seltit'l. amridg'A: tr4ITPrss.
ly v naturl sltion. Lngton,. G., d. |995, rtifiilLif:An t,eruie. mridg,A: IT Press.
t volutiorr'Bding rlats to l-nski,R' I].,. Ofri, R. T. Pnnok'nd . Adami. 2003.Th volutionaryorigin
..ttr,,'
nd diffrnt,possily of omplfaturs.Ntur 423: 139_144.
l i s . r l s o p a l o f g n r a t i n g Lvy,S. 1992.rti|t:ilLif: Th Qust for Ne rtio.Nw York: Pnthon
Books.
:apalof using an xplosiv
Ray,T. S. 1991'An approahto th synthsisof lif.In C. G. Langton'. Taylor,
ugh many ordrs of magnitud
J. D. Farmr,and S. Rasrussn,ds.'rtifiil Life II: Procdingsof the
ravgnratda omplity in- Workshlpo ArtifiilLife' vr>|' l1, 371_408.Rdwoodity, A: Addison-
f 4 0 m i l l i o n a r s .H a n s s i n g Wsl.
evolutionrquirs ompltly 1994a.volution' onrplity'ntropy'nd rtifiialrality.PhsiD 75:
2.9_26.
1994.An volutionrry approhto synrhtiioiogy:Zen and th art of
r to onvrt poor-quality wild
rating\if,rtifiil Lii 1, no. |l2: 179-209. Rprintdin . G. Langtn'
t krrow how to rd alga into d.,ArtifiilLii: u,i,|79-209. mridg,MA: IT Prss,1995.
1) if all w know is alg, w Smolin,L' 1997.TheI'ifof t orls.N,York: Oford Univrsityo'..,-'.*.
now all about orn, w do not
ng th rout to orn. Our p-
t guiding volution of spis
s not onsist of mnaging th
Ayala,FranisoJ. (b. 1934)
l most of th work of dirting FranisoJ. Ayala was on in Madrid' Spain, on Marh 12, 1'94.A distin-
'his ..wild,'' liv.
softwarwill guishdmmr of th intrnationl sintifi ommunity, h has bn a.
ordr to rap th rwards and krrowldgdfor hls studis in volurionary iologv and th philosophy of
mstiatsom of th wild digi- sin.H has bn Donald Bren Profssor of Biologial Sins at th Uni-
domstitirrgth nstors of vrsity of alifornia at Irvin sin 1987 nd Univrsity Profssor of th
Univrsityof alifornia sin 2003'
rattainal,and may always r- Ayala,s arr has oinidd with and' to a larg xtnt' has ld th ris and
f Artiliial [,if r still woking pansion of molr.rlrvolution nd th philosophy of iology. Indd, h
l mdium. hs n :r ladr in oth filds for many yars. Aftr his arly invstigtions of
434 l
variationin volution, pr- th An-razonasin anl tlrroughout th Caibban islands. dsignd labo-
was a gradutstudnt,h ratry xprimrrts t masuring tlr fftth amount of food, livirrg sp,
v<llutionmad possil a and lthernvironnrntaIfators hd on population fitnss.Rsults orrfirmd
is arr,Ayala usd mol- that natural sltion Was th driving fato imparing gnti polymor-
and population strutur of phisnr. H dvlopd Iaoratory mthods for masuriIlg dnsity.dpndnt
rsits.h impat of Ayala's and fquny-dpndntntural sltion.
..Biology In |97|, yIa and Dozlrnsky movl to th Dptmnt of Gntis at
in rnstwith his
..Tlologial tlr Univrsity of lifrnia at Davis. At UC Davis, Ayala ollaoratd with
1968)and his
of Sice,1970). palontologistJams \W. Valntin irr sris of invstigationspublislrd in
d i d dt o s r u d p h y s i s a t th 1970s that involvd divrs organisms, inluding tropial rid,n
p h i l o s o p h i al n d r l i g i o u s lams; Antarti, tmprtl, and tropial oani shrimp; and dp-oan
l t h o l o g y t t h U n i v s i t y starfish,sa urhins, and rahiopods. Tlr invstigationsdmonstratdthat
volutionist,orning to th th amlunt l gnti vriarlrty 1pnddnot On th abundan of r-
tizen in 1971. sours'ut on nvironmntal staility, whih allows gnti variants to b-
hrs, gntiistsFrnand om adaptd to diffrnt nvirnmtal fatursand rsours.Th rsults
lolution from a gnti pr- promptd nw hypothssto plain th major tintions osrvd in th
ssian-ornAmrian gn- fossil rord.
nl96i nd l964. Duing Duing th 1980s' rvorkirrgt U Dvis and latr at th Univrsity of al-
t i . rh a r r i d o u r i n v s t i g - iforni t lrvin, Ay:rl studid TrpLl'Iosomruzi, th agnt of hgas dis-
ian Drosophil ad disov- as,whih is a svr disas ndnri in many South Amrian ountris.
iation.H introdud nw H showd that it rprodus lonally, rathr than sually, as gnrally
monstratdthat th rate of thougl-rt at th tim. Ayale and ollaor;rtorihl Tiayrn formultd th
liatiorr'Ths studis wr lonal tlrory of parasiti protozoa. hy proposd that th population stru-
ii,G tis, L'olutio, tur of thsparilsits is prvailinglv lonal, vn though ths prvsiv hu-
m11sourgsrtain th apity for sxual rprodution.
ldud ltrophortith- or rntly, Ayala has ronstrutd th volution of Plsmodium fIi-
Dozhansky's laboratory at pru|' th gnt of malignant malaria' and rlatd spis.H has shown
,rofssorthr in 1967 'Wlt|t that th world pansion of P. flipltzzthroughout th tropial rgions is
hd th arlir prolms of rl1t'starting from :r singl Afian propagul rrrund 5,000 yrs ago, and
1 how this impatd th rat tlrat th spisis grrtiallylargly uniform, pt for mutations rntly
' i n v s r i g t i o no f t h g n t i volvdin rsponsto th human imt-nunsystm and to mdiins.
nd th volution of rpro- In addition to his ssayson iology as an autonomous sin,th onpt
ator.vxprimnts with th of iologial progrss' and th id of tlology, Ayala has xplord th dis-
'phil illistoi and rlatd tintivharatistisof th sintifirrrthod.H has m:rdothr ontiu-
firstomparativ dta ana- tions to th philosophy of sin,inluding xplorarions of how th pross
onomi lvl in Drosophil of r-raturalsltionprovids a ausl planation of tlr volution and dsign
of living forms. \/ithtnsivknowldg of th history of iology and, par-
onfimd thortial prdi. tiuIirrly,volutionry thoris, h has ontributd tnsivly to th Popu-
:anof gntivariaility in larization of volutitlnry thory as th ntral or of biology. H has
volutionaryiology was his forfullyargud thirt tlr tlror'l'of l,olution, as wll as sin in gnal,
i polymorphism in natural ndnot in orrtrzrditionwith rligious lifs.
La lading partiipant in th Ayala hs tndd Dozhansky's rnownd phras ..nothing in iology
rmorphisms wr adaptivly maks sns pt in the light of volution'' with th phrase ..nothing in
fofully for th lading rol volutionmaks snsxpt in th ligt of gntis.''
tsiv'ut gographially Frniso J. Ayala is th author of ro thn 900 spializd artils as
populations of Drosophil in wll s th author or ditor of mlr thn 20 ooks. is a mmr f th
46 Al
al populationsof DroslphiI
Atually, Hilair Bllo, who wrot thsamusing lins, Was svralnturis
ierrt groaP, uolutk |9:
out of dat. iros wr first sn in 1674 Antoni van Luwnhok,
,rs ontrolling ppulation growth loking at srapings from his tth through simpl mirosop. Thy wr
b mric Nturlist 1:. laldbtri in 1828 y th Grman iologist hristian Gottfrid hrn-
..small stik.'' In susquntyars' biol-
brg,from th Grk Word maning
:ntsin th produtivity nd siz of ogists disovrd th normous divrsity of atria. Th struggl to
and Drosophil birhii. Getis
undrstandthat divrsity, its volutionary history, and th ontinud rapid
meri Sitist5 6: 207 -22|.
volution of atria is on of volutionary iology's most hallnging and
.ionary iolog. Philosl1lh of iting aras of study.
Batria ar rlativly simpl, on-lld organisms' lald prokaryots'
innapolis,MN: Fortrss Prss. maning that thy do not hav a spial ompartmnt' or nulus, in whih
.lo. Washington, DC: Jsph
th gntimatrial is housd. Thr ar two typs of gnti information in a
atrialll. First, th is a small, usually irular hromosom within th
Sstmtisd th rigin of
.Washington, ll that primarily arris th gns rsponsil fo uilding th ll and its
,. D: National
omponnts.Sond, thr ar a numr of gns floating in th ll that ar
' 1998. uoluti-lrd olulr primarily involvd with rgulating th ll's mtaoli prosss. Batria d
:tio. atican ity Stat/Brkly, not hav mor sophistiatdll parts (organlls)suh as mitohondria (usd
.Thology and th Natural
-4,8, for powr gnration)or hloroplasts (usd' .g.' y ukaryots-organisms
with lls that do hav nulei-for phtosynthsis). Thy do hv riosoms,
organllsusd for uilding protins frm th instrutions givn y RNA.
hy rprodu y simpl ll division; on ll divids into two daughtr lls
and ah daughtr ll is a lonal rprodution of its rnothr ll.
Lik all organisms' atria ar opn to volution through natural sl-
tio: lls with advantagous faturs prolifrat fastr than thos without
thm.Advantagous faturs aris in atrial lls through th avnus.
First,th gns of atria hang spontanously,or mutat, and lonal di-
vrsityan aumulat rapidly. Sond' som atria ar apal of a kind
of sxual rprodution through th hang, twn lls' of som of th
gntimatrial on thir hromsoms. This gn hang plas mutants
thathav arisn in on l1lin into othr ll lins. Third, atrial gns that
47
48 Btril ullut
] transfrrddirtly from on onntrtions of oygn. Thy found thir nergy y nonoygn mans; on
gntransfr,an failitat th of th most ommon mans was y frmntation. Of ours, for th first half
olony and hang th hara- of th arth's xistn' thr wr no signifiant onntrations of oygn
yotsar simplr than ukary- availal.As th arth sustaind hmial hangs, oygn lvls gan to
.volvd flrth lattr, and ris and sltion prSsurtoward oxygn-tolrant (arotolrant)organisms
nldth formr. his assump- gan. Nt, oygn-tolrant organisms gan to us oxygn to prod n-
th natur of ll funtion, and rgy. Th first vid.oJ th faultativ aroi organisms-organisms that
t molular lvl. Th arth is ould swith frm on form of nrgy prodution (without oxygn) to an.
y yars th surfa was far too othr (usinglxygn)-an trad to aout 2.5 illion yars ago. Oygn
tssii lif dat ak to around lvls ontiriud to ris until thy wr aolt 1o of today's prsnt lvl of
Farsthr is solid vidn of 20%. Aout 2.25 illion yars ago' xlusivly oygn-usingorganisms (ar-
known as lu-grn alga. os)appard.
omatolits and livd in th sa ukaryots appard shortly aftr th mrgn of oligat aros. It is
L nssyondition of lif on now livd, thanks to th rilliant hypothsis of Amrian iologist Lynn
form still flourish today. Margulis, that ukaryots Wr formd through symioti intrations
\mrian miroiologist arl among various prokaryots. Som miroorganisms, prsumaly arhaalik,
nmon anstor-th parnt of asrd othr miroorganisms, som of whih wr atria. \Whil th
.Wos'
3.5 illion yars ago. DNA on th nular hromosoms of ukaryots indiat thir affinity with
think in trms of just a singl arhaa' it appars that th mitohondria-th powr plants of ukaryoti
trof lif is rally a pitur of lls-ar dsndd from th sugroup protoatria. (sherihi oli is
ritan through ll division today'swll-known protoatria; it livs in th guts of all mammals, inlud-
)ant horizontal gn exhang ing humans.) Th hloroplasts of plants, thos llular struturs that drive
l fous f th volutionary pro- photosynthsis,also appar to hav a symioti origin. Thus ukaryots ar
ls and small lustrs of gns. truly himras, whih ar th orgnisms arrying gnti instrutions from
nti mthods wr not likly many diffrnt typs of anstors.
:r, and mor aurat, to think Th symbioti rlationships that formd th ukaryots hav hos in th
, all of whom would hav n omplx rlationships that modrn ukayots hav with modrn atria.
or aurat pitur of th tr Batriatoday ar ubiquitous and othr organisms annot liv without thm.
stounding findings is that th any plants dpnd on atria to xtrat th nitrogn from th air so that
|ps:th atria (or tru at- thy an us it thmslvs. Aiding this pross is th aim of many artifiial
nalld, Arhaatria). frtilizrs,and it is wll known that tday griulturalists ar working hard
atria and arhaa. N4ost im- to modify plants gntially so that, thanks to gns transfrrd in from a-
:yotsin th way thy rad and tria, thy an thmslvs rat usal nitrogn. ammls us many at-
l i s a p p r st o r h f i r s t m j o ria in thir stomahs and intstins, usually alld gut flora, to digst thir
l and th ommon anstor of food. Batria in ths plas also hav othr funtions, lik prvnting r-
..N{olular
in th main ssay tain disassor stopping various potntial allrgis.or gnrally,humans
n). ny rhaa ar apabl us atria for thir own nds, as in frmntation. Tody, atria ar vn
xampl, hot or salty). This had usdin laning up oil spills.
n of th arlist lif forms, s- Unfortunatly, it is oftn diffiult to liv with atria. Many disass of
if was found in vry hot' dp- animls(spiallyhumans) and of plants ar du to atria.Rvrting ak
., atria gan to los ths to history,tooth day is du primarily to four kinds of atria found in th
n rgaind. mouth:Streptoous snguis, S' sobrinus, S' mitis, and S. mt.s.Othr un.
s in th ways in whih nrgy plasantatrial disasesin humans ar lprosy (Hansn's disas),ausd
Lnb dtrmind from molu- cobterium lepre; whooping ough, ausd Bordtll prtussis;
l rord, th first atri wr holra,ausd Vibrio holor; and turulosis' ausd cob-
uld not surviv in signifiant trium tubrlosis. Rntly, ppti ulrs wr found to ausd y
440 Br
Heliobctr plori. rtain intstinal disass, suh as Crohn's disas, a had n assumd to i
thought to th rsult of th ody's failur t rogniz its own atrial gut with fluid mittd .
symionts. his dirtor's pt dog
Batria ar known to rapid volvrs. Prft ampls of natural sl- ry with whih his n;
tion in ation, thy gntially put up arrirs against human prvntativ mryologial studi
masurs quikly. Antibiotis, spially pniillin, w th grat mdial t i k lgsgs h ih tt
disovris of th twntith ntury aus thy ntirly dstroyd.targtd 1 8 2 8 n d 1 8 3 7 .I n 1 t
batria without affting th human host. Unfortunatly' thanks to muta- drtook various kind
tion, high rats of rprodution, and horizontal gn transfr, atria d- Russian mpir. In t
vlop natural rsistans to antiiotis. For instan, almost as soon as though also a didd
.War
pniillin was introdud during !7orld II' it was found t'at Stphlo- Lik Gorgs uvi
coctts turswas dvloping gn-dirtd iohmial mthods of dfns. sparatd into fou
Th sam is tru of othr atria. Notoriously' vnral diss usd to urhins), th mollus
urd with just on shot of pniillin. Now it taks a oktail of drugs to and ras), nd th vr
omat it. It dos not hlp that humans do many things that nourgrsis- pitulation thory-th
tan dvlopmnt' suh as whn prostitutskp thmslvson ontinuous' through morphologi:
rlativly low doss of antiiotis. Batria sm to find this a hallng, find. ganisms lowr in th h
ing ways to surviv and thriv. imal mplifid fror
Th apaility of rapid volution in atria also prsnts a hallng for Urflrm of that parti
.W ..islrady at tJ
undrstanding thir arlist volutionary history. ar attmpting to r- srtd,
tra vnts that bgan at th dawn of lif, hanging rapidly and through man ftus, h hld, w.
mhanisms lik th horizontal transfr of gns with signaturs not asy to form of a gnralizd
tra. ]t4odrn advans in molular iology ar hanging our undrstand. and finally a partiul
ing, and th study of atrial volution is among th most hallnging and fronr th gnral to t}
iting topis in modrn volutionary iology. fo, nvr assumdtl
Dspit Bar's rj
|LIoGRAPHY
within a givn arh
Daon,l. Th icrobiI World. http://hlios.to.d.a.uk/t/miros/. arlir stagsof mr
Howland, J. 2000. The SurprisigArh.Nw York: oford UnivrsityPrss. attriutd to Louis A1
Univrsityof alifornia usum of alontolog'Itrodtilto tbe rh. tl mryos that h
http://www.ump.rkly.du/arhaa/arha.html/. -J.' d .R.
birds' or vry young
ths animls is quit
yos. But vn if th
Bar, Karl rnst von (1792_1876)
indiat anything; si
Karl rnst von Bar was prhaps th most minnt morphologist and mry- of irds, as wll s th
ologist of his day. H was orn in stonia to a Grman nole family. H at- mntal form'' (l828_
tndd th nw univrsity in Dorpat and ontinud mdial studis in Brlin Though h am
and Vinna. or intrstd in th thortial aspts of th sin, h pur- many Grmn morpl
sud postdotoral study at.!(iirzurg (1815), whr th rsidual influnof onrrtd him with t
Fridrih Shlling (-]"775_1854)ould still flt, transmittd thrugh th ratd y Shlling.Hr
gnial hands of Ignaz Dllingr (1770_184I). In 1817 h movd to Knigs- ampl, that th arh
rg to om assistant to Karl F-ridrih Burdah (1776-1847). Bar's guidd th ratur'
sor xprimntal approah to anatomy ld him to shd muh of his arlir ..t
mal,'' h dlard,
nthusiasm for a mor romnti and asthti intrprtation of natural ph. govrns its ntir dv
nomna. In 1826 h gan studying th fomation of th vrtrat mryo Bar ojtd to t
and initially onrnd himslf with th origin f th gg in mammals' whih rootd in Grman i
Ber 441
is'suh as rohn's disas,ar had rrassumd to idntifid with th Graafian follils of th ovirris or
o rognizits own trial gut with fluid mittd ,vtlr follils. Irrth sprir-rg of th nt yar h sarifid
his dirtor'spt dog nd loatd minut yllow ggs in its follils' a disov-
rftaInplsof natual sl- ry with ,|rilrhis.rrzrm would lrrrfotlr assoiatd.H ontirrud his
r s a g a i n s th u m n p r v n t a t i v mryolial studis, pulishing th two parts f his mastrwork, Die n-
niillin, w th grat mdial tiklgsgsllihtder hir (h dvlopmnralhistory of animals), in
thy ntirely dstroyd targtd 1828 and l837. In 18.34h ptd a hair in St. Ptrsurg,whr h un-
Unfortun:ltly, thnks to lnuta- dertllkvaious kinds of thropologial and thnographi studis for th
)ntal gn transfr, atria d- Russian rnpir' In his latr yars h am a modrat volutionist' al-
rr instan, alrnost as soon as thor"rgh also a didd opponnt of Darwinirr thory.
: lI, it was found that Stplo- l-ik Gorgsuvir (1769-1182), Bar hld that th anirnl kingdom ould
iohmilmthods of dfns. sparatdintl lur distiIrtarhtyps:th radiata (.g.,salrfish and s
.rsly,vnraldisasr'rsdto b urhins),th mollusa (.g.,lams and otopuss)'th artilata (.g.,insts
l it taks a oktil of drugs to nd ras),and th vtr.lta(.g.'fislr nd humarr bings).H dnid ra-
r n yt h i n g st h t n o u r a g r s i s - pitulation thory-th ida tlrat th mryos of mor mpl nimals passd
k pr h m s l v os n o n t i n u o u s ' through nrorphologial stagsorparal with thos of th adult forrns of or.
m to find this a hallng' find- ganisms lorvr in th hirarhy of lif. H maintaind that th mryo of an n-
imal xmplifid from th ginning of its gstation only th arhtyp or
:ria also prsntsa hallng for Urform of that partir"rlrorganisnl. ..Th mlrryo of th vrtrat,'' h as-
s t o r y .W a a t t m p t i n gt o r - s r t d.,. i sa l r a d ya t t l r g i n n i n ga v r t r a t '('1 8 2 8 _ 18 7 , 1 : 2 2 0 ) . S o a h u -
:, harrrging rapidly and thr<lugh man ftus, hld, w<>uld mov through stagsin whih it would tak on th
;nswith signaturs t-lotasy to forr of a gnralizdvrtrat'a gnrirlizdmamral, a gnralizd primat,
3y ar hanging our undrstand- and finally a partiula humn ing. Tlr frm of th growing ftus movd
among th most hallnging and fom th gnrl to th spifi. Th human mryo in its arly stags' thr_
'g. for,nvr assumdth mtu form of arr invrtrator of a fish.
Dspit Br,s rition lf strit rapitulation, h did allow tht spis
within a givn arhtyp displayd virtually i]ntialstruturs during th
.d.a.uk/to/miros/. arlirstgsof mryognsis.In a famus passagthat Darwin mistaknly
,York: oxfod UnivrsityPrss. ttriutdto Lois Agssiz (1s07-187)' Bar mntiond that h had two lit-
, ltrodutioto th rh' tl mryos that h hd flrgttn to lal: ..Thy might lizrds, small
raa.html/. -.I .T. nd .R.
birds' or vry young lrrmmIs.h fomation of th hds nd trunks in
thsanin.ralsis quit similar. Th trrnitisr not yt prsntin ths.m-
bryos.Brrtl,nif th wr in th fist stgsof dvlopmnt,th1.would not
indiatanything; sin th ft of lizards and mammals, th wings and ft
minnt mopholgist and mbry- of irds, s l,l,lls th l.randsand tto[ mn dvlop forrr th sm funda-
'o a Grman nl famil. H at- m n t a lf o r m ' ' ( 1 8 2 8 _ 18 3 7 ' I : 2 2 1 ) .
ntinud rdial studis in Brlin Though h m wrv of th romnti Nturpbillsophi hiitd y
al asptsof th sin,h pur- manv Grman morpl-rollgists, Bar rtaind rtain mtaphysial idas that
). whr th rsidual infln of onntdlrir .itlrtlr taditilnlf transndntalidlism tht was inaugu-
flt, transmittd through th ratdy Shlling.H smsnvr to hav aandond th onvition, for -
1 ) .I n 1 8 1 7 h m o v d t o K n i g s - anrpl,tlrt th :rrhtypof th organism, as kind of traphysial ntity'
:h Burdah (1776-1847). Bar's guidd th ratur's morphologial dvlopmnt. ..Th typ of vry ani-
d him to shd rrruh of his arlir ml,'' h dlard, ..tlth oms fixd in th rryo at th ginning and
:ti intrprtationof natural ph. g o v r n si t s n t i r d v l o p m n t ,(' 18 2 8 - 1 8 7 , 1 : 2 2 ) '
.mationof th vrtbrat mryo Bar ojtd to rlr transmuttionzrlthory th21tWas quikiy oming
;in of th gg in mmmals' whih rootdin Grmn iology. H undrstltldthat idas of rapitulation and of
442 Br
spis voltion gav sdutiv suor to on anothr) and h firmly op- flutuat in both p
posd thm oth: advan. N4orovr
swampd out. Bar
On gradually larnd to think of th diffrnt animal forms as dvlop'
plants appard sud
ing out of on anothr-and thn shortly to forgt that this mramor-
diatd, and tht in
phosis was only a mod of onption. Fortifid y th fat,tht in th
transitions within
oldst layrs of th arth no rmains from vrtrats Wr to found,
mammals-all arhi
naturalists livd thy ould prov that suh unfolding of th diffrnt
th prsnt tim wa
animal forms was historially groundd. Thy thn rlatd with om-
tntativly ut hop
plt sriousnssand in dtail how suh forms aros from on anothr. ,,for a tru undrst
Nothing was asir. A fish tht swam upon th iand wishd to go for a
i n t l l i g n ' ,( \ 8 7 6 '
walk, ut ould not us its fins. Th fins shrunk in radth from want of
volutionists and al
ris and grw in lngth. This wnt on through gnrations for a ou-
Bar.
pl of nturis. So it is no wondr that lut of fins ft hav finally
mrgd.(1828_1837, |: 200) |LIoGRAP1
Th ommunity of Grman zoologists livd along svral onptual fis- Bar' K. . von. 1828
surs. Som, lik th rdoutal Bar, rjtdth notion of rapitulation R|leio.2 vo|s
and th supportiv dotrin of spis dsnt. Gorg Hinrih Bronn \876. UbrI
(1800_1862)' Darwin's first Grman translator, thought that spis pro- Ntrissensh
grssivlyappard on th arth ovr vast priods of tim aording to a di- 1 1 8 8 6119 8 6
Oppnhimr, d
vin plan. H rognizd that mryoni dvlopmnt or a strong analogy Publiations.
to th morphologial dvlopmnt of spis;ut prhaps faring th ons. D a r w i n , . 1 8 5 9 .o
,l'96
quns of th rapitulational ida, h strssdth analogial haratr of oppnhimr,I.
th paralll and raffirmd th oundris of mryognsisthat Bar had amridg,:
Rihards, R. 1992.Th
stalishd. Othrs, lik th romanti Lornz kn (1779_1s51) and th
IdologilRo
asthtially drivn Gorg Gustav arus (1789_1869), unhsitatingly ad- Prss.
vand th thory of th rhtyp and its attndant notion of rpitula-
tion, whil th mrylogistsFridrih Tidmann (1781_1861) and Johann
Fridrih Mkl (1781*1833),undr th influnof Jan-BaptistLamark,
Bartholomw,
aptd th dotrin of spistransformation. Darwin adoptd rapitula-
Shmidt-Nils
tion thory dspit Br's striturs.H agrd that th mryo passd from
a gnralizd to a mor partiula morphologial stat. H simply hld that Although not mains
th phylognti anstor was of a gnral typ-for instan, that th vo- Knut Shmidt-Nils
lutionary prognitor of man was a gnralizdvrtratwhos form would tion. Bartholomw l
rapitulatd in th human mryo. ..Thus th mryo,'' Darwin o- pionrd th stud
srvd in the origin of Speies (1859, 38)' ..omsto lft as a sort of adaptations to sp
pitur, prsrvd by natur, f th anint and lss modifid ondition of thy dvlopd ask
ah animal.'' ampl' on of Shm
By th arly 1870s th volutinary thsis had swpt through urop' lav- watr and salt balar
ing only th most ralitrant in its wak. Bar undrstood tht th kind of among spis that
mpirial vidn Darwin itd, spially from iogography and palon- hmial proprtis
tology, pointd in only on dirtion, ut h would not ountnan Dar' varials. On of Ba
win's dvis of spis hang-han variation and natural sltion, rows living in salt n
whih ould not aount for th tlologial strutur of organisms. If Dar- a apaility not sha
winism wr orrt, intrmdiat forms should b produd y sltion, salt marshs. Th t
ut non wr found. han variation. whih th thor assums.should inspird, rval how
])rtoIomtad Sbmidt-NiIsn 44
alothr' and h firrly op- flutuat in botlr positiv and ngativ dirtions' thwartirrg volutionary
advan. Morovr, sltd traits would, through random mating,
swampd out. Bar rathr suppsed that th grat lasss of animals and
t nimal forms as dvlop-
plants appard suddnly in anint tims, as th gap-riddn fossil rord in-
forgt that this mtamor-
diatd, and that intrtral,tlologial laws lrad to dtrmin volutionary
fid ,vth fat that in th
trrsitionswithin ah of th largr groups of arrimals-fish' rptils, irds'
trtsw to found,
mmmals-all..arhing toward partiLllr gorlsand diminishing it ptlwr as
r unfo|dingof th diftrnt
th prsnt tim .wzlsapproahd. Br offrd his spifi hypothsis only
3y thn rlatd with orrr_
tntativly ut hopd that h had awknd in th radr rognition that
ts ros from on anothr. ..for a tru undrstanding of natur' W annot dispns with a govrning
h land wishd to go for a
i n t l l i g n ' (' 1 8 7 6 , 2 : 4 7 | . R m m b r i n g h i s a r l y w o r k i n m r y o l o g y ,
rk in radthfrom want f
volutionists and antivolutionists alik rvrd th rat Kl rnst von
lugh gnrationsfor a ou-
Bar.
rt of fins ft hav finally
|LIoGRAPHY
zrlongsvralonptl fis_ Bar,K. . von. 1828-1837' aikelugsgshiht dr Thir:Btlbcthtugund
th notion of rapitulation Re|leio.2 vols. Konigsrg:Borntrigr.
nt. Gorg Hinrih Bronn 1876. UbrDris Lhr.Inis St,lif dem Gbitdr
'r' thought that spis po- Ntrtuissshfte,2 vls.' 2: 235_480. St. Ptrsurg:Shmitzdorff.
[1886]1986.Autobiogrphof Dr. Krl rst uo Br.2nd d.J'
lds of tim aording to a di- Oppnhimr' d..;H' Shnidr' trans.Nlv York: SinHistory
pmt o a strong anlogy Publiations.
r.rtphapsfaring th ons_ Drwin,. 18.59. th rigin of Spis.Lorrdon:John urr1'.
c th analogial hartr of opnhimr,I.|967. sssi th istlr1,o| mbrologd Billlg.
:mbryognsisthat Bar had amridg,A: IT Prss.
-h
Rihards'R. 1992. ig lf uolutit:h orpbtlogilostrutild
okn (|779_1851) and th
lf Dri's Thlr.higo:Univrsit1'of hiago
IdellogilRrlstrutil
i9_1869|, unhsitatingly ad- Prss. -R./.R.
ndant notion of rapitula-
r n n ( 1 7 8 1 - 1 8 6 1 )a n d J o h a n n
,rof
Jan_BaptistLamark, Bartholm Gorge (. |91.9),and
. Darwin adoptd rapitula_
Shmidt-Nilsn,Knut (\9I5 -2007 )
tht th mryo passd from
:al stat.H simply hld that Although not mainstram volutionry iologists,Gorg Bartholomw nd
-._for instan, that th vo- Knut Shmidt-Nilsn Wr normously influntial for th study of adapta-
/rtratwhos form would tion. Bartholomw rrd Shmidt-Nilsn,with thir ollagusand studnts,
rs th mryo,'' Darwin o- pionrd th study of physiologial and iohmial traits of animals as
..oms lft as sort of adaptatins to spifi nvironmntal onditions. Th rsarh programs
to
rd lss modifid ondition of thy dvlopdskd how animals funrilrrirr hallngirrgsitr:arions.For -
ampl,on of Shrrridt_Nilsn,s projts zrskdhow dsrt rrinrlsmintain
l swptthough urop, lav- watr and salt alar-r.Ths typs of qustions ld to roadr omparisons
: undrstoodthat th kind of among spis that showd how wll diffrnsin physiolgial and bio-
m iogographyand palon- hmial proprtis wr mathd to diffns in partiular nr,ironmntal
would not ountnan Dar- varials.on of Btholomw'sproits :rmindhow popultiorrsof spar-
ation and naturl sltion, rows living in salt nrarshswr al to obtain watr y drinking sawatr'
ruturof orgnisn-rs.If Dar- apility not shilrd in populations of t sanrspisthat do llot liv in
ld produd y s|tion, salt marshs.Th umultiv rsults of thir work, along with th work thy
r th thory assums' should inspird,rval how wll th funtional svstmsof diffrnt spis'and vn
444 Bts
But iogograph rvas a triumph. Patiulariy important for Darwin was tlr liAll]t|D
Way that th inhaitants of islands ar so simila to th inhaitants of thir
losst ontinnts and not to th inhitants of lthr ontinnts farthr i'ALAYSIA
away. ..Th most striking and imprtnt fat for us in rgard tl th inhai-
sttGAPoRl
tnts of isl:rrrds,is thir ffinity to th<lsof th nsrrrrinlnd,rvitlrtrt-
ing atually th sanrspis.Nunlrous instansould givrrof this fat.
I will giv only tln, that of th Glapagls Arhiplago, situatd undr th
quator' twn500 nd 600 mils from th shosof South Amria. Hr
almost vry produt of th lnd and Watr rs h unmistakalstamp of
t h A m r i n o n t i n n t ' '( D a r w i n | 8 5 9 , 4 1 6 \ .
.s7all J
was likrvisvry snsitivto th signifinof iogography.His
first papr hinting t volution (in 1855) flusdon th ways in whih or- t 0 cs
ganisms sm to pttn thmslvsround th glo. H found that nwly Wi
arrivd organisms st-nalwys to onnet with livirrg organisms los to
thm...vr spis lras om into xistnoinidr-rtoth in spa arrd
t i m r v i t h a l o s l y a l l i d s p i s ' '( . W a l l a 1 8 5 5 [ 1 8 7 1 l , 2 5 ) . I n T h c e o -
t
N
i t i d i f l s
0 00Mi|es
FF
grphil Distributillt of ils (1'876)l/allaturnd to issus in io. 0 300Kilometers
gograplry,lolking oth at th spial issusthat suround islands as wll as 100B
1100
road qtrstionsof distiution. \Vll krrown is th sl-lldVallil's lin
(sfigrrr), brirk twntwo vr diffrrrtsts of organisr-ns, Asian and Walla,slin. Whn h w
lWalla rea|ized that thrt
Australasian, running hrough th Mlay Arhiplago, twn Borneo nd
Asia and Australasia.He <
Sulawsi, and twnBli (wst)and Lomok (ast).
in Th ogrphil Dist
A majlr qury for ths nintnth-nturyvolutionistsWas iltly how
a voagerwith Frdinnd
orgnisrs gt disps].Darwin spnt onsidrabl tinr running pri-
mnts trying to shorv how things lik sds anl slrils ould rvitlrstandslt
watr nd gt arrid y jtsam or y irds. H disussd his rvork tn-
sivly in te rigi. A liffrntapproeh was takn y Darwin's los frind, Aftr th synthti th
o t a n i s tJ o s p h D l t n H o o k r ( 1 8 5 ) . H a r g u d t h a t p l a n t d i s r r i u t i o n s tory of iogography w.
wr rtr xplaind r, on_istingland ridgs tlrt hav now disap- 1960s. Plat ttoni th
pard thnks to th ising sa lvls.This was prt of a thOry of Iimat, ontinntal drift. It now
proposd y goIogist harls l-ylt (1830-183), whih supposd that al. that had onrnd th n
though ontinntsannot mov sidwys' thy ar alrvayshanging shap - of lost ontinnts ut of
aus th arth's surf is forvr risirrgor flling. groups away from ah .
Btwn Darwin's tirn and th frlrrr-ringof tlr synthti thor in tlr th sam tim gology
1930s' iogographytook ak sat to palontology as most ffort was di- pothses.A lassixam
rtd toward th tring of arth'shistory. !7hn sltionthory am ak of Lstrosurs. This hr
into fashion' most attntion was peid to rirovolutionary usal issus' million yars go' is fat, r
prtiularl,vth ways irr r'',hihgns s;lrad and hrrgin popularirrs.Yt hav bn a world trav
thr ws som rvivd intrsti iogograplr,v. Partiularl implrtantWas found in th sam fossil
th influn of systratistand ornithologist rnst yr. In his lassi Sys- (Southast)Asia, and An
tmtis d th origill of Specis (\942) h disussdin dtail issus to do fat that, mor than 200
..rings thy wr part of Panga
with distriution' most notaly highlighting th irrrportantlf of
ras.'' Hr, on hs a rirrg of adjar-rt nd intrdingppulations of th gologial hypoths
..Th I
spis' lrut whr tlr nd points ar rprodutivly isolatd. Nlayr argud in th min ssay
that this is dfinitiv vidnof th formation of spisin a grdual rathr Th dvlopmnt of
thn a suddn pross. idas about how to r
Biogeogrph 449
0 300Miles
' [ 1 8 7 1 ] ,2 5 ) . I n T b G o . FF
turnd to issus in io- 0 300Kilometers AU$RAL|A
,120U
surround islands as wll as 1000
11 0
130! 1400
th so-alld \7alla's lin
!alla's
lin. Whn h was ollting spimnsin th Far ast' Alfrd Russl
ts o{ organisms, Asian and
!allarealizedthat thr was a lin of sparation twnt organisms of
lago, twn Borno and
Asia and Australasi.H disussdthis and othr iogographialphnomna
sast). tnTh Gogrphil Distributio lf nimls (1876). Th lin was first notd y
olutionistswas atly how a voyagrwith Frdinand agllan in 1521.
r l t i m r u n n i n g p i -
snails ould withstand salt
disussdhis work xtn-
ln y Darwin's lcs frind, Aftr th synthti thory was wll stalishd, th major vnt in th his-
udthtpln distriuions tory of iogography was th oming of plat ttoni thory in th arly
idgs that hav now disap- 1960s.Plat ttoni thory offrd gophysial xplanations for th ida of
r part of a thory of limat' ontinntaldrift. It now Was possil to addrss proprly th kinds of issus
3), whih supposd that l- thathad onrnd th nintnth-nturyvolutionists. It was not a qustion
:ahvayshanging shap - of lost ontinnts ut of ontinnts that mov sidways, arrying on-joind
tq groupsaway from ah othr. Biogography rivd a powrful tool and at
tlr synthti thory in th th sam tim gology rivd strong onfirmation for its audaious hy-
tology as most ffort was di- pothss.A lassi xampl of th two-way pross is givn y th distriution
n sltionthory am ak of Lstrosrs.This herivorous, mammal-lik reptil, found mor than 200
o v t l l u t i o n a 1 1a u s a l i s s u s . million yars ago' is fat, short, and squat. It is rtainly not a ast that wold
J hangin populations. Yt hav n a world travlr' rgularly swimming aross oans. Today, it is
,. Partiularly important was foundin th sam fossil dposits (Lowr Triassi) on th ontinnts of Afria,
nst Mayr. In his lassi Sys- (Southast) Asia, and Antartia. This would a miral if it wr not for th
sussdin dtail issus to do fatthat, more than 200 millin yars ago' all thr ontinnts tohd whn
.,rings of thywr part of Pangaa. Th fossils support th gologial hypothsis, and
.h importan of
trrdingpopulations of a thgologialhypothsis maks snsof th fossil distriutions (sFigur 27
:tivlyistllatd. ayr argud in th main ssay ..Th History of volutionary Thought,' y A4ihal Rus).
of spisin a gradual rathr Th dvlopmnt of plat ttoni thory oinidd with th advnt of
idsaout how to ronstfut spis phylognis using a st of xpliit
4s0 Biogogrphl
ruls, that is, idas aout ladistis. Th fusion lf ths onpts ld to th A mjor iogograpl
shool of viarian iogography, artiulatd in th 1981 volume Sstem- al illgist Rort
tics d Biogogrph: ldistis d Virie, ditd y Garth Nlson d w r d ( ) . W i l s o n 1| 9 t
and Nomn Platnik. Viariarr iogographrs, following Vnzulan lands in tlr s or oas(
Lon roizat, rrrphasizdth ir-nportanof sahingflr omnron iogo- iving and spis l
graphi pattrns mong unrlatd groups of organisms and dignlsingth funtion of suh fro
rol of gollgial vnts in rating thos pattns. For xampl, a lassi r h a r t l f t h i s | a n di
viarian analysis would idntify th ommon pattrns of rang limits and oIonizitln nd in
distriutions rf gnti variatitlrrmong s<-ruthstrn U.S. fish spis,pat- urv' a mathmatial
trns tied to oundaris t major rivrs' and diagns th formation of rivrs fourrd in lt lo:rti<ln inr
and th istn of old stuaris nd sa hnrrlss paramount vnts in SPis-lluvshlltl
braking on-ontiguous distiutions and ating prsnt-day pattrns. offrd th first quantiti
In ontrast, so-alld disprsal iogographrs argu that th disprsal of T h o l o g i |t h
spis aross road rangs and hangs in dispr:salrats aross major fa- usd th thory to rir
turs lik mountains or rivrs ar mor imprtant for prsnt-daypattrns. t h d i s t i u t i o no f s p
Th prlpr nswr is surly a omination of both viarin and dispr- a t i n 8a r . r g n l n r l
szrl.Drift and tlr susqr.rnt irrrirsar vry imptrrtant.Howvr, no on a u s l m h n i s m st
truly an say that ontinnt:rldrift is th only ky ftor in organi disprsals. of additionl ioggr
assiv disprsals hav ourrd twn Nortlr and South Amia as th rangS also tnd to hav
land links hav n stalishdtwn thm: th ..Grat Amrian Intr- Thr hav nsv
hang'' (rshall et a\. \982 s Figur 26 in t-hmain ssay ..Th History t h o y t l f M A t h u rl
of volutionary Thought'' y Nlilral Rus in this volum). This is th ra- S p k < l s k( i| 9 7 8 , 1 9 7 9 ,
son marsupils, suh as Opossllnls,xist in Notlr Ar-rrriaand r<ldntsp- argr.ringthat t rtin1
par in South Amria. (oviously, th golgil hngs hav tin part mpty r-rihsthat ar t
a funtion of drift; it is diffiult to kp th two fators apart.) rht mth th prdi
..Palontology
It is not orrly thanks to gology tht inrasing intrst is ing takn in and th
ausal fators in iogography.arly n, population gntiistsralizd that Th thlr of th ttl
gographial distriutions ar sigrrifiantin thi str-rdis. A lassi s was s t i k i n g x t r t s i o nI.n t l
th distriution irr Afria of sikl-ll nmia, a dissin whih th vitir and, on irlr islnd on
usually dis for rahing his or hr fifth birtdy. Why dos it tlur in i n i r lp o p u l t i O n x p e
rtain pats of th ontinnt trt not in othrs? In on dirtion, it was pos- d l i n l l n d t i n t i o nr
sil to pin th disas down tr th possssion of a rtain kind of gn- d i t i o n s . l t i s , i n f te,
homozygots for that gn show th diss-and it was shown that th gn prosss rvith volutio
.Why
auss th ollps f rd lood lls. But why th distriution? in th tions ol1stntlyo
Gold oast ut not in South Afria? Th answr am austh gnsin y Rort Riklsnd
singl doss (htrozygots)onfr a natural immunity to malari, and th this pt-tssin Wst lnt
mosquito is major pst in thos and only thos aras with th gn.In othr ogy and volution oul<
words, th distriution is linkd to sltionstting up arrirs against inf- T h s o n n t i O n s t
tion (Allison 1'954a,954). standing iogographi
\/ithth advnt of molular nrthods,population gntiistswr 1to t h t a n i m l I s i n r a i r
tnd si iogographil prinipls to amin larg-salpatrrns of pttn is vidnt in
gn distriutions in spa. Fron thspattl-ts, iologists hav n al to plnations not in sir
draw infrns aout th likly loations in whih a spis or group of h t I n t l r f | t i v | yl
spisoriginatd and th history of disprsalinto its prsntrang. Ths r- humidity, growing sas
onstrutions of history, alld phylogography, rprsntth modrn vr- Biogography ontin
sion of Darwirr's and !alla'spiorrringplratilr-rs. o n p u i l lu n d r p i n n
Bi-lgogrpb, 4'\ 1
rsrvs and ontinus tO play k rol in tlr onfirmatoy support tlf vo- h immdiat an
lutionary thory. that rsmld anima
holsts' Fathrs,whi
lLIoGRAPl.|Y
atually volvd rvitlr
Allison, A. . 1954a. Prottilt-t th sik|.]ltrait gainst sutrtinralriaI ovring in anir-nalssu
inftion. B rit i s h di l J rl r l 1,: 2.L)
.
loet pinnr sh
1954. Th distriutilrof th sikl-lltrait in l]st Afri:r arrd lswhr
heoptr. Thir pr
and its apprnt rlationship to th ini]nf subtrtianmli.
rnstios o| t Rrll,l ,\oit tli rtlpil cdil Hgict, 48: |2. would hav svd a
Avis, J. . 2000. Phlogeogrpl:y.mr.idg,MA: rvrd Univrsity Prss' t h i r i n s u l t o r yP r O p
Brgmnn, . 1847. Uer di Vrhiiltnissdr wdrmokonomi dr Thir zu ihrr rytially olord w
Gross. Gttigr Stdi.]. no. 1: 59-5_708. tion, or amouflag;s'
Darwin, . l8.59. () th Origi of Spal-..Londo: .)lhnlvlurv.
nough vidn of ol
Hookr, J. D. 185.]. Itrlduttt ss tl tb F|or f.Ntu Zelnd. l,ondot-t:
Lovll Rv. f a t h r so n t h f i n g r s
Lomolino, . V., B. R. Riddl, ltnd J. H. Blwn. 26.Biogeogrp.3rd d. gsts thi1tth fthrs
Sundrland, lr{A: Sinur Assoiats. posttl prsvdin s
Lyll, . 1830_] 83. Priipl,s tf Golog1,:Big- ttmpt to pli th I f t h p r s n o f I
Former hgs i th rth's Surf b Rfrc to ss Nola i
flight volv? rlttl
prtio.3 vols. l-ondon: Jhn urry.
i s t h 6 r s r a n i m l i r rt l
aAthrrr, R. .. r-r] . O. Wilsln. 1967. he h<t\'tf Isld Biogogrph1'.
Printon, N.}:Printon UrivrsitPrss. f o r m w i n g p l o
.!,
arshll, L. G., S. D. J. J. Spkoski Jr., nd D. . Rup. 1982. ammlin sit flight strok and u
vlution and tlr grat Amian intrhng, Silt2l5: 1.5\_157. faturs is indirt but
Mayr, . 1'942' Sstttisd th rigi tlf Spis, Nr,v York: (]olLrmi S i n t h l 8 8 0 s , w
Univrsity Prss.
hav n priodiall;
Nlson, G., and N. Platnik. 1981. Sstmtltisd Biogogrph: ldistis d
Viri.Nlv York: olLrmbi Univrsit Prss. and usd inipint win
Riklfs' R. ., and . Brminglrn. 2007, Th ussf vllutionrvrditins in Th sond is tht ir
arhiplagos:Pssritrirds in th l-ssrAntil|s. mrill Nturlist |69l. nt willgs to gin inr
285-297. p d r ( ) r s .V a r i o u s m t
Spkoski, J.J., J.. 1978' kinti modl of Phrrrzlitaxonomi divrsity.
nithr lrypothsisis tt
I. Analysis f nrirrinordrs. Plrlbi<ll<lg1,
4:22_251'.
|979. ^ kinti modl oi hrrrzoitaxot-tlrni divrsit. lI. arly
storsould lim r
Plozlifarilisand muItipl quiliria. Plo b i tllog 5 : 222-25 2. d o s n o t g U a a n t f l l
1984. A kinti modl of Phanrozi taxonomi divrsity.IIl. Post- trstrial ipds and 1
Plozoi familis and mss tintions. Plobilllgl,10: 246_2-67. liftoff?
.!(alla'
A. R. 18.5.5[l871]. Or-rtlr law lvhilr lras rgultdth introdution of nw Rnt rsahhas
spis. Als nd g:in of NtrI Histor 16: 184-196. Rprintd in
lution of th flight strt
ontributkls tll th Tlr of Nturl Sltio. Lndln: millan, 1_25.
|876. Th GogrphitlDistributitlll lf imls.2 vols. Llndon: movmnt ol, th for
Mrnillan. Deio,bus and VI
1905. 1' l-if: Rlrd lf uets nd o1liltions. Londn; hapman nd i n h r i t d :t h o n s
Hll. -I'. d .R.
o n s h d l n o d i f i dj t
thos prdatory dinos
Wr gly lollgtd
Birds
lik ird wings. Whn
Sin originating from small arnivorous dinosaurs in th Lat Jurassi whippd forward and
(aout 150 million yars go)' irds hv raditd into sm 9,600 spis gatd fthrs,dlivr
around th glo. Thir olors' songs' havios,and ologial adaptations th most frqurrtlyus
ar ndlssly olplx. What sorts of fators hav n ky in thir volu- Of urs, flight s
tionary radition? pod Di()hus \a
Birds 15
to row siz. Nw wok on th Gauthir, J. A., and L. F. Gall' ds.2001. Nl Perspetiues'lth rigin nd rl
arlybirdsshows that dinosaurs uolutil of Birds. Nw Havn, l.: Yl Univrsity Prss.
to th<lsof nrost nrmmals nd Padin, K., and L. . hiappe. 1'998a.Th lrigin and arly volution of birds.
Biologil Ruies (amridg) 73: 142'
ir.ds.suh s tl|'uiusornis.
|998. Th origin of irds and thir fltgt. Sitifi mri, Fruary:
sstags(juvniland suadult) 3847.
r rsult,irds fftivly am Prum' R. o.,4nd A. tl. Brush.2003. Whih am first, th fathr or th ird?
lould lrvhd a grtr ratio of ScintifiAmri,arh: 84-9. -K.P.
rovd wing loadirrg would hav
tion of spis, in th
std in th history of
rspt. First, Buffon's
dsnt, with rsml
This notion of spi
trms of tmporal on
Buffon pliitly adm
b modifid y th at
For instn, h prop
thir ounterparts in ,
should onsidrd
Daunon's anattlm
L'As (175) that th
rats' and h said th
ommon anstry.
Ths idas had a m
on his thr points' h
an indfinit transmut.
to a rtain point. Bu
quadrupds (200) to m
nlargd spis.But hl
spis wr rintrpr
largd spisould n
osrvd at th right l.
nnt' as Natur hsl
sidrd th hypothsis
( l 7 5 3 ) ' w h r h p o v i
world as dsnding fr
of hand for two rason
anothr spisand (2)
mannt lin of distintr
this point. ost of th
fat that h progrssiv
only what w would
to th lvl of a ..famil1
Pitursof a rhinorosand an lphant,... *'r", s Nturl istor.The tion etwn spis an
-;
many forms of animls that w apprnt y th ightnthntury fasinatd |n Lpoqus d I t
th savantsand ld thm to spultaut th rasonsfor th divrsity. tory of th living worl
proposd that th art
omt' had progrssiv
alls mad of diffrnt
gradual volution (in th sns of an irrvrsil) history of natur (Rogr 100'000 yars had
1988). In Tbelr of the rth (1749), Buffon ws mainly intrestd in th from an initial stat of
a t u a l a u s s 'w h i h , y t h i r O n s t a n ta t i o n ' a o u n t f r a n u m r o f g - anothr stimation: 3 r
ologial fats (.g., th trnal ation of watr, whih aounts flr major puts forward an asolt
aspts of gographi rlif). It was only aftr h gan to laorat his no- formation of sdimnt
Buffon 457
t i o n o f s p i s 'i n t h r i d - 1 7 5 0 s ,t h i r t B u f f o n g a n t o s r i o u s l y i n t r -
s t di n t h h i s t o r y o f n a t u r . T h r o f h i s i d s p l r y da m i o r r o l i r r t h i s
rspt.First' Butfon's onpt of spis was foundd on intrfrtility and
dsnt'with rsmlaning suordinatd to th ontinuity of linags.
This notion of spis suggstd a gnral vision of th unity of lif in
trms of tmporal ontinuity rathr th:rn in trms of trnal typs. Sond,
B u f f o n x p l i i t l y a d m i t t d t h t o t d o m s t i a t da n d w i l d s p e i e s o u l d
m o d i f i d y t h b a t i o n o f s v r l a u s s( . g . ,l r u r r r na t i o n o r l i m a t ) .
For instan, h p'roposd that th nimal spis of th old World had
thir ounterprts in Amria: th lattr dsndd from th formr, ut
should onsidrd today as distint spis.Third, on th asis of
Daunton's anaton]ial work on nimal skltons, Buffon argud in
L ' s e ( | 7 5 3 ) t l r a t t h r W a s a n r . r n d r l y i n gr - r r r i t 1 , otfy p o f r h v r t _
r2rts'and h said that this uniry of ryp suggstd th hypothsis of a
ommon anstry.
Tlrsidas had a massivimpat on ightnth-nturynatural historians.
on his thr points, howvr, Buffon did not rally adopt th hypothsis of
n indfinit transmutation of spis. Spies uld modifid' ut only
to rtain poirrt. Buffon proposd to rdu tlr numr ot spis of
qr"r:rdrupds (200) to mrly 32, intrprting th urrnt familis in trms of
nlirrgdspis.But h did not Want to go any furthr. ost of th urrnt
spieswr rintrprtdin trms of varitis' ut t limits of th nw n-
largd spisould not ovrstppd.Providd tat th naturl spisis
obsrvdat tlr riglrt lvl, thy ilr .,pduringniris,as anint, as pma_
nnt' as Natur hrslf'' (Scod Vue' 1765). Similrly, whil Butfon on.
sidrdth hypothsisof transmutation in his famous mmoir on th donky
(175),whr h providd an inrdily suggstivviw of th whol animal
wrld as desendingfom a ommon anstor' h rjtdthis hypthsisout
of hnd for two rasolls:(1)no true spisis known to hirv ppdfom
anothrspis:rnd (2) th infrtilitv lf hyrids provids dfinit nd pr-
manntlin of distirrtiontwnspis.Buffon nvr hangd his mind on
thls point. ost of th allgd Ontraditionsof his thinking rsult from th
fat that h progrssivlydoptd llotion of th spisthat inludd not
only what w would all today a ..spis''ut also vy possil taon r'rp
to th lvl of a ..ferrnily.'' But Br'rfforralways nraintaind an soltrtdistin-
Buffon'sI'Jturl i stor. T he tion twnspisnd varitis.
th ightnthntury fasinatd InEpoqus d l tur (7779), Buffon intgratd his idas aout th his-
asonsfor th divesity. tory of th living world into a gnral vision of th history of natur. H
proposd that th arth' aftr hving n gnrtd y a ollision with a
ot, hd progrssivly eom oolr' on th asis of xpimnts on
allsmad of diffrnt matrials (spiallyiron), lr alulatd tht at last
100,000 yars had n nssary frlr th arth to ool to its prsnt stat
:rsibl)history of natur (Rogr
from an initia1 stat of fusion. (In anothr manusript, h somtims givs
fon was mainly intrstd in th
anothrstimation: 3 million yars; s Rog l988.) In poqus, h also
L o n ' o u n tf r n u m r o f g -
puts forward an asolrrtgologil hronology on tl] asis of th stady
lat, whih aounts for major
formation of sdirnntary roks. p<tqusgav a massiv irpuls tl th
t r h s a nt o l r a t h i s n o -
458 Buffo
no dsntwith modi
a lly pr-evolutionist
a aa
aa . . .. Buffon's volutioni
aa a :ltion of spis witl
aa
a This is prisly w
rjtd,so many hist
tionist in th modrn
insight into Buffon's r
.tf.' a
aa a ta aa'
. .. BIBLIOGRAPH
a a
a
Br'rffon,G.-L. (Llrd
vols.Paris:Imprim
mntindin this
a
.... prtiuIir.|
a
Gyon,J., d' 1'992.Bu1
.. .. t. .... ... ...... .....3 .. .
.. d l mrt d Bffo
a G. anguilhm.Pr
I Lvjoy,A. o' 1959.Bu
I
HA|N nd W. L. Strauss J;
1749 1755 1761 17661767 1780 1782 I 789 Johnsopkins Pr
Rg'J. 1988.Intrldu
harls Darwin was far from th first to us th pitrrrof th tr of lif as ditionsdu trsL
rprsntation r mtphorfor th histoyf lif on erth.On rlirusr was Shnlitt.S. 2007.Bttff
Buffon. This diagm pl<ltsth frqunieswith whih he usd th tr mta-
phor and othr metaPhors'maps' and hins.Not that th us of th tr mta-
phor in fat dlind with tim' possily a funtion of th fat that Buffon am Burgss Shal
undr rligious prssurfor his toying with evlutionary spulations'and, as
Th fossil rOrd larg
on always ken t sty on th side of th authoritis,h modifid (or at least
onaled)his thinking aordingly. (Adaptd from G. Barsanti, ..Buffon et Alnrost invarialy, ll
l'imagd la natur,''in Gyon |992,290.I shlls) as wll as tral
ganisms imprintd on
S u h v i d n sa r s t i
idn also has its dis
volutionary viw of natur. This mmoir also put forward a hypothsis prsrval hard prt
aout th origin of th diffrnt animal and plant spis.Lif had appard fossils ar undiagnst
as a onsqun of th spontanous assoiation of organi moluls, tial tras an bui
whih thmslvs had rsultd from natural hmial rations whil th All, howvr, is not
trrstrialglo was ooling. Diffrnt assoiationsof diffrnt organi mol- wirrdow into th dist
uls had gnrtd th diffrnt living spis. From th onst' ths pisolslf fossiIizati
spis Wr as ompl as thy a now. As a funtion of th physial ir- parllld irrsightsint
umstans (i.',limat), spisapprd sussivly'ut thy did not d_ th potntial to rsolv
snd from ah othr. All wr spontanously gnratd. Buffo also tirrv dposits yilding
argud that th various spis migratd from on pla to anothr as a irnportant is th Burg
funtion of limati hang, and tht thy am tint whn th irum- notaly thos from so
stanswr no longr favoral. Thr ar svral r
This final spulation is haratristi of Buffon. On th one hand, his on- of intrst.In partiu
ptions aout th antiquity of th arth, its gadul hang,th influnof rian xplosiln,th st
physial onditins upon th history of lif, and xtintion look gnuinly of tlr arrirrr[s. Th x
volutionist.But his onptual shm for th arupt origin of sPis,With s k l t a l f o s s i l s n d t h
Burgss Shl 459
BIBLIOGRAPHY
Buffon,G.-L. ([,lrde).1749-1804.Histir turll,grlt prtilir.44
vols.Paris:ImprimriRoyl;Plassn.INot:All writingsy Buffon
mntiondin this ntry wr pulishdas part of Histoir turll,grlt
prtiulir.l
Gayon,.|.,d.|992. Buffo 1988,tesdu ctllloqitrtiol du bietir
d l mort d l3uffo,Pris-otbrd-Dijo.rfy . yr, postfay
G. nguilhm.Pis:Vrin.
Lovjoy,A. o. 19.'9.Buffonnd th prolmof spis. In B. Glass,o. Tmkin'
and \W.L. StraussJr., ds.,Forrurs
of Dri: 1745-18.'9'Baltimor:
.l
'1780 1787 1789 JohnsHokins Prss,84_] .
Rogr,J. 1988.Introdution. In Buffo:Ls poqusd l tur,i_lii.Pris:
hpiturof th tr of life s a ditions du Musum national d'istoir naturll.
lif on arth.On earlir usr was Shmitt'S. 2007' I]uf|o-(ut,rs. Paris: Gllimrd. -1.G.
ith whih h usd th tr mta-
. Not that th us of th tr mta-
ntion of th fat that Buffon am BurgssShal
volutionaryspulations,and, as Th fossil rord largly onsists of th rul and srapings of anint |if.
thoritis,h modifid (or at last
..Buffon t Aln-rostinvarialy, all that w find r roust pis of sklton (ons and
d from G. Barsanti,
shlls)as wll as traks, trails, and othr signs of ativity th on living or-
ganisms imprintd on th sdimnt nd ar now prsrvd as tra fossils.
Suh vidnsar still ntral pillrs in doumnting volution, ut suh v-
idnalso has its disadvantags.F.irst,many organisms, vn animals, lk
prsrvalhard parts. For ampl, onsidr th slug. Sond' many tra
ir lso put forward a hypothsis
fossilsar undiagnosti as tl th original typ of animal, and fftivlyidn-
d plant spis.Lif had appard
tial tras an uilt y unrlatd animals.
Lssoiationof organi moluls,
All, howvr, is not lost. This is auson oasion ntur throws opn a
ural hmial rations whil th
window into th distant past. Thls is possil on aount of ptional
oiatins of diffrnt organi mol-
g spis.From th onst' ths pisodsof fossilizatiln.Th rsultantsoft.odid fossils not oniy giv us un.
parallldinsights into th original divrsity of anint lif' but thy also hav
As a funtion of th physial ir-
th potntial to rsolv fundamntal prolms in volution' of th sdimn-
d sussivly'ut thy did not d-
tary dposits yilding xptionally prsrvd animals, argualy th most
tanously gnratd. Buffon also
importantis th BurgssShl (British olumia, anada) and its ongnrs'
l from on pla to nothr as a
notalythos from sluthwsthin and North Grnland.
l am xtint whn th irum-
Thr ar svral rasons why Burgss Shal_typfaunas ar suh a fous
of intrst.In partiular, thy throw rmarkal light on th sl-alldam-
I Buffon. on th on hand, his on-
its gradual hang, th influn of brianplosion' th sminglyarupt apprnand volutionary adiation
of th animals. Th plosion is slf-vidntfrom th dramati apparanof
[if, and xtintion look gnuinly
skltlfossils arrd th divrsifiati<lnof tr fossils, yt it is now apparnt
lr th arupt origin of spis,with
460 Burgss Sbl
voltion in th anrr
Shal_type faun:rsnray
thy rmin a ftil fil
ILloGRAP|-tY
Budd' G. .' and S. Jns
iltrianphy[a.8ill
onway orris, S. l998.
of Aimls.amrid
Llotr,..G., R. J. Aldridg
204. Th mbri
iml Lif. for1
Valnrin'J. W. 2004.
Prss.
Zhuravlv,A. Y., nd R. t
Rditio, Nw Ylk
|BLloGRAPHY
Budd,G. ., and S. Jnsn.2000.A ritialrappraisal of th fssilrordof th
ilarrianp|a.Bkllogil Ruitus75: 25-296.
onwy Morris, s. 1998. Th ruibl lf rtio:Th BrgssShlnd th Ris
of imls. amridg:Can.rbridg UnivrsityPrss.
Hu, '-G., R. J. ldridg,J. Brgstrm, D. J. Sivtr,D. J. Sivtr,and ..H. Fng.
2004, Th mbri Fossilsf bgjig,hi: Th Floeriglf rl
Aiml Lif. Oford: Blakwil.
Valntin,J. !. 2004. o th rigi of Phla.hiago:Univrsityof hiago
Prss.
Zhuravlv,A. Y., and R. Riding,ds.2001. Th llogof th mbri
Rditio.Nw York: olumiUnivrsitvPrss. -s...
BILIoGRAPHY
Darwin, . 1859. ()n th, rigi of Spies.London:John urray.
Howard, D. J., nd S. . Brlhr,ds. 1998.dlssForms:Spisd
Spitio.Nw York; Oford UnivrsityPrss. ain, Arthur Jams
Mayr' . 1942.Sstemticsd th rigi of Spcis,Nw York: olumbia
UnivrsityPrss. -^4.R. A. J. ain (h always us
of th group of British .
Ford's oIogil gni
.!Var
tr sorld II. Thy f
as a major ( t h yw o u l d
duatd at Oford f
sltion ws rgardd'n
gists, lik Linar Profs
on volution, and whn
drift wre muh favord
ti) rasons for distrustr
lin of thinking riv
| 9 4 6 o f h s p i r i r u l i st
Profssor of Zoology (;
Togthr with fllow
light of thory ing dv
ampl of Ford, ain st t
of ntur. Thir most
emorlis, widly found
trysid round oxford. l
ing and with plain shlls
takn to a pardigma
fturs Wr thought fal
ain nd Shppard sh
th akgounds in whi
xampl, is diffntfron
from th grass of a fild 1
a major lmnt in th dj
to th akgrounds th l
portant tool in making t
nvils (stons)on whih
ould ollt nd ount t
d i f f n tl l i r i s( i n
ml Speis d hir ]
sujt.
ln dmand for Darwinian
bsrvdgroup atually rak-
i if th work of Bush ndd
Lso oftn th ritis r mo-
a rligious natur rathr than
final word on th topi.
n:JhnMurry.
llessForms:Specisd
ain, Arthur Jams (92|_|999)
ies.Nw York: olumia
-^4.R. A. J. ain (h always usd only initials for profssionalpuliations) Was on
of th group of British volutionary iologists who wr mmrs of . B.
Ford's ologial gntisgroup at Oford Univrsity in th arly dadsaf-
tr NorldWar II. Thy firmly stalishdth signifianof ntural sltion
as a major (thy would hav said th major| aus of volutionry hang.
duatd at Oford for th wr, ain had ntrd a world whr natural
sltionwas rgardd with suspiion, if not outright hostility. Most iolo-
gists' lik Linar Prlfssorf Zoology . S. Goodrih, did not ltur at all
on voluti<ln,and whn th topi Was mntiond altrnativssuh as gnti
drift wr muh favord. Not a fw hd fairly ovrt ligious (at last vitalis-
ti) rasons for distustinga purly naturalisti approah to origins, and this
lin of thinking rivd almost offiial support with th appointmnt in
\946 of th spiritualist Alistr Hardy as sussorto Goldrih as th Linar
Profssorof Zollogy (ain and Provin 1991; Rus 1996).
Togthr with fllow Oonin iologist Philip Shppard, wrking in th
light of thory ing dvelpdy Ronald A. Fishr and th xprimntal -
mpl of Ford, ain st out to show that natural sltionis a powrful for
of natur. Thir most lratdand lasting work was on th snat| p
morlis,widly found in th woods, madlws,and hdgrowsof th oun-
trysid around oford. Th snail oms in many diffrnt ftrrms,with and-
ing and with plain shlls, and with pink nd rown and othr olors. It ws
takn to a paradigmti ampl of gnti drift; suh simpl and vrying
fatursWr thought far too insignifiantto hav any tru adaptiv valu.
ain and Shppard showd that in fat th shll ollrs ar tightly tid to
th akgrounds in whih th snails a found. A hw.lodforst floo, for
ampl, is diffrntfrom a dappld dith sida hdg,and again diffnt
frm th grass of a fild grazd y ttl. Thy disovrdthat th snails ar
a major lmnt in th dits of irds' thrushs spially' and that aording
to th akgrounds th snails ar apturd and atn diffrntially. An in-r.
portant tool in mking ths dtrminationswas th fat that thrushs us
anvils (stons)on whih to smash th snail shlls, and hn th rsarhrs
ould ollt and ount th rlativ numrs of forms takn y th irds in
diffrnt loalitis (ain 1954; in nd Shppd 1950, 1954). ain's zl-
ml Speis d Thir uolution (1954) m a standard work on th
suit.
165
466 i
latural history that bgan as a of gntis.Lik nrany of his gnration, h gan as an mryologist, swith-
oung ollgstudnt mmr of ing to prolms of hrdity whn Mndl's work was rdisoverd at th -
ginning of th twntieth ntury. astl workd mainly n mammals, although
s at th Univrsity of Hawaii's h was also th first to us th fruit fly Drosophil mlogstr,whih bam
vry ativin rsarh and on- th volutionist's workhors in th twntith ntury. Thomas unt Morgan
o volutionary biology. H is Won th Nol Prize for diphring th gnti natur of this organism.
y o f H r w a i i . sm o s t m i n n ts i - astl's most famolls sris of primnts Wr on rats of varying lak
) oursof his distinguishd a- and whit oat olors (s figur on pg 47). |1 prformd sltion x-
gus nd to his many studnts. primnts tht showd how on ould hang th ovrall rang of hus in a
rt sintist.on, whn h was population, from mixd to virtually ntirly whit or lak (astl and
' a symposium hld in honor of Phillips |91'4)..Whatwas signifiantaout this work ws tht on th sl-
humling.Sinis a slial n- tion was finishd, th ts' oat olors ontinud to hovr round th final
vry whih dirtion. . . . Th point ahivd. In othr words, astl showd what harls Darwin had
Hwaiin Droslphila Projt] laimd and what many ritis had dnid: seltion an hav lasting if not
[Hawaiian word for loals] stu- prmanntfftson th ovrall natur of a population of organisms.astl's
8 5 ) .I n d d , a r s o n ' s H a w a i i a n work provd that on sltionis rlad, vrythingdlsnot rvrt ak to
ining gror.rndfor mor than 400 th starting point.
studntsin all spts of volu' Throughout th twntith ntury thr wr numrous othr primnts
intistsfrlm all ovr th wold showing that sltion an hv lasting ffts. Paradoially, howvr, al-
laustlf arson's ladrship in though Castl's work is rightly haild as a major xprimntal stp on th
lility t ring togthr th st Way to modrn volutionary thinking, h himslf did not ntr into th -
. 'w h o h s n l t o m o t i v t primnt intnding to slrow th worth of sltion. Rathr, as a gntiist' his
lns to unlok th srtsof vo- first intrstwas in th ntur of th hrital fators that Mndlism pr-
rink out how knowldg an supposs.astl was trying to show that ths fators, what ar now alld
:1nt.Ths qualitis mad this gns'ar not ntir and unhangalin th ours of gnration' ut split
up as prodution ours and romin in nw variations in th ours of
tim. It was only aftr Morgan's work was ompletd in th sond dad of
th ntury that astl ondd that h had n wrong and that (apart
dings. 1989. Hmpton Lawrn
.W. from variations ausd y spontanous hngs' or mutations) gns ar
y. In W. Andrson, K. Y. tansmittdunhang.
s, Spitkltt'd tb Fldr
In th ours of his long ar, astl had many studnts' th most fa-
y Prss.
] wY t l r k : | u m l ' i U n i v r s i t y mous and important of whom Was gntiist and volutionist Swall \/right
(sProvin 1986).In anothr paradox, Wright is proprly known as on of
f spis.Si 168: 1'414_141'8' th foundrs of modrn volutionary thought, yt th ulk of his mpiril
S. Karlin nd . Nvo, ds., rsarhwork was on th pattrnsof hrdity as shown y guina pigs' an in-
). orland, FL: Aadmi Prss.
trstthat wnt k to his studntshioundr astl.
n :J o h n u r r a y .
rvolutionsin mirovolution nd
|LloGRAPHY
Andrson,K. Y. Kanshiro, and
.W..
d th FtludrPriipl, 4_76. astl, ., and J. . Phillips. |L)14.Pibld Rtsd Sltio: primtl
-.Y.. stof th fftiusslf Sltiod of th Tlrof GmetPurit i
.Washington, .lfashington.
dlirosses. D: arngiInstitutionof
rovin,!. B. 1986'StullWrightd ullutior Biolog.hiago:Univrsity
of hiagoPrss.
Rus,. 1996. od to : h optlf Progrssi uolutilrBillog.
vrsityand latr at th Bussy In- amridg, A: Harvrd Univrsity Prss -.R.
ltriutdmuh to th arly yars
470 stle
fifrq
The usesof I
f,
-.2 - !
T 'r
iq
(i
J.B.s.Haldan's
is th apston txt
haild primarily for
hitts of populati
\/
( 1 9 3 1 ) ,a n d a l d n
I a synthsis of lassir
hmistry rvith popr
disussion is onfin
ial nratrial drawn
Arystwyth, ntitl
Haldan's prolr
n of natural sl
shas th prolrna
answr) intrwovn
part follows Fishr i
dt1 to low sltion
dan's support lf Fis
rrid th Inathml
hanging nvlrnm
and in part as a rlt
ovr, and important
ltion, r<lgnizig
,'.* populations may oI
Ths disussins
di of uss. Y|a|
uss of within- an
trs and a road ss
nal hrptr.
.Working
throug t
ry of Mndlism in
and btwn spi
pral 1oi, diffn
mosoms' or o{ wh
Gtics d th Ol
gu tht th sam s
Willianrastl,sdmonstrtionoftfftsofsltitrnforoato1or.op within spisou
r o L u : | e l a t o l o r . . . t . u , " d f o r l a s s i f i a t i o n . i l r l t r tht
l z : sastl's
l t i owork
n-forlight
olr. Th pradx roadns th ndt
o|or.Btlttrlt,ulz'.,.t....l. r",...t tat th units f
was aimd at disp.ov,nj .'"r.r *ntis(v showing Th onlusion
.."^.::pJ:u.c th ffts of natrtral
hrdity lnd in ah gnrtigl), thought sin Darw
a s t l a n d P h i l l i p s l 9 1 4 , P l t 1 . )
'.-i.t'om th sarn tim, Hal
dominant rativ {
rativ aus.hr
volution, inludin5
that mirrds ar sur
Th aussof volutin 471
+4
Tb uses of uolutio (I. B. S. Haldan)
J. B. s. Haldan's (7892*1964) The ses of ullutio' pulishd tn 1932,
is th apston tt in th origins of thortial ppulation gntis. uss is
haild primarily for its important, ritial disussion of th work of th ar-
\
hitts of population gntis' R. A. Fishr (1930 [195s]), Swall \/right
! (1931),and aldan himslf. It is lss widly, yt just as dsrvdly,haild as
I
a synthsisof lassil gntis,hromosomal mhanis' ytology, and io-
hmistry with population gntis.This is ironi austh mathmatial
disussionis onfind to an appndi addd to original disussionsof iolog-
.W.als,
ial matrial drawn from Haldan's 191 ltursat th Univrsity of
Abrystwyth, ntitld ..A
Rxamination of Darwinism.''
Haldan's prolmati may posd as, ..What is th natur and signifi-
an of natural sltionin Mndlian popultions?'' In this sns,Haldan
sharsth prolmati of his ontmporaris Fishr and \/right.And in his
answr, intrwovn with supporting mpirial vidn, Haldan in larg
part follows F.ishrin th viw that umulativ volutionary hang is mostly
du to low sltion prssursting on mutations of small fft. But Hal-
dan'ssupport of Fishr is not in th form of an unritial summary. aldan
arrid th mathmatial ploration of sltion in finit population sizs,
hanging nvironmnts, and multipl dimnsions furthr thn Fishr had,
and in part as a ritil onsidration of !right,s work on th topi. Mor-
ov, and implrtantly,Haldan emphasizd th problm of th units of s.
ltion, rognizing that sltiv fors ating on gamts' organrsms' Or
populations may om into onflit.
Ths disussions ar arrid orrt in th middl haptrs and th appn-
di of ss. Haldan mds thm in rih disussion of th gnti
auss of within- and twn-spisdiffrnsdisussd in arlir hap-
ts and a road ssssmntof th sin of volution as whol in th fi-
nal haptr.
\Vorking through mpirial vidn amassd from th tim of th rdisov-
ry of Mndlism in th 1900s, Haldn dmonstrats tht diffrns within
and twn spis ar du to diffrns twn individual gns at om-
paral loi' diffrns in numr of individul hromosoms or sts of hro-
mosoms' or of whol lrromosoms. Antiipating Thodosius Dozhansky's
Getics d th origi of Speies (197), Haldan uss this vidn to ar-
. sltionfor oat olor. Top
gu that th sam sorts of prosss that aount for th gnti diffrns
iddl rop: sltionfor light
within spis aount for suh diffrns twn sis. Furthr, Haldan
' paradois that astl'swork
roadns th ndlian ontxt of volution.
y shwingthat th units of
ng th fftsof natural Th onlusion of uss mphasizs th progrss mad in volutionary
l.) thought sin Darwin and th powf of volutionary planations of lif. At
th sam tim. Hldan is lar that whil h ss natural sltion as th
dominant rativ volutionary us, h dos not lim that it is th only
rativ aus. Througout ses Haldan highlights th multipl auss of
volution, inluding dvlopmntalonstraints ln volution. Haldan insists
that minds ar surely produts of volution;thy also ar in all liklihood not
472 ulliSforz
ussionin ldan's onltrding Gs (1994) fittingly slll.Ilsth radth of lris ndvor. Haild s brak.
in a distitlymatrialist philos- through in th urrdrstanding of human volution, it offrs th first full-sal
ronstrution of whr humn populations originatd nd th paths y
for him' omplishd a ron_ whih hy sprd throughout th world. By mapping th worldwid go-
hrdity.Haldan forgd ahad graphi distriution of gnsfor ovr 110 traits in ovr 1'800 primaily ao-
rvidswath of iology with pop. riginal populations' migations we hartd and lok was dvisd to dat
part and parl f th origins o{ voiutionry history. ssntial information was ntriutd by arhology.
lat is ommonly alld th mod- In Th Nolithic Trsitio d th Geetis of Popltios i urlpe
. r . i t i s g o u n d r k i n gi n i t s (1984)' h had addrssdth qustion of wthr ultural diffusion was du
to th spad of idas or of popl. Th issu was invstigatd at a world
sal wih spial attntion to th link twn gnti divrsity and lan-
guagdiffrntiation.Although linguists hav not rirhdonsnsuson lan-
f Spis.Nw York: olunrbi guag lassifiation at gloal sal' an intriguing ovrlap btwn th
distriution of gns and languags Was found at diffrnt gographi
f NturISltitllt.2ndd. Nw sls.
. l,ondon: Lngmans.
Givn th brdth of his rsarh intrsts, it is not surprising to find a
tions. Gtis |6: 97_1'59. gratdal of wok olltd in a ook ddiatdto th ploration of modls
-R..s. that miglrt hlp us undrst;rnd how ultur:rl traits trr trnsmitted (ulturl
Trsmissiond uolutil' oallthord with . w. Fldman' 19s1).
avalli.Sforz has njoyd a rmarkaly rativ sintifi longvity. R-
)\ ntly his laboratly dvlopd nw thniqus for disoving DNA varia-
tion usd to dfin nw gnti markrs of th Y and othr hromosoms.
] a modrn rinarlrationof a R- Th study of ths uniprntally tansmittd marks trnd out to esp-
to voltionaryiollgy mgd ially fftivirr luidting popr:lation origins.
ang of filds that h ultivatd at Cavalli-Sforza has natural gift for ommuniating his passion for r-
otr for undstandingth human sarh. H is th author of mny gnral audin ooks and nWspapr
olurns. Th ttook The Getis of Humt Popultios (oauthord
1940s,h rifly pratid md- with !. Bodmr, 1971| was awardd a prize as th most quotd tt in th
s;rrh'Attatd y th statistial f i l di n 1 9 8 1.
ndrth guidanof R. A. Fishr
rm tim (in th arly 1950s), h ILIoGRAPHY
. i l s .a h i g h l y i n n o v a t i v t o p i . L. L.' and A. Ammrman.,l,984.h Neolithicrsitiond tb
availi.S{oza,
utd lasting mthodologial ad- Gtiso|-Popultiotlsi rop. Printon,NJ: ritonUnivrsityPss.
.W.
'_dwardsgnti distan, an s- avalli-Sforza,L. l-., nd Bodmr. 1971.b Gtisof Hum Popultios.
SanFrniso:W. H. Frmn;inola,NY: DovrPulitions.
grphi studis (irnothr fild h
avalli-Sforza,
L. L.' and . W. FIdmn.198l . ulturITrnsmissil d
gaphyof gns',). uolutkl,Printon'NJ: PrintonUnivrsityPrss.
rihnssof historial rords in valli-Sforza,L. l-', P. nozzi, and.Piazza. 1994. he Histor nd Gogrph
. dispnsations)in lrurnan popula. of 17t,ttGs,Prineton,NJ: PrintonUnivrsityPrss.
lutionary sintiststo liv that avalli-Sforz,
L. L., A. oroni, and G. Zet.24,osguinit, Ibreeding,
d
GticDrift i [tl.Printon,NJ: PrintonUnivrsityPrss. -P.'
rrranpopulations. In 24, o-
i ltl' organiirlly prsrrtdth
of gntisat StarrfordUnivrsit
Rort(Is02_1s7I)
hamrs,
mritusprofssor. Rort hambrs Was ol-l of tlr most sussful usinssmn in Vitorian
f modrn human gns for ron- Britain. Togthr with his rothr, William, h had pulld himslf up from
Llistlrd Geograph of Humn abjt povrty to found a thriving pulishing businss. Thy wr major
474 hmbrs
dsignr.To Ply, th hoi was ovious; th intriat omplity of fa- population of alls
turs suh as th y was inompatil with anything ut a dsignr,a thsis nal population-y
ommonly rfrrd to as th Argumnt from Dsign. whr th gnotyp
Darwin's us of han variations distinguishd his idas not nly frm ltiv prssurs.
thos of rationists ut also from othr advoats of ompting thoris of Thr ar two g
volution, partiularlyJn-BaptistLmark (1809). Arding to Lamark, a atastroph or ot
mutations ar dirtd toward an organism's nds; thy aris with rspt t suh s a forst fir
th us and disus of an organism'sparts. For xampl, giraff nks r lon- ass it dos so indi
gatd aus giraffs strthd thm to rah trtop foliag. opposing this ganisms. h smal
modl, Darwinian volution lims that mutations ar undirtd, han gnration may hal
vnts;mutations for long-nkdgiraffswr just as likly as mutations for ltion. A similar for
short-nkd giraffs. Howvr, on nk-lngth mutations ourrd, th is, an ur wh
long-nkd giraffs had a highr fitnss valu, thus sprading thir long-nk poulation and sta
gns throughout th population. Th ritial point in Darwin's argumnt is Th migrating grou
that although mutations aris y han, it is not y han that som indi- ity with thm. Th
viduals surviv and rprodu whil othrs pish. Natural sltion distin- rat a nw popul
guishes twnnfiia1nd dltriousmutations. land prognitors,
hos of ths argumnts mrg in modrn ritiisms of Darwinian vo- rol. This pross l
lution that ar fousd on th sming surdity tht han variations importan ar link
ould vr produ intriat struturs or wll-ordrd iohmial pro. A sond form o
sss.Ths argumnts ar asd on mistakn idas of th rol of han th nxt, in all limi
that ho th argumnts in Darwin,s tim. Fist, han variations produ akin to Dozhanky
raw matrial that is moldd by natual sltion;without natural sltion qunis hang gr
to st th dirtion and spd of hang, hn variations ould not au- alll oms th '
mulat with nough spd to produ muh mningful hang in any stru- ours usth
tu or pross. Sond, han atually has a spifi maning with rspt ll frqunis.
to Darwin's han variations nd th gnti mutations tht aus thm. mon' and an unO
FIr,han mans that spifi mutations our at random with rsptto beoms xtrmly
th nvironmnt so that, unlik th l,amarkian argumnt' mutations ar aus allls may
not dirtd y nvironmntal fatursor th liklihood that thy will prov whih on will vr
usful. Som spifi mutations ar indd mor likly thn othrs-for x- Swall !right, r,
ampl, som mous oat olor mutations ar mor likly to aris than sponsil for fusin
othrs-ut thir potntial usfulnsshas no aring on thir liklihood of first to mphasizt
apparing. gud that thr wr
\Whnontmporary iologists rais th notion of han thy likly ar r- advantagous for tl
frring to rndom drift. Random drift rfrs to th hang in gnti ompo- of ths ominati
..shifting alan''
sition that ours in a population from on gnrationto th nxt ausof
.s7right
th random smpling of gns that ours in a finit population. A familiar rr, would
ampl of random gnti sampling is th s ratio within a family; in th- 1940s through th
ory, half of th hildn in a family should oys and half girls, ut it is not lution (Provin 198
unommon for a fmily of four hildrn to onsist of thr oys and on girl Th most influr
ausof th random draw of th s hromosoms.Th lassi illustration atd argumnt for
o f r a n d o m d r i f t w a s g i v n y T h e l d o s i u sD o z h a n s k y ( 1 9 7 , 1 2 9 ) ' w h r h thory of volution
dsrida snario of lindly drawing alls fom an urn. Th alls wr ol. wr disovrdan
ord diffntly, ut othrwis indistinguishal.If on lindly draws alls gl protin that s
from an urn and plas thm in sond urn, it is likly that th sampl wr two or mor
hn d uollttion 477
i; th intriat omplity of fa- population of alls diffrs in frqurry'wit gard to olor, from the origi-
r aIrthingt a dsignr' a thsis nal popultion-y han lon. In short, rarrdm drift dsisa snario
n Dsign. whr th gnotyp frquny of a population hangs in th :rsnof s-
rguishdhis idas not only from ltiv prssurs.
Jvoiltslf ornpting thoris of hr two gnrl kinds of random drift: isd smpling as rsult of
:k (1809).Aording to Lamark, a atastroph or Othr norrsltiviasing, and gnti drift. atastroph'
's nls;thy aris with rspt to suh as forst fir' may kill a larg portion of a population, ut in most
r ampl,girff nks r lon- assit dos so indisriminatl1,rvith rsptto ny prtiular rit in th or-
lh trtopfoliag. Opposing this ganisms. Th srll num of survivors tht ar lft to produ th nt
xutations r rrndirtd' han gnrationmay hav a diffrnt gnti omposition than th original popu.
vr just as likly as mutations for ltion. A sinrilar fom of drift, thar is, n rupt hng in gnotyp frqun-
k-lngth mutations ourrd, th is, an our whn a srnall gror.rpof organisms migrat from th main
lu,tlrrtsspradingthir long-nk population and start a nw on' suh as ours whn an island is olonizd.
| p o i n t i n D a w i n ' s i l g u n r n ti s Th migtinggroup may only tk :r fration of th original gntivariail-
t is not y han that som indi- i t y w i t h r h m . T l r u n i q u g n i s u t u r o f t h f i l u n d i n g p o p u I a t i o n a n
.s prish.Natural sltion distin- rat a nw popr"rlationthat is ognizaly distint fom its oiginal' main-
11tlt1tlons. land prognitors!vn though natual sltio my av playd a minimal
l d r n r i t i i s m s f D a r w i n i a n v o - rol. Tlris pross has n alld th foundr fft, and argllmnts for its
a s u d i yt h a t h n v a r i a t i o n s importnar linkd to th influnof rnst Mayr (1954)'
rr wll_ordrdiohmial pro- A sond form of randorn drift ours grdually, fror on gntion to
;taknidas of th rol of han th nt, in all limitd populations through indisrinrinatgamt sampling
. First, han variations produ akin to Dozhanky's analogy of th alls in tlr urn. Gnotyp nd alll fr-
lltin;withut naturl sltion qunishang gadually aross ]nrations'irnd, givn nough tim, on
han variations ould not au- alll oms tlr only gn in th populatiln (that is, it is fid). Fiation
:h maningfulhang in any stru- ours austh dirtion of andom gnti drift is dtrmind y th al-
:as spifimaning with rspt ll fqunis.Tvpially' a ommon lll is liklv to om mor om-
:nti mutations that us thm. mon' and an unommon on is likly to om rar. vntually on lll
rs oLlrat random with rspt to omstrmly ommon and thn Ad. Th han lmnt rmains .
. uttions
r n r k i n r g u m I . l tm aus allls ma qually ommon' thus rrr:rkingit impossil to prdit
th liklihood that thy will prov whih on will vntually bom fid.
c mor likly than oths-fr - Swall Wright, who along with J. B. S. aldan and R. A. F.ishrwas r-
ns r rnor likly to iris than sponsilfor fusing Mndlian gntiswith Darwiniar-rvolution' was th
; no aring on thir liklihood of first to mphasiz th rol that random drift plays in volution. Wright ar-
gud that thr wuld ftrr diffrnt ominations f gns that would
notion of hn thy likly ar r- dvantagousfor thir arirs and that random drift would influnwhih
: r sr t h h n g i n g n e t i o m p o - of thsomirrtionswould mrg and sprad through a spis(Ii7right's
..shifting
gnrationto th nxt aus of alan'' thory of volution). Howvr' throughout his long a-
.!7right
s in finit population. A familia rr, would vaillat on th importan of rndom drift and in th
l s ratio within a family; in th- 1940s through tlr 1960s h dlgtdrandom drift to minr for in vo-
[ oys and half girls, ut it is not l u t i o n ( P r o v i n 1 9 8 6 ) .
o onsistof thr oys and on girl Th most influntial appliation of gnti drift-and th nrost sophisti-
omosoms.Th lassi illustration td rgumnt fo th rol of h:rrrin vlution-is th so-alld nutral
D o z h a n s k (y1 9 7 , 1 2 9 ) ,w h r h thoryof volutiln.As prtins and th DNA squnsthat odd for thm
'lls from an urn. Th alls wr ol- wr disovrdand plord, iologistswr finding nrany variants of sin-
rishbl.If on lindly draws balls gl prltin that smd to hv quivalnt funtionirrg.In som ass' thr
Ld urn, it is likly that th sampl Wr two or mor variants in :r singl ppulation and, in othrs' diffrnt
478 hne d uolutio
spis wr fixd for diffrnt vriants. otoo Kimura rognizd that ths Mayr, . 1954. hang
osrvationswr fats of a singl prolm unitd y th onpt of gnti Hardy, and . B. Ft
.$right's Alln and Unwin.
drift. H xtndd original mathmatial work on modling of gn-
Millstin, R. L.2000. (
ti drift to show how thspattrnswould mrg (Kimura 1983). This work
no.4: 603-624.
ld, in turn' to using DNA squnsas ..molularloks'' and to a rvolu- Paly'
.!.
l1802l 1819.
tion in undrstandinghow spis,gnra' familis, and vn ntir phyl ar Rivington.
.!.
r l a t d( K i m u r a 1 9 8 3 ) . Provin, B. 1986. Se
From a marovolutionary prsptiv'som iologists argu that vnts of hiago Prss.
suh as tintions, spis divrsifiation' radiations, and inrasing om-
plxity an st xplaind stohastially,without rfrnto a partiular
usal story. This is not to say that thr ar no undrlying auss to
Charlsworth, Br
marovolutionary vnts, ut that stohasti mthods sm to aptur ths Brin harlsworth il
vnts ttr than dtrministi mthlds (Millstin 2000). For ampl, is 50 yars. H is knowr
th inrasing mplxity of th organi world du to passiv or ativ imntal work has
trnds, that is, y han or dirtdvolution? Although it is diffiult to pro- for svral rasons:a
vid nssaryand suffiintonditions for iologial omplxity, it is gnr- tion of th onntior
ally agrd that lif originatd as simpl organisms and grdually am and numrial appro
muh mor ompl. Baus lif gan nar th minimum oundary o{ work of oths, and al
omplxity, rndom variation alon would produ mor ompl organ- Brian harlswortl
isms; this would a passiv trnd. An ativ trnd would mplifid y BA in Natural Sin
dvlopmntal routins or phnotypi novltis that driv volution toward honors in 1966.He m
inrsing omplity, yond that of hane (arroll 2001). orah harlsworth' l
A final sns of han in volutionary thory is that arriving at any pati- graduation' oth ha
ular nd point in volutionary spa is a han vnt. For xampl, Stphn .
studying undr J. .
Gould (1,989)fmously laimd that humans ar an improbabl nd point on Upon riving his
th volutionary tr;many unrlatd (han)vntsulminatd in th is- with Rihard Lwonti
tn of humans. Gould did not imply that thr ar not sltivrasons an offi with Td G
(mong othrs) xplaining th volutitln of humans. Rathr, h argud tht populations. B.. r
humans ar a rsult of a vast numbr of improal vnts.This is th sns lm; this ffort produl
of han, attriutd to Aristotl, whr indpndntausal hains omin fin fitnss and dsr
to produ an unintndd fft. A ar aidnt, for ampl' is n instan hangs. His ook,
of spartausal hains omining and produing an unintndd rsult. rmains th primary s
In 7971B.. was h
BIBLIOGRAPHY
tiatd xprimntal
Batty'J. 1984.hannd naturlsltion. Phil<lsrlph
lfSi51, no. 2: sions with D.. ld t]
183-211. hangs that afft tht
arroll' S. B. 2001. Chanand nssity:
Th volutionof morphologial
ifirs f rominatior
omplityand divrsity.Ntttr409 (Frury22|:1102_1109,
Darwin, . 1859. th rigi of Sples.London:John uray. immdiat rsult was
Dobzhansky,. |97. Gtisd th Origi of Sples. Nw York: olumia tht launhd D..'s
UnivrsityPrss. Th harlsworths
l,G. J. 1999.On th dul naturof hrrin volutionaryiologynd Maynard Smith (B..
paloiology.Plobiolog25 75*87. la at th National ln
Gould' s. 1989. WonderfulLif: The BurgssShld tb Ntur o/Hls/ry. Nw
olina bgan B..'s wor
York: W.'W. Norton.
Kimura, 14. 1983. h Nutrl Thrlrof olulr uolutio.mridg: tr' Jan, Was orn in
ambridgUnivrsityrss. Aftr attnding a sy
Lamark,J. B. 1809.Philosophizoologiqu'2 vols.Pris:Dntu. no-Darwinism in 19
hrlsorth, Brin 479
o Kimura rognizdthat ths ayr, . 1954. hng f gntinvironmnt and volution. In J. uxly, A. .
unitdy th orrptof gnti Hardy, and . B' Ford, ds., t.'olutions Proess, 157_180. London: Gorg
Allrr and Unwir-r.
Ltialwok orr modling of gn_
illstin, R. L. 2000. han zrnd mrvo|ution,Philosoph of Sine 67,
r r g ( K i m u r a 1 9 8 3 ) .T h i s w o r k no.4: 603-624.
llularloks'' and to a rvolu- Paly,W. [1802l 1819. Nturl Thollg (ollted Wlrks' vol. 4). London:
milis,and vn ntir phyla ar Rivington.
Prvin,W. B. l986. S,ll Wrigt ld uollttior1,Biolog. hiago: Univrsity
of hiago Prss. -].Z.
lm iologistsargu that vnts
r l i t i o n s. n d i n r s i n g o m .
without rfrnto a partiular
' r no undrlving auss to harlsworth,Brian (b. 1945)
mthodssInto aptur thS Brian Charlswoth is on of th frrnostpopultion gntiisrsof th iast
u{illstin2000). For xampl, is 50 yars.H is krrown primarily for his thotialwork, although his pr-
w o l d d u t o p i r s s i v O r a t i v imntal Work has n sustantial. harlsworth's work hs n influential
ln? Although it is diffiult to pro- for svral rasons: willingnss to tak on diffiult prolms' an appreia-
l i o l t l g i l o m p l x i t y .i t i s g n r - tion of th onntion twn tlrory nd data, lvr us of mathmatial
rganismsand gradually am and numrial pproahs, honst assssmntsof th shotomings of th
lar th minimum boundary of work of othrs, ad an unusually road knowldg of th sintifi litratur.
l produ mor omplx organ- Brian harlsworth (B..) was born in ngland in 1945 and otaind his
/ trnd would mplifid y BA in Naturl Sins from Qun's ollg, amridg, with first lass
ltis that driv volution tward honors t |966. H mt his futur wif, Dorah Malrby (. |94, now D-
: ( a r r o l l2 0 0 1 ) . orah hrlsworth' hraftr D..), in his first ar at amridg. Upon
: o r r .i s t h e t a r i v i n ga t n y p r t i - graduation,blth harlswots Wnt on to PhD studis at amridg, B.C.
tn vnt.For ampl, Stphn studyingundr J. NI. Thoday.
s a an improal nd point n Upn riving his dgr in |969, B.. took a postdotoral fllowship
) vntsulnrinatdin th xis- with Rihard Lwontin at th Univrsity of Chigo. At hiago, B.. shrd
rt thr ar not sltiv rasons an offi with Td Gisl, wo was simulating volution in ag-struturd
'hutlans.
Rathe, h argud that populations.B.C. realizd tht h ould otain analytial rsults on th pro-
prolvnts,Tlris is th sns lm; this ffort produd a sustantialody of work that showd how to d-
dpndntausal hains ombin fin fitnss and dsri tlr rlationship twn gnti nd dmographi
i d n t ,f r a m p | .i s a n i n s t n hangs. His ook' uolutillt i g-Strtured Popultios (1980, |994),
oduing an unintndd rsult' rmainsth primry sour on th subjt.
| 1971B.. wils hird s a lturr at th Univrsity of Livrpool and ini-
tiatd primntal work on th voiution of rornination rats. Distls-
hilosophof Si51, no. 2: sions with D.. ld thm t study modifir thory' whih onrns gnti
hangs that afft th volutionary fors ating on othr loi, suh as mod-
:volrrtionof morphologial ifisof romination, dominan' mating systms,and nrutation rts' Th
.ur2v 2 ) :l 1 0 2 _ 1l 0 9 .
immdit rslllt was a ollaorativ sris of paprs on Batsian mimiry
rdon:JlhnN{r-rrr;ry.
of Spis. Nw York: olumi tht launhd D..'s arr irr volutionar biology.
Th harlswlrths movd to sussx in |974, thanks to fforts y John
iologynd
in vllutinary aynard Smith (B..'s hD aminr).A saatialyar in huk Langly's
la t th Nationirl Institutof nvironmntal Halth Sinsin North ar-
\Id th N,tturr;f Hrrr,. Nw olina ganB..'s work on tri1nsposal lmnts.Tli harlsworths' daugh-
tr,Jan, Was born in 1981.
ulruoltklz. amridg:
Aftr attnding a symposium at hiago,s Fild Musum on altrnativs to
vols.Paris:Dntu. no-Darwinism irr 1980, B.. oauthord a papr with Monty Slatkin and
h rlesulorth,D eb orh
Russ Land that sharply ritiqud hallngsto no.Darwinism. Ths hal- of Livrpool, D.. was
lngs prinipally stmmd from Stphn J. Gould's tnsions of Nils l- miriry y thir oll;
drdg and Gould's onpt o{ puntuatd quiliriunr. Tis papr hlpd in Batsian mimiry wa
atapult B.C. from a rsptd prt on mathmatial thry to a ladr in trn ould rought
t h f i l d o f v o | u t i o n .B . C . h s s i n t k n m a n y o } p o r t u n i t i tso r i t i i z B.. ld to ir ollaora
idas outsid orthodo no-Darwinism' frrn Garil Dovr's suggstiontht D..'s arr in volut
transposal lmnts might play a ausal rol in volution to rnt at- harlswort |975).
tmpts to rt a sinof IntlligntDsign. Undrstanding th r
Th Chalsworths movd to th Univsity of Chiago in 198.5,whr drstanding how volut
B.C. was dpartmnt hair fom 1986 to \997' The harlsworths' transi- att:rkd oth fats in
tion to th study of molular volution Was mntdy thir 1993 disov- stirnding of how gns
ry, with artin Morgan' that dltrious mutations rdu th n.runtof othr loi. This work w
variation in parts of th gnom with low rmination. This providd a gnOm volr-rtion.
novl planation fo a wll-stalishdpattrn that was thn asrid to th D.C. pursud th stt
hithhiking ffts of nfiial mutations. n-roststriking prolms,
| t991, B.C. was namd a Fllow of th Royal Soity,whih failitatda rorphisms that ford
rturn to th Unitd Kingdom via a Royal Soit Rsarh profssorship. and harlswort |979
B.. ontinus to work primarily in mlulr volution and th volLltionof brding systm itslf '
s hromosoms. s a powrful frgui
(Charlsworth 2006).
IBLIoGRAPHY D..'s alrivmntst
harlsworth,B.1.978.A modlfor volutionof Y hrnrosoms and dosag sity of hiago (1988_1
ompnsation . Prodigsof th Ntiol dmlf SiesUS 7.\: as wll as a srisof hol
5618-5622. low of th Royl Soi
Ilgsol |994. uolution i g-StruturdPlpultios.2ndd. mbridg: Biology and volution (
amridgUnivrsityPrss.
gnom volution, part
harlsworth'B.' R. Land,and Nl. Slatkin.1982.A no-Drwiniirn ommrltar
on marovolutj'on.uoluti on 36: 47449 8, and nourags studn
Chrlsworth, B., ' T. Morgn,nd D. harlsworth . 199.Th fftof thir minds to aompli
dltri<lus muttinsn nutralmlulrvariatiln. Gtis|4:
1289-|33.
B|LloGRAPHY
harlsworth, B., P. Snigowski'and !. Stphan.7994.Tevolutionary dynamis
of rptitivDNA in ukryots. Nture37|: 215_220. -D.H. hrlsworth'D. 2006..
R726-R735.
2007.Q&A:D
harlswrth' Dorah (b. L94\ harlswth,D., trdB' l
mimiry.1. Singllo
Dorah harlsworth (nalty) is a pionr in the study lf rdingsys. 1'979a.volution
tms and gnom volution. onntratingon how r'o|uriortmolds stions t979. Mdl for
of th gnom to allow th gns for omplx traits to o.inhritd so that 467-498.
th traits fntion proprly without th disruption that rmination and
sxual rpodution might othrwis produ.
Dorah Charlsworth (D..) otind hr undrgraduat dgr from htvrikosrg
ambridg Univrsity, whr sh mt hr husand, Brian harlsworth Srgi Srgvihhtv
(B..). Aftr taking hr dotoral dgr in iohmial gntis, D.. ws sian ntomologist, an
not al to otain a position in sin and workd s B..'s unpaid assis. ntis. Born into a w
tant for th nt 10 yars. Aftr B.C. tok a position at th Urrivrsitv th Univrsiry of os
hetuerihou 481
of th twntith ntury. H is spially known for his synthti and intr- sts of rlatd x1
disiplinay attmpts to undrstand th totality of prosssinvolvd in th till gldulos,
origin of plant spis at a ruial tim in th history of volutionary iology. family), lausn,
Jnsn was orn in skilstrup, Dnmark, th son of farmrs and hous of otyps (gn
uildrs. His formal shooling took pla twnth ags of 8 and 14,ter- mhanisms of s
minating whn h had to tak on th rsponsiility of managing th family ologial fator
farm. H dvlopd an intrst in gntis and for aout ight yars duatd and arfully
himslf in th asi sinswith th aid of a shoolthr' gan to tak rst th lif in
an ativ intrst in th nw sin of gntis, studying Mndlian gntis inhritan) y s
along with Darwinian volutionary thory. In 1913 h ntrdth Univrsity y ringing to r
of opnhagn to study otany, gntis, and ology, oming in onrat world. Mor nal
with som of th shining lights in Sandinavin sin that inludd ompl intrpla
hrstn Raunkir (his major profssor in otany), P. Boysn Jnsn, Wil- polyploidy (hro
hlm Johannsn, and August Kogh. At on point h vn studid physis gnral thory ol
with Nils Bohr. His arly rsarhon gntisand ology in th Violaa Throughout tl
(th pansy family) ld to a mor dtaild undrstanding of hyridization in thir rsults in a r
plants and th phnomnon known as introgrssion,whry a smallr num- Institution. Thir
r of gns r introdud through th mhanis f hyridization. His hD spiation and 1
dissrtationon th Violaa' pulishd in \926, was on f th first mono- a viabl spis(
graphs to omin systmatis'ology, and gntisin any plant group. iosystmatis) t
was susquntly appointd assistant profssor to notd gntiist jvid gentis and
.Wing
at the Royal Agriultural ollg in opnhagn, ut his ral rak Dozhansky and
am with th granting of a Rokfllr fllowship in 1927_1928 that took i Plnts (1950)
him to th Univrsity of aliforni to work with . B. Baok on th gn- th nw fild of 1
tis of th gnus rpis. Whil thr, h mt with Harvy onroe Hll' a dis- lausn's owr
tinguishd alifornia otanist, who latr invitd him to partiipat in th the uoltio o,
nw intrdisiplinary projt h was organizing with th arngi Institution somwhat rif
.W.ashington
of to undrstnd mor prisly th mans y whih nw plant sngr Lturs
spis originatd undr natural onditions using nativ alifornia flora. rally n ovr
(Hall had arlir ollaoratd with logist Frdri lmnts on suh a ut nonthlss
projt.) Apting th offr' lausn arrivd in alifornia in 191, but in- ntury voluti
stantly found himslf th ladr of th nw tam that Hall had assmld th viw that v
whn Hall did unxptdly. nomna suh as
Th nw intrdisiplinary tam at th arngi Institution of !ashington In 1951,la
that was assmldat Stanford Univrsity inludd taonomist David Kk tird in 1956 ar
and physiologist \/illiam Hisey. Until th mid-1950s th tam of lausn, his wif, Anna I
Kk, and isy (as thy am to known) dsignda srisof lassial- rsarh positio
primnts to driv gnral prinipls for undrstanding plant volution un- in trms of stu
dr ontrolld, ut natural onditions. Adapting som of th lratd mtiulously
mthods of nintnth.nturyurpan workrs suh as Gston Bonnir, tory ttooks
and arly-twntith-ntury workrs suh as Gd,t Tursson, who sought to
undrstand th pross of adaptation, lausn, Kk, and Hisy prformd BIBLIOG
a numbr of transplant xprimnts involving thr distint loations along l a u s nJ, . 1 9 5 1
altitudinal gradints in northrn alifornia. On loation Was at sa lvl at UnivrsityP
Stanford, anothr at Mathr (at aout 4,600 ft)'and yt anoth at Timr- Frnh'. S. 19
lin in th Sirr Nvada mountain rang (at aout 10,000 ft).Prforming Biogrphi
Iuse 48.'
)Wn for lris synthti nd intr- stsof rlatd prirntaistudison nativ Cliforni plants suh as Pottt-
ity of prosssinvolvd in th till gldulos, chille spis, and th Madiina (th hayfild tarwd
historyof volutionary iology. family), lausn, Kk, and Hisy plord hyridization and th forration
th son of farmrs and hous of otyps (gntiallydistint ologial rs of plnts). Thy hartd th
wn th ags of 8 and 14, tr- mhanisms of spiation and rfind th undrstnding of th gnti and
sibility of rrrnaging th falily ologial fators that ld to th origin of lant spis.Thir wll dsignd
l for aout ight yars duatd and arfully xutd transplant studis ar gnrally rgardd as laying to
lhooltlrr. gan to tak rst th lif irr th irrhritnlf aquird hartristis(o Lanrarkiarr
: s ' s t u d y i n gM n d l i a n g n t i s inhritn)y showing th distintnssof th gnotyp and phnotyp, and
1913 h ntrdth Univrsity by ringing to rlif th importan of plrnotpi plastiity in th plant
nd logy, oming in ontat world. Mor narowly within plant volution, tir studis highlightd th
inavian sin that inludd omplx intrplay twnlryidization,apomiis, and phnomna suh as
.Wil-
otny), P. Boysrr Jnsn, polyploidy (hromosorrrduling), all of whih Wr ssnrialto driving a
point h vn studid physis gnralthory of plant volution.
;s and olgy in tlr Viola hroughout th 190s nd 1940s, lausn' Kk' and isy pulishd
Jrstandingof hyridization in thir rsults in a srisof important paprs and monlgraphs with th Carngi
ssion,whry a srllrnum_ Institution. Thir work was rad widely y othrs inrstdirr mhanisms of
rnis of hyridization' His PhD spiation and y thos intrstdin applying this undrstanding to dvlping
|26, was on of th first mono- a vial spis dfinition usful ir-rfornrulating th ..nw systmatis'' (alld
]ntisin ny plant group. H iosystmatis)that sought to intgratapprohs and undrstandingfrom
sor to notd gntiist jvtnd gntis and ology to traditionl taonomy. Thy influend Thodosius
Jopnlragn,ut his ral rk Dozhansky and Gorg Ld1,rdStins.Stins's Vrititllt d uol'utio
wship in 7927-1928 that took in Plts (1950) rought otany into th volutionarysynthsisand organizd
rith . B. Baok on th gn- th nw fild of plant volutionaryiology.
,ith Llarvy Monro Hall, a dis- lausn's own synthsislf plnt volution appard in 195] as Stgesi
'.itd him to partiipat in th the uoltil of PIt Spis'An imaginativ and pinring work, it was
l g w i t h t h C r n g i I I l s t i r u t i o n somwhat rif and intodutory in natur ausit ws asd on th Ms_
th mns y whih nw plant sngrLturs h had givrrarlir at ornll Univrsity. Th ook has gn-
using nativ lifonia flora. rally n ovrshdowd Stins'smor omprhensivarlir synthsis,
Fdri lmnts on suh ut nonthlss rmins on of th lassi works in th history of twntith-
l in aiifornia io 791, r.rtin- ntury volutilnayiology. Among its mor imporrant ontributions wes
tam that Hall had assmld th viw tht volution low th lvl of th gnus is rtiultdu to ph-
nomna sulr as hyridizatilnompanid y poIyploidy.
rgiInstitution of \/ashington In 1951, Clausn was appointd profssor of iology at Stanford. r_
:ludd taonomist David Kk tird in |956 and did in Plo, Alto, lifornia, in \969. H was survivd y
i d - 1 9 5 0 st h t a m o f l a u s n , his wif, Anna Har-rsn,who-lh rnarrid in 192\ thy had no hildrn' In a
Jsignda srisof l:rssial- rsarhposition for muh of his lrf' lausn did not lav rnuh of a lgay
r s t a r r d i npgl e n t v o l u t i o n u n - in trnrs of studrrts,ut his insiglrtful studis, iraginativly dsignd nd
pting som of th lratd mtiulously utd, am lassi xampls usd in narly all introdu-
krs suh as Gston Bonnir, tory txtooks of gntis'logy, arrd volutiorr.
Got ursson,who sought to
r, Kk, and Hisy prformd |BLIoGRAPHY
; thr distint lotions along
lausn,J. 1951. Stgsi th ulluti<l
lf Plt S1lt..is.Ith,
NY: orrrll
ln loation ws at sa lvl at
Univrsityrss.
t),and yt anothr at imr- Fnh,. s. l989. Jns hristirrIusn.Ntil Acdcntof Scines
rout 10,000 ft).Prforming Bilgrphil emoirs 58 7 5-107.
486 op
)
'ltivand ontfovrsil palon-
r. H playl :r lading rol irr
:sriingth nw spisdisov-
s alld th Arr-rrianShool of
:dismissdharls Darwin's s-
)rossof hang. H optd in-
tdalong prditalelins'
wzrlrhQuakr family. coP oN Fosstl litl o lv J&sY
surprisingly larg
row, JamsFranklin (.1'916)
lthal mutations. T}
Ovr th ours of a arr that spans ight dads,Jams Franklin row has ous ffts ar not m
mad svral sminal ontriutions to oth thortial (mathmatial) and - dividuals haror a
primntal population gntis. Althugh his primary rsarh organisms rssiv natur of 1
hav n fruit flis of th gnus Drosophil' row has also mad important sion. row ontinut
ontriutions to th study of humans and agriultural rops. Although row humans.
has workd on prolms ranging from th gntis of DDT rslstn to - Crow was th r
amining gns that violat ]Vlndl'slaw of fair miosis to dvloping thory Kimura dvlopd tJ
aout th bnfits of sual rprodution, his most prominnt ontriutions that th vast majorit
rvolv around th rol of mutations in volution. with rsptto sl
Born in January 1916 in ollgvill, Pnnsylvania, row grW up mainly asily gnratsman
.liihita,
in Kansas, whr h rivd his undrgraduat dgr from whthr and what f<
Frinds Univrsity, a Quakr shool. I 197, row wnt to th Univrsity of Laying th foundati
xas at Austin for graduat work, initially to study with prominnt gn- Crow to dtrmin
tiist Hrmann Josph (H. J.) ullr, who would latr win th Nol Priz this variation was tl
for his studis of X-ray mutagnsis in Drosopbila. Mullr, howvr, was to variation and ran
about to migrat to th Sovit Union' so John Thomas (J. T.) Pattrson . row dvlopd an
am row's major advisr. Just prior to th Unitd Stats' ntry into.!7orld that vry mutatior
\WarII tn 1941, row landd a faulty position at Dartmouth Collg, whr modl, th xpt
.W.artim
h stayd until 1948. and th drafting of his ollagus at Dart- was hetrozygousa
mouth ld row to add ourss as divrs as parasitology, navigation, and quals four tims th
mryology to his initial tahing rsponsiilitis of gnral zoology and g- mutation rat. rol
'Wisonsin'
ntis. In 1948, row movd to th Univrsity of whr h has r- ook, z lntrodut
maind for si dads. y gnrationsof st
Mutations ar not an unmid blssing.Although mutations ar th ulti- apparan in t970
mat sorrr of vriation and ar thus ndd for volution to prod, dl- In addition to hir
trious mutations far outnumr nfiial mutations. row and his studnts mad svral othr
usd th gnti thniqus availal in Drosophil melnogstr to masur studnt undr th t
th input of nw lthal mutations apparing on singl hromosom ah Austin, row was
gnration.From this valu, thy alulatd that th lthal mution rat pr isolating arrirs
gnti lous (gene)pr gnration was around thr pr million. utations study th gnti
nd not hav lthal ffts to b dltrious; in fat, row and his ollagus mappd largly to
hav shown tht nonlthal, dttaldltriousmutations aris at last 10 D. ldrihi-2.
tims mor oftn than lthal mutations. Rnt studis suggst that th rat row has also h
of mildly dltriousmutations is vn highr than thos mutations that hav Mndlism. Invest
ffts that an masurd in th laboratory. qual sgrgationh
Crow's group and othrs hav shown that most dItriousmutations ar part a fair gam.
almost ompltly rssiv;that is, th ffts thy hav in htrozygots mioti driv syst
are far smallr than th ffts thy hav in homozygots. Baus muta- Dr>sophilmelo
tions ar mostly rssiv,thy ar largly shltrdfrom sltion and thus at studnt, Yuih
ar not asily wdd out from th population. Ths dltrious muta- row, his studnts
tions, whther thir ffts ar lthl or mild, drasth man fitnss of gntis,and volu
th population that harors thm. ullr alld this rdution of fitnssth row's influn
.With
gnti load of th population. ullr and othrs, row invstigatd tion to th sors
the tnt of th gnti lod in oth Drosophil and humans, finding it to dirtly trained, rr
ro 489
row has ditd th ..Histoial Insts'mollusks, and hliratsd rustaanswith rgard to num-
. (As of mid- 2008, row has r of spis,ut rustaansar unquivoally th rnost morphologially
r of t National Aadmy of divsanimal group on arth. h fw, vry gnral faturslistd aov in-
nitrs fr th NAS, inluding adquatly onvy th rmrkal divrsity of a grrr-lpthat inluds wll.
rting. has srvd s hair of known forms lik lostrs, shrimp, as, and arnls, as wll as lss
of th dial Shool at th familiar amphipods, isopods' fairy shrimp' watr flas, and ostraods (or
..sd
s prsilntof th Gntis So- shrimP''), to nam ut a fw (s figur). In ft, th familiar forms
ompris only snrll propt>rtionof th roughly 68,000 known rustaan
spis.Thy ar pimarily a marin grop ut som also oup frshwatr
or moist trrstrialhaitats ('g.,trrstrialisopods, ommonly alld wood
. Sintifimri 74, no.2: li, pill ugs, sow ugs' or ..roly-polis,''and trstrialras). Intstingly,
th oughly 150 rnillion rd trrstrialrs of hristmas Island in th Indo-
nal virv.l Rt,'itulf G. aifi annually migrat from inland forsts to th s:l to spawn; mny mor
tims that numr of juvrrilsmigrat in th rvrsdirtion. Sonr aquati
Ioad. Oford Suru of
groups (.g., larg ranhiopods rfrrd to as fairy, tadpol, nd lam
: ls it a hlth risk? Prodigs shrimp) hav adptd to living in tmporv ponds or ..phmralpools'';
8380-8386. ths rustaansundrgo urapid iif yl and pr<ldudorrant mryos
n to tllultirlCtis Thor. nystd in a had pottiv as that n withstand tndd priods of
drying, only to mrgand ontinu thir dvlopmntwhn a pond rfills.
ss28,no. 6: 660-678 [doi:
Crustaansrang in siz from lss than a millimtr (minut zooplankton
lllstht n nr:rintindirr and nthi spis living within marin sdirnents)to th Yry larg. Japa-
ns spidr ras an hav [g spans ding 3 mtrs, or roughly 9 feet,
tfftingfitnss in Drosopil and lostrshav bn aught that d 20 kilograms (40 pounds). It is an-
9-78 -N../.
tiipatd that many rustaansawait disovry (at last as many spis as
th 68,000 spisurrntly known), spilllyupon ftrrthrplor:rtion of
vry rmot hiritats lik frshwatr nl marin av systms and th dp
sa-and nw findings ar not nssarily ptd to onfind to mr
s ar uiramians (insts,rnilli. spis. Aordirrg t artin and Davis (2001), almost 200 nw familis
rs, mits, tiks, and lrorssho hav n dsibd in th 20 vars that lrv lpsd sin th lst widly
s' arthopods ar Onsidrd to usd lassifiation,that of Bowman arrd Abl (1982). Rmarkly' an n-
l ommon anstor).and rus- tirly nw lass of rustaans'th Rmipdia' was disovrd in a martn
ringlysimpl qustion, ...Whatis av in th Bahrnas as rntly as 1981, and nw ordrs (Splaogiphaa,
.,and this is atlywhat mks Thrmosbanaa'Mitaa, ilnd Bohusaa)of minut forms (0'5 to 2 mil-
pods' rustaans:lr asiall), limtrs)of prarid rustnshav n disovrd in rmot avs and
d a stiff yt flil osklton the dp sa just sin th mid-1950s. Not surprisingly,som of tlrsnwr
J, or moltd, to aommodat ordrs ompris only a fw' rirr spis.\what is surprising is th rmarkaly
ftursdistinguish rustaans disjunt distriution of som of ths nrvly disovrd groups, sulr as th
:alld nauplius and had r- splogriplrirans-only our spisr known, disovdin singl avs
Lrfound two pairs of antnna in Brazil' South Afria, and nrthwstrn Australia! It is susptdthat ths
'f mandibular appndags, and populations psntrlis of an anint marin group that was on mor
d mailla-but vn ths fw widsprad;hangs in s lvls might hav ld to individuals tllrringn-
ik som othr artlrropods, th trirrdin, and ltr adapting to, grounlwtand a'',haitats.
th had' or phlon' may or ustaans td and rpr<lduin alrnost vry \4l1lyimginal for ani-
tions of fusd sgmnts,suh as mals. Som ar filtr fdrs and siv atria or tiny zooplankton from
.g., a lostr's..tail''), and ap- thi Watry nvironmnt' othrs savng for thir food, nd lany a
of soilization. prdtos. h lattr mer1,h:rr,s()m rInkl adpttions, srrh as th
492 rsts
..lalv'' .
*:;;;.Sflffi'v
r1,-|],\.
modifid
smsh mllusks
rustilan hs a t
) ('.,' gthr, produ
'r @I1
d -_--,' shrimp''). Th sol
.g
stun or vn kill sr
also notl for >
mits. Nutnrltts
spollg lll1d f()rm
phnornllon of
ntly mmtt
Ss ar usu
known) nd su
as watr fl:rs,wil
as parthnogns
vlop frorn unfrti
nvironmnt o s<
swimmir-rglarva 1
morphosirrg intl t
spis, sh as f
mrg (dirtdv
Th rlv rrr
liotl ,virrsgo) sa
xhiitillg n inr
this ..plosin''O
tonishingi bif l
phyla hav long si
an quivalrlt[rllr
a topi <lfd:rt
plosion tlf ody p
haps gratr oxy
mtaboli rtsan
gn.dupliation t
what wr tl-rnr
s
progrms <lfanim
Rprsntativs of som si rr-rstantyps. () opoda; () ostrod; nonlthl l-najor:
() nostraa; (d) Notostraa; () ladoa; (f) Lptostraa; (g) Arraspidaa; mnt onstitutdi
(h) irripdi; (i) Stomirtl1.lod;
(j) ysida; (k) Tanaidaa; (l) lslpo1a; filld by adapt
(m) Amphipoda; (n) uplr;rusiair;(l)Dndrrnhiat; (p) arida;
an adptiv radiat
(q) Astaid; (r) Anomur; (s) Ahlata; (t) Thalassinid; (u) BrahyLrra;
Fossil vidnr
(v) Polyhlida. (From Polr 2004, Figur 2.)
suphyl (or arth
taly, on oi th ft
vi]nthat rus
this tirlr' Th las
or ..sa r-nonky
som of th ail
49
modifid ..law,, of a stomtopod mantis shrimp tht allows .^:':
smash mollusks or spar fish with grat for and spd. Anothr typ of
rustaanhas a rndifid lw with pinrs that' whn quikly rought to'
gthr, produ loud snapping nois (hn thir nam, ..snapping
shrimp''). Th sond is assoiatd with a ..shok wav'' strong nough to
stun or vn kill small pry, suh as fish. Som spisof snapping shrimp ar
alslnotal for hiiting soia1havior mor typial of ants, es,and tr-
mits. Numrous offspring of a singl fmal ( ..qun'') will oist in a
spong and form asts to a for th ylung and prott th olony. This
phnomnon of usoiality in snapping shrimp ws dsrid rltivly r-
ntly y mmtt Doff (996) and is a first among rustaans.
Ss ar usually sparat (although som hrmaphroditi spis ar
known) and sual rprodution is th typial mod. Som rustaans, suh
as watr flas, will asionally rprodu asxually via a mhanism known
as parthnognsis,whry fmals will prldu young (also fmal) that d-
vlop from unfrtilizd ggs. rustaan ggs may rlasd into th aquati
nvironmnt or so.tims roodd, ut th young usually mrg as fr-
swimming larva that will undrgo a sris of molts and stags for mta-
morphosing into th adult form (indirt dvlopmnt). Th fmals of fwr
spis, suh as frshwatr ras, rood thir ggs until miniatur dults
mrg (dirt dvlopmnt)' molting as thy grow largr.
Th arly amrin priod of gologial history (approimatly 550 mil-
lion yars ago) saw th mrgnof numrous multillular animal phyla
hiitirrgan inrdil divrsity of ody plans. Palontologistsnow liv
this ..plosion'' of lif forms ourrd ovr only 5 to 10 million yars (an as-
tonishingly rif amount of ti"l,volutionarily spaking). any of ths
phyla hv long sin gon tint, and nvr again in th arth's history has
an quivalnt urst of volutionary innovation ourrd. Th rasrnrmalns
a topi of dat and spulation.Possilontriutors to th amrian -
plosion of ody plans inlud hangs in th arth's gohmistry (.g.'pr-
haps gratr oygn availaility favord marin organisms with highr
mtaoli rts and lrgr sizs)and suddn, larg-salgntihangs (.g.,
gn-dupliation vnts and a rshuffling of gn rgulatory pathways in
what wr thn rlativly simpl organisms).Thraftr' prhaps th gnti
programs of animals had rahda lvl of omplity that prludd furthr
s. (a) oppoda; () Ostralda; nonlthal major hangs. Anothr viw is that th pr-amrian envlron-
f) Leptostraa; (g) Anaspidaa; mnt onstitutdn ologial taula ras of mpty nihs simply waiting to
<)Tanaidaa; (l) Isopod;
filld y adapting organisms. Thn' as now, suh nvironmnts Dromot
l n h i t (l p ) r i d ;
an adaptiv radiation lf spis.
halassinid;(u) Brahyu;
Fossil vidnonfirms that rustaansWr on of numrous arthropod
suphyla (o arthropodous phyla) prsnt in th amrian and, mor no-
taly' on of th fw that hav survivd to this day. vn mor striking is th
vidnthat rustanshad alrady divrsifidinto svrlmain lineagsy
this tim. Th lasssOstraoda (sdshrimp), Branhiopoda (rin shrimp,
or ..s monkys''; fairy, tadpol, and lam shrimp; and watr flas), and
som lf th aillopoda (spifiallythostraans
[arnalsIand wormlik
494 rustns
form; diffrnt urrowing spis may ppar morphologially similar ut pattrn thory,'' also r
not shar a rnt ommon anestor. volutionary affinity among spis irr losr affinity to t
may also misonstrudif any of thm has undrgona rent volutionary all thr viws onsid
rvrsal to a mor anstral morphologial fatur. Th DNA approah to and omplling mol
infrring rlatdnss avoids th problms prsntd y ths instans of rustaan nstor' I
,
morphologial homoplasy. Thrmosanaans r minut rustaans found som ar alling th
in groundwatr and springs' and they hav variously n plad within th Sustarrtially mor da
suprordr Praarida (in part austhy hav a rood hamr, a fatur th National Sin
found in som praarids)and in a ompltly sparatsuprordr,th Pan- Arthropod Phyl<lgn
arida (austhir rood hmr has a distintiv dvlopmntl origin 120 diffrnt gnsfrt
and loation). Similarly, mysid and lophogastid shrimplik rustaanshav pnding and r high
bn onsidrd ithr to ompris an ordr of praarids or to sprat
ordrs and ludd from th rarida. In ah of ths ass, DNA vi- BIBLIOGRAPI
dn has n usfirl in rsolving long-standing phylognti unertainty Bowman,T. ., and L.
and has shown that morphologial faturs (.g.,th rood pouh and fa- l,. G. Al,d.,S1
turs of th arapa' forgut irulatory systm' and ys) shard y som tsiologof rust
so-alld praarid spis may in fat hav volvd indpndntly or n a l m a n ,W . T . 1 9 1 1 .
Duffy, J. . ] 996. us
lost ntirly.
orty,R. A., and R.
Som rustaan spis (.g.,on ntir infralass of rustaans rlatd to ssoiationSpi
arnals, th Fatotta or ..y.larva'') ar known only from thir larva N{artin,J. W., and G.
(adult forms hav nvr n found). othr spies ar known from only mal rL|std. Sin
or only fmal forms, so knowldg of ky morphologial faturs is similarly Anglsounty.
laking. In th mphipod family Luothoida' mls may undrgo a adial Poor,G. . B. 2004..
l dtifi tiltt' 'ol
transformation during a singl molt, suh that pr- and post-molt mals wr
Shrm,F. 1986.rus
rronously lassifid as not just diffrnt spis ut as longing to diffrnt
familis. Individuals of som spis of aridan shrimp ar squntial hr-
maphrodits, ginning lif as mals and latr transforming int fmals (a
uvir, Grgs
ondition trmd protandry). Taonomi rognition and assignmnt is sp-
ially ompliatd y a spisthat has supr-malswith nlargd gnitalia Although Gorgs
(ths nvr transform into fmals) in addition to th mor typial mal- playd a prominnt r
phas hrmaphrodits that will vntually transform. In all of ths ass voluntarily. In rtros
DNA vidn is somtims th only mans y whih som spis an lishd yond doubt
idntifid auratly and plad within an volutionary hirarhy. that this had ourr
Finally' in addition to th intratility of dtrminingrustaanrlation- drmind th ida of
ships, th qustion of rlationships among rrrstaansand th othr major lassify organisms ir
tant arthropod suphyla (Hxapoda, or insts;Myriapoda' or ntipeds philosophr ihl I
and millipds; and hliriforms, omprising spidrs, mits, tiks, and volution than Jan-
horssho ras) is prpling, for many of th sam rasons notd aov. tion thory. owvl
Varius altrnativphylognishav n proposd on th asis of diffrnt uvir's allgd ont
intrprtationsof morphologial vidn.For instn' th ..uniramian hy- tt. som of uvir
pothsis'' onsidrs arthropods to polyphylti (i..' unlikly to hav had a th sin of volut
singl, ommon anstor) and sparats hlirats (.g., spidrs) and rus- drstood in light of
taans (possssingiramous appndgs)from uniramus insts' myriapods, uvir was primr
and onyophorans. Th ..mandiult hypothsis'' taks a monophylti (sin- him was lassifiati
gl anstor) viw of arthropods and units rustans' insts, and myri- prviding unifid
apods within th andiulata. Yt a third viw, asd on an ..arthropod was losly rlatd t
uuier 499
svralsintistslaimd that
s p i sB. u t r h q u s t i o n r -
d disppardor still isted
who dsid hug nutnr
)xtintion.uvir prop<lsdto
:rrstrial nimals as a tst. In
i m . r i s o u l dh d I y m a i nu n -
dtaildproof of suh a as.
y, h idntifida giant fossil as
Ildthis niml t'gthrim.
irs disivlysupportd y th
-
nnt to a known family (th
,'t-
rt th disapparanof num-
mastodon,Plotrim, and Prt of Gorgs uvir's argumnt agtrinst volution was sd on th
:r, t.) irr diffrnt prts of murrnifid organisms rought ak y Napolon's sintists,whrl went rvith
him on his amp:rign tl gypt. his is th skltlrrof n iis, whih uvir
I r.rnprdntd prision, h
pointd out is idntil to th skltons of iiss tlda,v,thus apparntlv showing
animals no longr istd.
tht thr is no h:.rngeovr tim. (Fom uvir 1804.)
'othssould aount for th
rwhn l-amark gan pu-
:r statd thy had lr ithr disappardin th nt stratum. Futhrmor,thy osrvd an altrntion
3uvir rjtdth modifia- of frshwatrand marin forr:rtions.This ould b aountd for y sus-
to vidnto support it. H siv vnts of lvation and dprssionof th ontinntal aras. Baus th
, h o u g h t h r w a s ' " . i d n i n juntions twn th two kinds of strata Wr rthr sharp, uvir on-
rr gypt y tinn Goffroy ludd that suddn gographi hangs had n rsponsilfor th tin-
'ovidth slightst
lu of th tion of sussivtrrstrialfaunas and for a rpopulation by xisting faunas
th psnt(sefigur).If no frorrr othr plas.
uld Lamark and ohrs - ..uvir,s
As shown y artin Rudwik (1976, 117), rjtiorrof volu-
tin-r? In ntrast' uvir and tion . . . was primarily a dfnsof titio' not of spial crtion.,' :u.
sussivstrata aov th vir's gnral rprsntationof th history of lif onsistdin a srisof loal
lt disapparanof loal fau. rvolutions,whih thmslvsimplid th tintilnof som spisand th
ntrs),thr was a onstant migration (in and out) of othrs. nvr mntiond ir gnral atastoph
its htristifossils, that that ould hav dstroydall xisting spison arth. Suh a h,vpothsiswas
t02 uuir
BILIoGRAPHY
uvir,G. 1804.6moirsur Llbis dsninsgyptins.
lsdu usum
tiold' HistoirtrIl
A(1804):116-1.5.
uuir 50
ens.lsd usm
known and appriat
as a man of solid wort
ntifi laims a fair an
Darwin's first systn
h was traind y ar
th world is formd 1
tastrophs),th grat
Lyll' whos thr-volt
grly dvourd y th
thsis, arguing that tht
Darwin, Chrls (1809_1882) Lyll hypothsizdtha
harls Rort Darwin, th nglish naturalist and volutionist,was orn in ing transitory and a
th town of Shrwsury on Fruar 12, 1809, th sam day as Araham th distriution of lan
Linoln aross th Atlanti. H did on April 79,1882, at his hom in th vil. kind of watr d, wh
lag of Down (Knt). H was th author of o th origi of Spies' pl- rings on lvation in
lishd in 1859, and h is onsidrdth fathr of volutionary thinking. Darwin aptdth
.
Darwin's patrnal grandfathr was physiian asmus Darwin, author o{ in the Lyllian mod.
Zoonomi (1794_7796), on of th arlist works prolaiming volution. islands? Lyll argud t
His fathr' Rort Darwin, was also a physiian as wll as a mon man' ar- argud that it was im
just rak th sa surf
ranging mortgags twn aristorats with land and a nd of ash and
usinssmnwith ash and a nd of saf loans. Darwin's matrnal grandfa. lands nd thn thy si
thr was Josiah \Wdgwood, th man rsponsil for industrializing th pot- th sa surfa,ven a
try trad in Britain. Darwin mntd th !7dgwood onntion furthr pltly nth.
.W.dgwood. harls Darwin's g
whn h marrid his first ousin, mma and his wif had 10
hildrn, svn of whom livd to adulthood. on was Gorg Darwin, th In Gln Roy, a small v
minnt physiist and authrity on th tids. indntations around t
harls Darwin was duatdfirst at Shrwsury Shool, on of ngland's as thy wr knowni
lading privat shools, and thn snt to dinurgh Univrsity in th foot- around wat that on
stps of his fathr and grandfathr, intnding to train as a physiian. Un- lat 1830s, Drwin a
h,ppy in this dirtion' h nrolld at hrist's ollg in th Univrsity of ara had mvd upw
Camridg, intnding now to an Anglian lrgyman. Whil at univrsity' th Swiss sintist Lo
oth in Sotlnd and ngland, Darwin startd to show a strong intrst in watr Was a lak, da
sin,and y th tim h graduatd in 1831, senior sintistshad startdto appriatd his mista
spot oth th intrst and th talnt. Th sustantial Darwin-!7dgwood for- thinking in a atastr
tuns mant that Darwin nvr had prssingfinanial ndsand ould always of God.
indulg his passion for mpirial inquiry. Darwin would not I
Divrtd from th lrial arr y th offr of a voyag on HMS Begl, As a young man prpz
undr th aptainy of Rort Fitzroy, Charls Darwin spnt fiv yrs going tsgl voyag his lit
around th oast of South Amria and vntully irumnavigating th strongly in a God wh
glob. Although h was ngagd s a gntlman ompanion to th aptain, trfr with his rati
Darwin soon bam th full-tim ship's naturalist, aumulating xtnsiv th writing and puli<
olltions that h shippd hom' thus adding to th alredy good opinions lik many mid-Vitor
of his Camridg mntors. Darwin lso mad tnsiv travls inland. Th ism. h Anglian b
diaris h wrot on th voyg Wr th foundation for a popular tavl ook aus it insistd that (
titl'edTh Vog of th Bgle, first pulishd as part of th offiial rord of of organisms, suh at
th ship's travls and thn indpndntly. Thr is no dut that ing Arhdaon William I
-t04
Dri, hrls 50.'
rad as an undrgraduat, Darwin livd that th most signifiant fatur thinking pulily. Th rt
of th living world is tht it is ompl and dsignlik,t th nfitof indi- also (aftr a dd or t
vidual organisms. ism. lWhat is lar is th
Lyllian gology dirtd Dwin to thinking aout organisms; fossils wr tion than for Darwin's <
vital vidn of subsidnand lvation. Thus primd, in 1835, whn th to dny th lttr outr:
Begl rahd th Galipagos Arhiplago' a group of islands in th Paifi, auss) ranging fom L
)arwin was astoundd to larn that th dnizns, partiularly th irds and tis) to saltationism (r
th giant tortoiss' r diffrnt from island to island. volution from a om- som kind of guidd v
mon anstry is th ovious answr' spiallyto sot-neonwho livsthat Darwin had intnd
God's glory is shown through the working of law. Howvr, it was not until would now writ, filling
h rturnd to ngland in th following spring of I87 nd was onvind only on of ths oo
that th Gal6pagos varitisa truly diffrnt spisthat Darwin mad th Plts dr Drllesttc
lap to tansformism, as it was thn alld. inadquat hypothsis
Darwin rea|ized tht h ndd a mhanism to plain his nwfound - that partils ar givn.
lif. As a graduat of th Univrsity of amridg, h was awar of th tri- y h s r g a n s .T h i s
umphs of its grtst sintist, Isaa Nwton. Darwin thrfor sought for to whih Darlvin alwa
th organi world a kind of quivalnt of Newton's univrsal law of grvita- Apart from a sris
tion. H found his answr in Sptmr 1838. H startd with th althu- tl ook on orhids, a
sin lulation that populion prssurswill tnd to outstrip supplis f plants, yt anothr on
food and spa. H notd that thr smsalways to b a sour of variation yars Wr on th hum
in natural populations. Hn, arguing analogially from th susssof th and a kind of supplm
rdrsof animls and plants, Darwin onludd that th will e a natu- mls (|87z\.In th or
ral form of sltion, with som winning and othrs losing, and suss ing aout humans, othr tl
a funtion of th winnrs' puliar faturs. Vitally, this natural sltion, o that h thought his th
..survival th topi unintrsttng
of th fittst,''as it am known, lads not just to hang but to
hang in th dirtion of adaptation. thory on th tal ant
Darwin soon Wrot up his thinking as a skth of a thory' ut for rasons mnt about humans.
still undetrmind h did not pulish. Undoutdly a factor was that h fll In this,Darwin was'
sriously ill nd for th rst of his lif ws an invlid and a rlus. (Th x- th impliations for Ho
t aus of th illnss is unknown. Som spulatthat it was psyhologial, pl wantd to talk a<
th strss of having suh an volutionary thory. Othrs think it might hav prssur from frinds a
n th rsult of n illnss,hagas disas,pikd up on th Bgle voyg.) that lWalla had on
For mny yars, Drwin sat ol.l his volutionary thinking and was divrtd to plain hulnan vo
into a massiv taxonomi study of arnales. physial and soial, rs i
.!alla,
Finally' in Jun 1858, Alfrd Russl a young naturalist witing tion was th ky ausa
from th Fr ast, snt Darwi a short ssay ontaining th sam ids Dar- ary mhnism, sxu
win had 20 yars for. Thus spurrd, trats of Darwin's thinking and Darwin livd that l
.lfalla's mining th diffrns
ssay Wr pulishd at on. Darwin now rapidly Wrot up his
thory, pulishing o the origitl of Speist th nd of 1859. Although h did not
Te rigin was immdiatly ontovrsial, ut it is important not to xag. last dad.H ontinu
grat this ft. In Britain and n th ontinnt, and vn in Amria, many growing orrspondn
popl, oth sintissand th gnral population, quikly aeptdth ida rs ross th wrld as
of volution. Darwin himslf hd laid th groundwork for this y arfully with rspet and aff
ultivting mn-notaly anatomist Thomas Hnry Huly and otanists h did' y gnral a
Josph Hookr in Britain and Asa Gray in Amria-who would dfnd his from Nwton and a p
Dri, hrls 507
th most signifiant fatur thinking pulily. Th rligious tndd to mor rlutant, ut thspopl
lik, t th nfit of indi- also (ftr a dad or so) mst proaly aptd som form of volution-
ism. What is lr is that thr was onsidraly mor nthusism for volu-
ut organisms1fossils wr tion than for Darwin's own mhanism of natural sltion.No on wanted
primd' in 1835, whn th to dny th lattr outright, ut most wantd to supplmnt it with othr
up of islands in th Paifi, auss' ranging from Lamarkism (th inhritan of aquird haratris-
s' partiularly th birds and tis) to saltationism (volution y instantanous mjor rrw variations) to
land. vlution from a om- som kind of guidd volution whr God plys a ontinuing and ativ rol.
) somonwho livs that Darwin had intndd t'e rigi to a tsr for a sris of ooks h
v. Howvr, it was not until would now writ, filling out in dtail all th laims mad rifly in te origi.
of 1837 and was onvind only on of ths ooks was vr writtn, Th Vritio of Aimls d
lis that Drwin mad th Plts under Dlmstitil (1868\, notl hifly for Drwin's somwhat
inadquat hypothsis aout hrdity, alld pangnsis. (Darwin argud
to plain his nwfound - that partils ar givn off all ovr th ody and olltd in and transmittd
g' h was wr of th tri- y th s ogans. This would thus nsur Lamarkian ffts, hypothsis
)arwin thrfor sought for to whih Darwin always susrid.)
l n . su n i v e r s l w o f g a v i t a - Apart from a sris of inidntal works that Darwin now authord-a lit-
{ startd with th althu- tl ook on orhids' anlthr ln liming plants, a third on instivorous
tnd to outstrip supplis of plants, yt anothr on arthworms-th major laors of th post-rigi|1
,s to souof variation yars wr on th human spis,rsulting in The Desnt of (1871)
Llly from th susssof th and a kind of supplmnt,he prssio of th motios i d i-
d that thr will a natu- mls (7872|' |n te rigiz, dliratly, Darwin had said virtually nothing
rs losing, and suss ing aout humans' othr than an unamiguous ommnt at th nd suggsting
lly, this natural sltion, or that h thought his thory also applid to humans. It was not that h thought
.ds not just to hang ut to th topi unintrstingor unimportant' ut h wantd first to gt th si
thory on th tal and not gt vrything at on lost in th gnral argu-
r of a thory, but for rasons mnt out humans.
:dly a fator was that h fll In this, Darwin was only partially sussful.At on, vryon sizd on
valid and a rlus. (Th x- th impliations for Homo spiesof th ..monky thory.''This is what po-
rtthat it was psyhologial, pl wantd to talk aout. For a whil, Darwin stayd aloof. Thn, undr
. othrs think it might hav prssurfrom frinds and supportrs,and prtiularly glvanizd y th fat
d up on the Bgle voyag.) that \/allahad om a spiritualist and had turnd to non-natural auss
y thinking and was divrtd to xplain humn volution, Darwin argud in dtail that human natur'
physial and soial, is a produt of unrokn law. As always, natural sl-
' a y o u n g n a t u r a l i s tw i t i n g tion was th ky ausal featur, ut Darwin supplementdit with a sond.
:rtainingth sam idas Dar_ ary mhanism, sual sltion, whr organisrs ompt for mats.
ts of Darwin's thinking and Darwin blivd that sxual sltion was partiularly important in dtr-
n now rapidly wrot up his mining th diffrns twn human ss and ras.
r ndof 1859. Although h did not liv to a vry old ag, Darwin's halth improvd in his
t it is important not to xag. last dad. ontinud to work until th nd, inluding answring an vr.
' and vn in Amria' many growing orrspondn.Aknowldgd both y his ountrymn nd y oth-
ln, quikly aptd th ida s aross th world as On of th grat sintists of all tim, he was rgardd
:rdwork fr this y arfully with respet and afftion. hr Was thrfor no grt surpris tht whn
{ n r yH u x I y n d o t n i s t s h did, y gnal aIaim,h was urid in $7stminstrAy, just aross
ria_wh would dfnd his from Nwton and a pa or two away from Lyll.
508 Dri, rsmus
r Theories of ind d Be. mahins. mphsizing th univrsal appliation of harls Darwin's funda-
ophy of history-a sltion mntal ir-rsights,Dawkins drW attntion to th possiility th;rt whn gnti
rt th gnalogy(who got what naturl sltion ratd a spis,Hoto spies, with an xtndd phenotyp
ruting onptual gnalogis that inludd languag and thnology' this in fft stablishd a nw mdium
rtl for similarity of idas, and of volution y natr'rralsltion:human ultur, in whih th diffrntial rpli-
flit, thy sid with similarity. ation of salint ltual itrr-rs'alld ms' uid aot-tntfo rrrany of th
sition modl of sintifi faturs of ultural volution that ar Othrwis prpling. Dawkins initially
ry is all th ttr for it. prsntd th id of mms as a sort of thought xprimnt to illustrt th
bstratnssof th fundametal ida of natural sltion. nd h has not
laimd to making a major ontrir-rtionto th sintifi invstigation of th
gompioto Drtui. volution of humn ultur. Othrs, howvr, hv takn up th onpt n-
thusiasti:rlly,with rid rslrlts to dat.
{ ullutionrTboris of ind
In 1986, Dawkins pulishd anothr pdggialtour d fore,h Blid
Prss.
phologiclonstructiod Wthkr,in whih h illr.rstratd many of th most lusiv impliarions of
r,.hiago:Univrsityof higo volutionary thoy via his pionring Blind \Wathmakrsoftwar, on of
th rly triumphs f th Artifiial Lif 1ovmentin thortial iology' In
rtation:pogrssivityand 1995,e was app<lintdto th first Charls Sirnonyi Profssor of th Pu-
. andJ. ainshin,
ds., li Undrstanding of Sin t Oford. In this idal rol for a writr of his
l ambridgUnivrsityPrss.
't Hu Purpos?mbridg' talnts,lr has susquntlypulishd a srisof luid ooks and atils-
plaining sptsrfsinto h gnralpuli. L{ also has takn tlr lad in
d Lrni g:'', u o.:([:,n. th puli disussions of ontrovrsial issus of sin poliy and politis.
"'
Most rently, his roadsid on rligion, Th God Dlusio (2006), an at-
tmpr to rais orrsiousnssout th waknsssand follis of traditional
rligious lif,has attratd oth nthusiastipris and vilifiation' playing
a lading rol in what is bing alld ..thnw tlrism.''Lss wll known to
llutionry thorist bst known th lay puli r Dawkins's important analyssof fundamntal prolms in
ld phnotyp, and th mm. kin sltionand th volution of ornrnuniation,among othr topis.
ford, whr h has livd and
z is on of th lassi works of |BLIoGRAPHY
lrging onsensus mong suh Dawkins, R. 1976, Th SIfsll G. oiord: oford Univrsit1'Prss.
lorg!illiams, Rort Trivrs, 1982.Th tededPhotp.Oford: Oford Univrsitrss.
or' thologistNiko Tinrgn' 7986. Th tslid WtbmaAr.Nlv York: \W.W. Norton.
;tion in sh vivid nd ssi- 204.Tb ttt,stlr's
?ll/c.Boston:HlughtonNlifflin.
206.h (]ldDlsitl.Bostorr:ughton ifflin.
as mrly a ..popular'' ook.
Grafn,A., nd M. Ridly,ds.2006.RihrdDtukis:Htttu Sitist hgd
lamntal, and hn most - th W |XtTik. ofrd: Oxford Univrsityrss. -D.D.
ral sltion is th ..gn's-y
:oups, and vn individual or-
rry nfiiaisof all adapta.
The Descent of NIn, nd Selectio in
:, volution dsigns and uilds
'litionsof gnsy improving
Reltio to Se (harlsDarwin)
Lsdos not stop at th skin of Tbe Dst of , d Sltili Reltio to Se, pulishd in 1871,
d onstution of an xtndd was hrls Darr.in'swork on th evolution of th human spis.Initially,
mnt nd vn othr spis. h hd r"rnwillingto gt into this ast of tlr volutionary dbat, ut
iust as importnt parts of th whn Alfrd Russl !7all (1870), who had om a spiritualist, dlard
loths,and parasitsoftn - that humans ould not hav n produd y natural mans' Darwin flt
hijking othr gens' survival that h had to gt involvd.
514 Th Desntof n
llution is original, as Drwin r- promoting an unsuppotal tho|ogial thsis. Although Darwin is vry
hrs.Lik Thomas linry u- Vitoria' he is a vy liral Vitori:rn. For all that h worrid aout th
r' 186) for him, Darwin's fftsof mdiin, h immdiatly strssdthat lr did not think w should
nans with othr nimals' show- stop using it. H also thought that th hif rasorr whit popl tnd to
n o might imagin' Thrfor, ovromothr popls is baus othr popls suum to th disassof
) not part of th natural world, whit popl mor rpidly than whit popl do to th disass of othrs.
nals,ruh rason to think that or gnrally' although thr indd pssagsin ei Kmpf (1925)
rot think that humans wr de- that do sound lik Soial Darwinist' Hitlr drw somwhat haphzardiy
s inlindto support th ida of on many sours' and th nti-Sritism of his youth in Vinna nd th
.Wagnr
Volkish sntimnts f th group promoting th mm<lryof wr at
lan asonand soiality. H ar- lastas signifiant,or mor (Fridlandr 1997; Krshaw 1999).In an as,
outomsof volutiorr.Bause th Nazis gnrally did not muh lik volution aus it strssdth on-
naturl sltionwoks lways n s so f h u m a n k i n d , i i n k i r r gA r y n s w i t h J w s a n d G y p s i s . ( R i h a r d s 2 0 0 8
th group, h was onrnd to disusssths hargs' putting thm in ontt.)
lthrsould nvrthlssvolv.
|LIoGRAPHY
Lndof sltionmight hav pro-
]stion that morality ould Darwin,. 1871'.The Dstof , lld Seltilli RItil,oSe.London:
th rodrnmhnism of ..r- Iohnuray.
plld:No lntllignclltl'd,2008.NthrrFrnkowski,dirtor.90 mirr.
N.p.:PrmisN{diaorporation.
r odd natur of Darwin's ook,
Fridlandr,S' 1997. Nzi Germd the |es.Vol. 1, Th Yrsof Prseution,
at ll. It was rathr aout Dar- 193_9.London:Widnfldand Niolson.
hang,that is, sxual sltion. Grubr,. . 1981.Dri o .2nd d. hiago:Univrsitof hiagoPrss.
mts' and hn thy dvlop Hitlr,A. |925. in Kmpf.Munih:VrlgFranz hNahfolgr.
rthsof th pok) to aid in Huxl,' H. 1863.uir'lc s ttl 's Plcei Nture'Lon]on:l/illimsand
Nrgat.
:s lik human hirirlssnssand
Krshaw, |. 1999.itlr.o|.1,1889-1L).J6: I]ubris.Nr'York:Norton.
rdy natural sltin and so Rihards'R. J. 2008. h Trgi Sseof Life: rst lIhld th Struggllur
n rplid that sxual sltion uolutiorThoght' hiago:Univrsityf hiago Prss.
nnd ody shap. For many Rus,M. 2008. brlsDnui. Oxford: Blakwli.
ual sltion as somthing d- Trivrs,R. L. 1971.Th volutionof ipro:rl altruism.QurtrlRuituof
Lrglrtof it as sornthing st in- Biohg 46: 35_57.
Wll,A. R. 1870.Th limitsof nattlrlsltionas applidto man.ln his
ris of intrstin nimal soial
otributios to th Thorof NturlSltio, 332-71' London:
of gratthortial and mpiri- amillan.
Wikart,R. 2006. From Dui to Hitlr: uoltiorthis, ugis,d
las ld to mak som vry on- Rismi Grm\,,Nw York: Palr]rar. \4amilln. -.R.
iorityof mals ov fmals,
lso worrid that modrn mdi-
rn rdingstok, ut flt al
The Deuelopmet of Dui's Theor (Dov ospvat)
sud popl of ultur whos Th Deulopmet of Dri's Thor ws pulishd in 1981. It appard
of ths sorts of rmarks, th justa fw months aftr th dath of its author, Dov Ospovat, at th ag of 33.
t h r a t i o n i s tas n d t h i s s o - Th ook ahivd an important milston in what am known as Dr-
:orists)' frustratd in thir at- win Studis,tlr histoil study of harls Darwin's lif nd work. Darwin
lifsar gnuin sien, hav Studisam a spial ara in th histlry of sirrshortly aft th 1959
in lads straight t th vil id- ntnnialof th puliation of hrls Darwin's o the origin of Spis.It
y National Soialism (s sp- soon grw into a mior fild of historial rsarh.Th xplosiv growth was
-nrratd fr|m plld: No due to two fators' Fist, until th 1950s Darwin was l.lotrognizd as th
i arnpi of d sholarship xrmlyimportant siIrtisttht h is today. Th modrn synthti thory
516 Th Dvlopmntof Darwin's Thry
lf volution (th thory fomd v omining Drrvinian natual sltion This nw viw lf Dar'
with Mndlian gntisand, m<rrrntly' with rnolular gntis)gan pholog and mryoltlg
during th 1930s nd 1940s. At Iirst it dm<lnstrtd tht a numlrr of diffr- ginning in th 1980s, wl
nt influns' inluding natural sltion and gnti drift, ontriutd to and mbryologial dvlo
volutionary hang' ut th rlativ strngths of ths influns Ws un- th stag fo roadr his
known. By th 1950s it Was gnrlly ptdthat r-raturalsltionwas th gathrd y historian D;
dominant voltionry for. This onsnsusoinidd nily with tlr l- Th D r'itti Her itg
rtions of th ntnnil of the rigi. B:lt'ts Drwin Was th fathr of portant was Ospovat's s'
natural sltion,h now am th fathr of th synthti thor of volu- o m l n i t t d t o t h n o t i r l nt
tion. Th nw interst in Darwin oinidd with th disovry tht Darwin jor intrst to historins
hd lft an immns sto of prsonal paprs that inludd prsonl orr- 2 0 0 3 ,2 0 0 4 ) .
spondn, rsrh nots, and prliminary drfts f his ooks and artils.
Urrlik Gr;orMrrdl (rvhospps wr dstoydy lris stlssor))Dar- lLtoGRAPY
win's paprs offrd immns opportunitis fo rsarhin th histlryof si- Amundson,R. 200.5.TDe/
n and, indd, in th lif story of a sintist whos importan was Rlotsof uo-duo.al
suddnly ing rognizd. Burian, R. . 2004. list
Th first wav of Darwin Studis onntrtd on his voyag on MS tis. amridg:lr
K o h n , D . , d . 1 9 8 5 . h D
B g l a n d h i s n o t o k s o f 1 8 7 - 1 8 9 , w r i t t r rs l o n a f t h r t t r r r r d .
Prss.
T.his priod ovrd th girrnirrgslf Darwin's volutionary thought and O s p o v tD , . 1 9 8 1. T b D e u
his dvlopmnt of th prinipl of natural sltiln.This phas of Dar- UnivrsityPrss.
win's lif was importnt to sintists aus natural sltion was th as- R i h a r d sR, . J . \ 9 9 2 ' h N
pt of Darwin's thought that fit most losly wirh th synthti tlrory of d ldollgiIR ls
hiagoPrss.
volution. Th priod from 1840 to l859 was oftn rgardd as a pr-rzzl. If
200, he Rlmtltt
D r w i n h a d d i s o v r d n t u r l s l t i o r r f o r 1 8 4 0 , w h , v d i d h d l a y of Glth. l.rillgl: LII
pulishing until 1859? ospovat;lrovidd an llswr to that qustionnd' 2004.MihalRus
simultanously, gan th study of Darwin as a lnAn lf his ag-a Vito- 2s-38.
rian naturalist. Rus, . 1996. od to ]
arnridg, MA: rv
Ospovat was bl to provid this nw prsptiveaush was tlr first
2004.h r<lmatt
historian to gain assto Drwin's paprs from th so-alld lak <l,th
isttt lf tsiolog.r'7:
st of portfolis that Darwin had ssmldin th 1850s in ppratiOnfor
writing his major work on volutiln.(This work ',vas,of ours' intrruptd
.Walla's
y Alfd Russl odisovry of natural sltiln,and the ri.
Dinosaurs: Th mor
gi lf SpiesWas th rsult. S th main ssay ..Th History of volutin-
ry Thought'' by ihal Rus in this volum.) Th fist pratitionrs of Bas of thir immt.t
Darwin Studis hd sn ntural sltiorras Dirrwin's major lrivmnt first ol to mind wl-r
and intrprtdim as uniqu gnius.Ospovat, in ontrast,dpitd him as pld with th fat that
having bnfitdfrom wid adingsin th sinof his day, inluding pal- ntury ngland) wlrr
ontology' systmatis,morphliogy, and mryolgy. This plaind Dar- t h i r h i s r o r i e l l y i n gu
win's hsitation. vn though n.rtural sltin ws in pla, Darwin's t t , d i n o s u r sh a v p I
vidnthat volrrtionhad atuall .ltrrrd Was rlarivlwak in tlr rly our volution. Nvr
yars' It was gradually strngtlrndduring th two dzrdsl>for1959. d v l o p i n g m o s i
..Darwin was and undrstandingvol
th first rputal lraturalist to argu for transmut.ttion who
was l to tak advantagof th dvlopmntalviw of natllr that mrgd Dinosaurs ar a unlq
in th mid-nintnth ntury. This, as nruh as lris nw thory [natural thir rt postu and '
sltion],sprtsDrwin's work frlm [Jan-Baptistll,amark's'' 1Ospo. most sLlssfultrst
v a t 1 9 8 1 .i 6 8 ) . spirnnd th piod fror
Diosurs 517
:or rsarhin th history of si- Amundsln, R. 2005. Th hgig Rol of tb mbro i ullutior Biolog:
intist whls importan was Roots of uo-duo. amridg: mridg Univrsity Prss.
Burian, R. ' 2004. 1listmllogil Pprs o Duelr\lmet, uolutio, d G-
tis. amridg: amridg Univrsity Prss.
ltratd on his voyag on HMS -l
Kohn, D., d. 985. h Drii Hritge. Printon, Nj: Printon Univrsity
writtn soon aftr h rturnd. Prss.
win's volutionary thought and -he
ospvt' D. 1981 . Deulopmt of Drtuin's hor. mridg: ambridg
l sltion.This phas lf Dar_ Univrsity Prss.
s natural sltion ws th as- Rihards, R. J. 1992..h nig of uolutil: The orphologil ostrutio
d ldologil Rlstrutio of Dri's hor. hiago: Univrsity of
ly with th synthti thory of
'as oftn rgardd as a puzzl. If hiago Prss.
-l840, 2003. The Romti lptil of Lif: Sci d Phiklsopb i te g
lfor why did h dlay of Goethe. higo: Univrsity of hiago Prss.
Ln answr to that qustion and' 2004. ihal Rus,s dsign for |iving, Jourl of tb Histlr of Biolog 7:
l as a man of his ag-a Vito- 2s-38.
Rus, M. |996, r-ld to : h opt of Progrss i uolutior Biolrlg.
amridg' : Harvard Univrsity Prss.
sptivaus h was th first
:romth s-alldlak ox' th 2004. Th romanti onption lf Robrt J. Rihards. Jurl of th
Histor of Billog7: -2' -R..
l in th 1850s in prpaation for
work was, f ours, intrruptd
rtural sltiln,and o th ori-
..Th History of volution- Dinosaurs:Th modl systmfor volution
:ssay
lum.) Th first pratitionrs of Bausof thir immns popularity, dinlsaursar oftn th organisms that
as Darwin's major ahivlnt first om to mind whn on thinks aout animals from th past. This, ou-
Ovat,in ontrast' dpitd him as pld with th fat that thy wr disovrd at th pla (mid-nintnth.
;inof his day, inluding pal- ntury ngland) whr modrn volutionary thory took root, has ld to
nyology. This plaind Dar- thir historially ing usd s xampls in volutionary dbats.In this on-
ltion was in pla, Darwin's tt, dinosaurs hav playd oth positiv and ngativ rols in argumnts
d ws rlativlywak in th arly aout volution. Nvrthlss,dinosaurs rmain an important atalyst for
g th two dads for 1959. dvloping a mor sintifiallylitrat soity, a rquisit for appriating
to argu for transmutation Who and undrstandingvolution.
ntalviW of natur that mrgd Dinosaurs ar uniqu linag of rptils st apart from othr groups y
tuh s his nw thory [natural thir rt postur and distintiv hip and hand strutur.Thy ar also th
an-Baptist]Lamark's'' (ospo. most sussful trrstrial animals to vr inhbit th ath. Thir tnur
spannd th priod from 225 to 65 million yars go. Furthrmor' if ids
' 18 Dilsrtrs
sfiguron pag 520), thir latr Rihard own, who purhasd th spimn for th British usum (in
to thprsntday, whr thy l863), alr dsrid th spimn. Both' prhaps du to thir antievolu-
lS. tionary thinking, fild to attritany volutionary signifianto th atypi-
humbl ginnings. For in- al avian attriuts. Susquntly,Thoms Hnry Huly ntrd th pitur.
ts giants' th arly forms wr Frsh off n xtnsiv study of avin anatomy and systmatis, h rognizd
lsrnivorous ipds wr this spimrrimportan nd usd it s modl organism in his famd l-
ls. ovr tim thy divrsifid tursand puliationsin support of volution (.g.,Huxly 1s6s). To this day
m a j o r l i n a g sg i g a n t i s m o - rhoptery is lratd as on of th most lar-ut ampls of an vo.
hr or mor tons. Som di- lr'rtionrymissing link'
i p d l 'w h r a sh b i v o r o u s So how do dinosaurs figur into th rheoptr stor? Aout th sam
oad sal,dinosaurs appar to tim that Arbeoptry, WaS disovrd, a hikn.sizd thropod dinosaur
.al rprtoirs.Physio1ogially known as ompsogthgus was also disovrd irr Grmany. This tiny di-
ptils.Biologially thy wr nosaur provd for th first tirrrthat not irli dinosar.rrsw lphant-sizd,as
nals, thus possily plaining own had ontndd. Dvlpmntal iologist arl Ggnbaur in 1864 as-
tutly pointd out that th ankl strutur of ompsognthgus was akin to
urs ourd in 1824 whn that of irds. H lrrluddthat it was n niml of doul lationship-
argarnivorous thropod di- both ird and rptil. Amrin paleorrtologistdwin Dinkr op latr
)toussdimntsin ngland. ahdth sam onlusion rgarding th lrg arnivorous tyrannosauroid
s o v r i os f o t h r d i n o s a u i a n Llps (= Drptrlsurus)'In a srisof paprs pulishd from 1866 to 1869
r p t i l i . r na f f i n i t i s ,i t w a s i n i - h pintd out that th longatd nk vrtraand light skull strutur in
d . u p v a r i n t so f l i v i n g s u i a n Lelps wr also avian faturs. Shortly thaftr, Huly, inspird y
not until l842 that this pra- Ggnbaur'sfindings and apparntly unawar of Cop's rsah,notd th
anatomist Sir Rihrd Own rmarkai similritis btwn th hips of eglosrus and tant irds.
l lingin and of thmslvs. Using rhopter as guid, h was al to idntify many attriuts in
arfully grat rptils.'' own, ommon twn irds and dinosars. By th 1880s h had rahd an in-
ut rhet ttriutsof ths ani- sapal onlusion-I.lot only r ids rptils' thy r in fat living
usion of th hip vtra' ar thropod dinosaurs. rbeoptrr is simply a fathrddinosaur.
irds and mammals. F{n,h This viw of avian anstry is now almost univrsally aptd among pa-
and physiologially mor ad- lontologists.Ironially, dspit ing naly a ntury and half old' it has
that living rptilsa dgnr- nly rrrlyn doptd on a road sal. By th turn of th nintnth
1qustiondth plausiility of ntury' palontologists instad favord th ompting thory of stm rp.
;. any sholars liv that tils from whih th mjor lads of ahosaurian rptils (dinosur rla-
,lutionaythoris that wr at tivs)' roodilians, hrivorous ornithishian dinosaurs, ptrosaurs (flying
rptils),saurishian dinosaurs (thropods and long-neked hrivors),and
to figur prminntly in evolu. irds indpndntly ros. It was flt tht th ommon ftures shard y
ston qllarry in Solnhofn, G- irds nd dinosurs stmmd fom mor anint ommon nstor than
r. Tts finding showd tht within thropods thmslvs.
51,two yars aftr harls Dar- In th 1960s, Huly's harg was rnwd whn Amrian palontologist
3w spimn of rbeopter, John ostrom disovrd wll-prsrvdspimnsof Dinonhus, a mm.
tru,ovrall natur of this r- r of th Domaosauria (small to rndium-sizd, agil rnivoros di-
sportd wings, fathrs' and a nosaurs with sikl-lik laws on thi ft). ostrom and othrs idntifid
possssdtth, a ptilian tail, narly 200 dinosaurian attriuts shard twn irds and dromaosaurs.
..missinglink'' (n animal in- inluding nmrous fusd sral vrtr,hollow lg ons, and long for.
)arwinand his followrs ndd arms (s figur on pag 520). Furthrmr, ladisti analyss-a mthodol-
zing|, tlris rvltion was not ogy whry shard-drivd(volutionailyhangd)haratristisar usd
AndrasWagnr (in 1861) and to stlishrlationships-showd tht th gnsisof ids am from within
520 Diosurs
portd glidr, appars to hav had wings. Finally, thr was th disovry of
irlrptor, th smallst known dinosaur-rcheopter-sized and a
asal dromaosaur to oot. larly miniaturization had ourrd for th
volution of irds. Additional findings solidifying th argumnt for dino-
saurian anstry for irds inlud th disovry that gg mirostrutur of
nlnaviandinosaurs is mathd in birds, that thropods roodd thir ggs
I i k i r d s , a n d t h a t x t n s i v a v i n s p i r r o ya i r s a s y s t m s x i s t i n d i -
nosaur skltons. !(/ith ths latst findings, th dat ovr avian anstry
has all but ndd. Dinosaurs did not go tint at th K- oundary (th g-
ologial bordr sparating sdimnts from th sozoi ra, whn th di-
nosaurs ruld, from mor modrn nozoi ra sdimnts)'dstroyd by an
astroid or othr atalysmi hang. Thy in fat rsid fluttring about
outsid today.
Dinosars hav playd an important rol in th dvlopmnt of modrn
volutionary thory y providing on of th most lbratd ampls in sup-
port of volution-rhopterx and th volutionary stags lading to its
r, and th dromaosurlan
gnsis.But do ths animals hav anything to offr for pomoting volution-
:s twn th ird (left) and te
h t p o s u r .s k t d a n i v o r u s ary undrstanding in th futur? I liv th answr is ys. Alng with x-
t' thI main tos and fingrs, an ampls of human and quin (hors) volution, studis of dinosaurs will
lngatd and rtrovrtd puis (hip srv as a valual vhil y whih th puli an om vrsd in the si-
,d fmur (thighon)' longatd ntifi mthod and at th sam tim grasp th workings of volution. Th r-
a prforatatula (a tunnl beopter story is not th only xample that is workd out in intriat
ronsattah),n antrital dtail, although argualy it has sen mor xamination than any fossil vrt-
; o k r )' n d h l l o w o n s ' j u s t t
rat. In fat it is just on of many. Sintists now know how many tims di-
nosaurs rahd lphantin proportions, how thy did it through sltion
for variant dvlopmntal trajtoris, and how thy am dwarfd in is-
.s
land sitations. know how thy divrsifid with ky innovations suh as
rnliklythat irds ould hv d- dntal attris (th volution of hwing using multipl tth) in onjun-
tion with floral hangs. And w hav a good undrstanding of how limati
;t univrsal.Nvrthlss,during hangs' ontinntal drift, and th dmis of nonavian dinosaurs-plausily
ly avian palontologists and or_ at th tim of an astroid impat-ld to volutionry hang that shapd th
t anstrystill might found in world.
l that similaritis btwn dino- Th aformntiond studis of volutionary vnts hav rivd onsid.
lpndntadaptations to similar ral attntion in th worldwid mdia and hn in th publi y.
thir dissnsionwas th asn Through dinosaurs th volutionary mssag is gtting through. As suh, for
:riutsof irds. Ths inludd roadiy promoting voiutionary undrstanding' dinosaur rsearh is ar-
ula) and fathrs,as wll as th gualy on of th most important filds today. owvr, dinosaur rsarh's
ght to th anstors to birds' influn in promoting sintifi undrstanding and its potntial for hlping
l smdunlikly andidats for volution gain roadr aptan ould onsidraly gratr. }yIor of-
)pter. Spurnd y thir on- tn thn not our hildrn's first introdution to sin is through dinosaurs,
on th study of ths faturs. In and thr is no ttr opportunity to tah th sintifi mthod than to
r dinosaus.Rmination of young' urious, opn minds. Thos who ask for data in support of what thy
thy had nt n prviously r- ar told will ttr quippd to judg for thmslvs whthr what thy
linosaursfrom hina. Prsumaly ar told is a rality or not. It is for this rason that dinosaur rsarh is an
rrgulation.on spimn, a ur- important sintifi pursuit, on that an failitat sintifi litray and
522 Dobzhnsh
provid th rquisit akground for undrstanding and iudging th lgiti- that fild and st its arl1
may of astrat onpts suh as volution. Tahing sien at an arly Getis d the rigm
ag should of primary importan for sintists intrstd in promoing a Was that th study of g
mor sintifi litrat soity. Dinosaurs an provid th modl ampls to duil to gntispr s
mak this a rality. th ntirty of volution.
gard h stipulatd y d
B|LIoGRAPHY gn frqunis'' (Doz
op, . D. \867 . An untof xtint rptilsthat aproahird,s.Procedigsof Th main prolm of
th cdem of Nturl Siecesof Phildelphi19: 234_25. ..parado
Gtics, is th
Darwin, . 1859. on the origin of Speles.London:John Murray. to hav variation prsn
nhantdLarning.Zoom Dinosaurs.[.!bsit for hildrn.]http://www
othr hand, th partiul
.nhantdlarning.om/sui ts/dinosaurs/.
ar vry likly maladapt
Ggnaur,. 1864. Untersuhge 7ur uergleichnd Atomider ..volutt
,1., mati fashion,
Wirblthir.o|. rpus und rsus.Lipzig:!7ilhlm nglmann.
Huxly,T. H. 1868.On th nimalswhih ar mostnarlyintrmdiat btwn lssly dar pri of ont
irdsnd rptils.Geologilgzi5:357_365. unfavorabl mutations''
ostrom, J. H' 1969. A nw thropoddinosaurfrom th Lowr rtaousof D o z h n s k y . so w n i n s
]Vlontana.Postill 128: T-17.
variation in natural (nor
I975 . T origin of irds.ul Reuit,c'
of rth d PltrSins3:
55-77 . in Russia. At that tim
Own, R. 1853. on theArheoptero| Von Myr, with a dsriptionof th btls,ut it was his stt
fossil rmainsof a long-taildspisfrom th lithogrphistonof Solnhofn. f|, Drosrlphil, that w
PhilosophiclTrsctionsof th Rol Soietof Lodo |53: 47. drtakn in th 1930s,
Univrsityo{ California N4usumof Palontology.www.ump.rkly.du. working with th Drosa
.Wagnr,
J. A. 1861. Ur in nus,anglihmit Voglfdrnvrshns Rptil aus
Dozhansky,s arly Wor
dm SolnhofnrlithgraphishnShifr.Brishkdmider
'Wissshft (unih) 2z 146-154. -G.'. 1981).
Drosophil at first s
thr smd to no int
gan's itmnt at finalL
Dobzhansky'Thodosius(|900-975)
i n h r i t n o u l d s t t
Thodosius Dozhansky is prhaps most famous for th slogan, ..Nothing in gratrnumrs in his la]
iology maks sns xpt in th light of volution,' (Dozhansk 1973|, ations wr artifats. D/
whih is ndlssly invokd y volutionists in thir attls with rationists. traspifi variation' and
But that is only half th attl that Dozhansky wgd on half of th im- volution. In trying t
portan of volutionary thinking. H had two targts in mind: rationists wild Drosop hil' Dozhl
and thos would- volutionists who naivly suppos that molular iol- who proposd that most
ogy is th ky to lif whn in fat nothing, not vn th prosss of molu- al diffrns only wh
lar biology' an undrstood pt in th light of volution. Ironilly, our mor oftn in la<
Dozhansky himslf was osssdin a rligious or at last a moralisti way as thr is signifianti
with th ..maning'' of vrything, volution inludd (Batty 1'987a, 1'9871 way, natllral population
Rus 2001; van dr eer 2007)' rnst ayr (196) may hav blivd that Dozhansky, originl
volutionary iology providd th ultimat planations of lif, whil mol- W a y t o o s r v t h r w i s
ular iologial explanations ar at st proximat' ut for Dozhansky, th rosop on variations l
ultimat understanding of lif was dpr still than volution (sth disus- that involv diffrns
sion latr in this ntry). gn diffrns as Wll.
To thos familiar with his work, Dozhansky is known not just as a flag f o v o l u t i o n a r y h n g
arr for volutionary thinking, ut in partiular for his ontriutions to Dozhansky at first part
population gntis.H did as muh o mor than anyon ls to stlish to p|in rhis varitit
Dolzhsk 52
andingand iudging tlr lgiti- that fild and st its arly agnda, spiallythrough his influntil ttook
. T a h i n gs i n a t a n a l y Gnetics nd tb rigin of Spis.on of th urrnt thms of Gnetics
l t i s t si n t r s t di n p o m o t i n g a ws that th study of gn frquny hangesis an autonomous fild, not r-
providth modl ampls to duibl to gntispr s (i..,th gntisof individuals). And its domain is
th ntirty of volution. What Dbzhnsky ould not prov in th lattr r.
gard h stipulatd y dfinition: ..volution is [no rrrrthn] a hang in
gn frqunis''(Dozhansky 197, l t; s also susquntditions).
rt approah irds. Proedigs of Th main prolm of population gntis' as Dozhansky l:rl.loratdin
hi 79 234_235. Genetis, is th..parado of viailiry.'' On th on hand, ah spisnds
rn: John urray. to hv vriation prsntin ordr to adapt to nvironmntal hangs.On th
lr hildrn.] http://www otlrr hand' th partiular vriations that might pov adaptiv in th futur
r vry likly ml1aptivat prsnt.As h onludd in his typially dra-
h,1tomi dr
'zig:Wilhlm r.rglrnann. ati fashion, ..vrlutionaryplastiity an puhasd onl at th ruth-
lst nearly intrmdi:rttwn lssly dar pri of ontinuously srifiing som individuals to dath from
/-5b). unfavoralmutations'' (Dozhansk 797, 126-127).
,m th Lowr rta<lusof Dozhansky's own insights into th paradox wr asd on his studis of
variation in naturl (nonlortory) populations, whih gan in th l920s
of ,trth d PItr Sis 3:
in Russia. At tht tinr h ws studying intraspifi variation in ladird
yr,with a dsription of th btls,ut it was his studis of virriation in naturl populations of th fruit
lithographi ston of Solnhfn. f|, Drosophil, tat would prov most influntial. Ths studis Wr un.
et of Ldo 153: 47. drtakn in th 1930s, aftr h had movd to th Unitd Stats nd was
. w.tvw.ump.rkly.du. working with th Drlslpbil gntiist Thomas unt organ (onrning
Voglfdrnvrshns Rptil aus
Dozhnsky's rly w<lrk,s Btty |987a, 1'994 Lewontin 1981; Pror,in
kdmi nn'
|rist-,|l
-..'. o. r98r).
Drosophil at first smd vry diffrnt from ladyugs. Phnotypially,
thr smd to no intraspifi varition at all. on nd only rall Mor-
gan's itmntat finally disovringan osrvalmutnt whos pttrn of
/Jl
inhritan ould studid. Susquntly,variations wr found it-tvr
..Nothing in
rous for th slogn, gratrnumrs in his l:rortory.But it was livdy rrranytha thsvari-
v o l u t i o n ' '( D o z a n s k I 9 7 ) , ations wr artifts. Drosophil in th wild smdto hav littl if ny in.
t n t h i r a t t l sw i t h a t i o n i s t s . traspifi variation, and hn prplingly littl rrrrialfor futur adaptiv
rsky wagd on half of th im- volution. In trying to mak sns of th diffrn twn laoratory and
] W t r g t si n m i n d : r a t i o n i s t s wi|d Drosophil' Dozhansky followd th suggstilnof Srgi htvrikov,
ly supposthat molular iol- who proposd that rost mutatios wr rssivand rvor-rldsult in osrv-
ot vnth prosssof molu- al differnsorly whn dorrld up in th homozygous stat.Tlris would
h light of volution. Ironially, our mor oftn in laoratory stoks than in natllr' bus in th formr
,iousor at last a moralisti way s thr is signifiantly mor inrding than in wild populations. In this
Linludd(Batty 1,987a,1987; way, natural populatins sak up vaiations lik a spong.
,r (1963\ may hav livd that Dozhnsky, oiginally in ollaoratiln with Alfrd Sturtvan' found a
:planationsof lif, whil mol_ Wy to osrv othrwis hiddn variatiorri Dros<lpilay fousing lris rni_
intt,ut for Dozhansky, th Osopon varitilnsin hromosom strutu1.g-.(inyg15i6n''yx1i;11i1;n5-
ill than volution (sth disus- tlrat involv diffrns in gn arrangmnt and rroftn assoiatd with
gn diffrnsas wll. In oth rspts'ths verietions onstitut rntrial
Lnskyis known not just as a flag for volutionay hang. But how ws th osrvd vriation minturind?
..shifting
t r t i u l a rf o h i s o n t r i t i o n st o Dbzhirnsky at first partly adoptd Srvall!7right's balan'' thory
)r than anyon ls to stalish to plain this virriation. H rirsond that tlr variations Wr adaptivly
524 Dobzbnsk
insignifiant and that thir frqunis wr drifting randomly. This might As muh as h laim
lad to a rdution in variation within ah population of a spis' ut dply tzz\ed and un
would lad to an inrasin variation twnpopulations and hn within prodd. H had four
th spis. t h p r s r v a t i o no f v i
Dozhansky was quit surprisd, thn, to find vidn that th invrsion that ronild th go
diffrns wr adaptivly signifiant aftr all, lading him to plor a rang gratr fitnss of htro
of variation-maintaining forms of natural sletion. H finally sttld on th tinu to orn in ah
ida that htrozygots ar adaptivly suprior to homozygots us hav- undrstood this situatlo
ing two gns for a ttalt, instad of two opis of th sam gn, allows mor fr will, whih givs us
adaptaility at th individual lvl. Sltionfor adaptaility at th individual Prhaps. But h also x1
lvl in turn prsrvsvolutionary adaptaility at th spis lvl. ntur'' tht so many ir
..alan''
Dozhansky's aount am to known as th thory of vo- of th group (Dozhan
lution. His main dtrator was rmann Mullr' whos viw of volution 1_1'22;van dr r
..lassial'' (old-fashiond)
was dud y Dozhansky th thory (on th Dozhansky drw p
lassial/alanontrovrsy, s Batty 1987 [and othr ssayson that is- sours in his invstigat
su] and Lwontin 1974). Aording to Mullr, thr was no parado of via- was a major rason for t
ility. Th vast majority of mutations ar dtrimntal for th individuals in txt. Dozhansky's und
whih thy our, and lso for th spis in whih thy aumulat. Thr intraspifi variation a
was, in ullr's opinion, only on spis-humans-in whih mutations a- dr Mr 2007), lthug
umulat to any grat tnt. Through th amnitis of ivilization, w hu- and many of th sam
mans hav managd to sap natural sltion,to our ultimat dtrimnt. also n raisd in th l
Only through a onsious ugnis program an W now sussfully rvrs in onvrsations with D.
th rising tid of dltriousmutations. Dozhansky, for his part, ritiizd
..th B|LIoGRAPHY
ugnial Jrmiahs'' who ovrlook th importan of volutionary
plastiity and who sm unawar that natural populations of many spis Adams, M. B., d. 1994..
possssonsidralrsrvsof varition (Dozhansk 1'97, |26). PrintonUnivrsityI
Btty,l. 1987a. Dozha
Dobzhansky and ullr wr also at th ntr of ontrovrsis from th
biology.In L. Kriigr
1950s through th ear| |970s onrning th impat of radiation-indud amridg,A: Har
mutation from atomi Wapons tsts. (Both mn rivd onsidral fund- |987..!ighint
ing from th Atomi nrgy ommission during this priod.) Adding yt Jourl of th istor
mor mutations to th human spis was straightforwardly ngativ from 1994.Dozhansk
signifian of gn
ullr's point of viw, but lss so from Dozhansky's viwpoint. Intrst.
ThodosiusDobzh
ingly' Dozhansky was strongly opposd to th tsting and us of atomi
Dozhansky,T. 1'937.C
Wapons' whil th on pro-Sovit ullr livdthat, up to a point, suh UnivrsityPrss.
Wapons had an important rol tl play in th <rntainmntof ommunism 1.962,Mnhidl
( B a t t y1 9 8 7 ) . avn' T: Yl Unrv
Dozhansky also usd his studis and prsptivson variation to suppot l 9 7 . N o t h i n i r r
mri Biolog T
liral dmorati idals. As h and Lsli Dunn argud in thir popular
Dunn' L. ., and T. Doz
Hredit, R, d Societ (Dunn and Dozhnsk 1'946,45), ..thsolut Pnguin.
uniqunss of vry human individual . . . translatd into mtaphysial and Lwontin'R' , 1974,h
politial trms is fundamntl for this nd dmoray.'' In his widly rad Harvard UnivrsityPl
kid uoluig, Dozhansky argud that, far from gnti divrsity un. I9 8 l . l n r r d u t
drmining qulity' it atually srvs as an llnt rational: ..quality of Lwontint al.,ds.'
N w Y o r k : o l u m h i
opportunity tnds to mak th oupational diffrntiation omport with th
Lwontin,R. . t al.,ds
gnral polymorphism of th population, nd would maninglss if all I-XLIII. Nw York: (
popl wr gntiallyidntia|" (|962, 244). N4ayr,. 1963.ausnd
Dobzhsk 525
drifting randomly. T'his might As muh as h laimd to undrstand volution, Dozhansky rmaind
r population of a spis, ut dply puzz|ed and unomfortal aout th mans y whih evolution
l populationsand hn within prodd.H had found a Way to ronil th sltion of th fittst with
th prsrvation of vaiation for futur adaptiv volution, ut not in way
i n d v i d n r h a r t h i n v s i o n that ronild th good of th spiswith th wlfar of its mms.Th
, lading him to xpl<lra rang gratr fitnss f htrozygotsmans that lssr-fithomozygots will on-
:tion' H finally sttld on th tinu to orn in ah gnration.It hs n suggstdthat Dozhansky
r to homozygots aus hav- undrstood this situation rligiously, as a nssaryvil-along th lins of
; of th sam gn' allows mor fr will, whih givs us moral frdom' ut at th ost of many sad hois.
r adaptailityat th individual Prhaps. But h also pliitly rgardd it as an ..unplasingimprftion of
y at th spislvl. natur'' that so many individuals should suffr suh ..misry'' for th good
..lan''thory
rs th of vo- of th group (Dozhansk197,1271Dozansk 1962, 35; Rus 2001,
ullr' whos viw of volution |00_\221van dr 2007).
(old-fashiond)thory (on th Dozhansky drw produtivly on his Russian hritag and Russian
7 [and othr ssays on that is. sours in his irrvstigationsand planations of intraspifi variation. This
:r,thrWas no paradox of via- was a major rason for th originality of Gtics in an Anglo-Amrian on-
jii'mf;..f, . r.tl * ;nci;.'ict r]s-;' txt. Dobzhnsky's undrstanding of th rligious/moralisti signifian of
l whih thy aumulat. Thr intraspifi variation also rfltd his Russian hritag (s spilly van
u m n s - i n w h i h m u t t i o n sa - dr Mr 2007), lthough this influn has n mor diffiult to ronstrut,
rmnitisof iviliztion, w hu- and many of th sanr onrns-suh as th vils of natural sltiln-had
: i o n ,t o o u r u l t i m a t d t i m n t . also rrraisd in th Anglo_Amrian ontt (..g., y th athist organ,
an W now sussfullyrvrs in onvrsationswith Dozhansky; B21tt|994,207 and not 3).
lzhansky,for his part, ritiizd
|BLIoGRAPHY
h importan of volutionary
. l p o p u | a t i o n so f m a n y s p i s Adams,M. B., d. 1994. Th ullutioof ThodosisDobzhsk.Printn,NJ:
o b z h a n s k yt 9 7 ' 1 2 6 ) . PrintonUnivsityPrss.
Batty,J. 1987a.Dozhanskyand drift:Fats,valusand hanin volution;rry
ntrof ontrovrsis from th
iology.In L. Kriigr t al., ds.,The ProbilistiRuolutk-l,olr.2.
himpat of radiation-indud ambridg,A: avrrdUnivrsityPrss'
mn ril,donsidalfund- 1987.!7ighingth isks:Stlmt in th lssil/alan
ontrovsy'
Juring this riod.) Adding yt lourl of th istor of Biolog 2:289_|9.
;traightforwardly ngtiv from \994. Dozhanskyand th iologvof dmoray:Th moral and politial
signifianof gntivariation.In . B. Adms, ed.,Tbe uolutio of
lzhansky'sviwpoint. Intrst-
TheodosiusDobzhsk'Printon,NJ: PrintonUnivrsityPrss.
) th tsting and us of atomi Dozhansky,. 1937, Geticsd the origi of Sples.Nw York: olumia
rclivdthat, up to a point, suh UnivrsityPrss.
: h o n r i n m n to f o m m u n i s m |962. nkitl uoluig:Th uolution lf the um Speles.Nw
Havn' T: Yl UrrivrsityPrss.
;ptivson variation to support 1973.Nothingin iologymakssnsptill th lightof vllution.
triB iollgTrhr5 : 1 5-129.
Dunn argr.rdin thir ppular
Dunn,L. ., arrdT. Dozhnsk'1946.Hrdit,R,d Soiet1'. Nw Yok:
: h a n s k y\ 9 4 6 , 4 5 | , . . t h s o l u t Pnguin.
anslatdinto mtphysial and Lwontin,R. . 1974. h Geti Bsisof uolutirlrhge.Camridg,MA:
dmoray.'' In his widly rad Harvard UnivrsityPrss.
, far from gnti divrsit un- 1981.Introdution: Th sintifiwork of ThdosiusDozhansky.In R. .
Lwontint al., ds.,Dobzhsk'sGetisof Nturl PopultiosI-XLIII.
llntrational:,.quality of
Nw York: ColuriaUnivrsityPrss.
diffrntiationomport with th Lwontin,R. C. t al.' ds. "|981. Dobzhnsk'stisof Nturl Popltions
rnd would mninglss if all LXLLI]. Nw York: olrrmiaUnivsityPss.
41 ayr, . l963. irusand ittin iologv.Si134:1501_1506.
526 Drosophila
.v.
Provin, B. 1981.origins of th Gntisof Natural Populationssris.ln R. .
Lwontin t al., ds.,Dobzhsk'sGtisof Nturl PopltionsI_XLI.
Nw York: olumbia UnivrsityPrss.
Rus, . 2,|.ster of stris:Is uolutio Slilostrutio?
ambridg,A: Harvard UnivrsityPrss.
van dr Mr,J. 2007.ThodosiusDozhar-rsky: ..Nothingin volutionmakssns
xptin th light of rligion.''In N. Rupk, ed,,mint Lius i Tt,utith-
tur Sic d Religion. Frankfurt: Ptr Lang. -1.8.
DrosophiI
Drosophil is a gnus of flis in th family Drosophilida that ar widly
usd for volutionary rsarh. Th origin of this rsarh lis in Thomas
Hunt organ's Noel Priz-winning work on inhritanc in Drosopbil
mlogstr (s figur) starting around 1908. organ's intlltual
dsndants hav pandd th sop of work on D. melnogstr from
gntis into virtually vry aspt of iology, turning it into on of th
most usful modl organisms. volutionary rsarh has n a roust
part of this tradition from th bginning. urntly' th omination of
th as and spd with whih many spis an rard in th laora-
tory and th tnsiv akground knowldg of gntisand dvlopmnt
mak thm usful in addrssing a wid vrity of qustions in volutionary
gntis.
Drosophil adults ar usually only a fw millimtrs long' with prominnt
rd to rown ys, and ompl trnal mophology that has rmaind
fairly onsrvativ ovr volutionary tim. Drosophil sow holomtaolous
dvlopmnt, undrgoing a omplt transformation from th fding larva
to the adult form during a pupal stage. Thy ar ommonly alld fruit flis,
although most spis fd on miroorganisms that inhait rotting plant ma.
trial. Th gnus Drosopbil urrently inluds twn 1,150 and 1,500
spis, dpnding on whih groups ar inludd in th gnus, and has a
worldwid distriution. Phylognti rsarh has larly shown that th
genus is paraphylti, maning that othr gnra ar mor losly rlatd to
som of t'e Drosophil than thy ar to othr groups. In th road sns,
thn, Drosophil sou|d inlud many other gnra' inluding th spis-
ri Sptomz (approximtly 260 spis), Hirtodrosopbil (approi- Drosophil mlogs
matly 150 spis), and codrosophil (approimatly 120 spis). Th way in whih it wl
N{olular vidn sttggststhat th gnus is at last 50 million yars old, \994.
a onlusion supportd y drosophilid spimnsin ambr that ar around
this ag.
Around 1910, organ disovrd a singl D. mellgsler with whit
ys' th inheritan of whih turnd out to s linkd. This simpl obsr- undrgraduats Alfr
vation inspird Morgan and his laoratory studntsto isolate othf mutants, N4ullr, who all playr
with whih thy wr al to onfirm th hypothsis that gns rsid on Morgan, Bridgs, an
hromosoms. In thos yars, Morgan's tiny (16 ft 24 feet)..fly room'' whn thy all movd
at olumia Univrsity had ight dsks; thr wr oupid y olumbia whr thy rmaind
Drosophila 527
aturalPopulations sris.In R. .
of NturI Popultios I-Llll.
SoiI ostrutio?
.Nothing in
vlution mks sns
d.,liet Lius i Ttuentietb-
:r Lang. -1.8.
r i l l i m t r lso n g , w i t h p r o m i n n t
morphology that has rmaind
r osophil show holomtaolus
rmation from th fding larva
.ar ommonly alld fruit flis,
ns that inhabit rotting plant ma-
u d s t w n1 , 1 5 0 a n d 1 , 5 0 0
ludd in th gnus, and has
h has larly shown that th
|nraar mor losly rlatd to
'hr groups. In th broad sns,
)r gnra' inluding the spis-
ies\, Hirtodrosopbil (approxi- Drosophil mlogster'th fruit fly, th workhors of volutionarygenetis.
r (approximatel 720 spis). Th way in whih it was tailord for suh rsarhis disussdin Kohler
is at last 50 million yars old, 1994.
imns in amr that ar around
sturtvant produd th first gnti map asd on th raliztion that th lvls of natural variz
frquny of rossovr vnts ould b usd to dtmin th ordr and loa- whthr gnti varr
tion of gns. H also workd on a wid varity of topis with volutionary maintaind by natur:
impliations, inluding hromosomal invrsions, th gntis of spis dif- ontrasting Dzhan
frns, th gntis of dvlopmnt, and th phylogny of th gnus. with ullr's mpha
Bridgs was among th first to xploit th disovry that th salivry glands was a major fator in
of Drosophil ontaind polytn hromosoms onsisting of thousands of of Drosopbila to stu
DNA strands rsulting from rpatd rounds of rpliation without ll divi- quning of Dros<lpb
sion. Th rsult is hromosoms visil at low magnifiation with hartr- of volutionary patt
isti anding pattrns that allow homologous rgions to idntifid aross Th as with whi<
individuals and spis. Bridgs's dtaild dsriptions of th anding pat- in th study of volut
trns allowd linkag and physial maps to alignd. ltion xprimnts <
ullr is bst known for his Nol Priz_winning work on mutation. B- Knnth Mathr and
ginning in th mid-1920s, ullr was th first to ploit th fat that hro- Susquntly' sus
mosomal invrsions-hromosoms in whih a pi appars in rvrs of haratristis, no
th normal ordr-supprss romination whn in a htrozygos ondi. gg siz, dvlopmn
tion with an uninvrtd hromosom. Mullr found that h ould us an in. totais' gotais' m.
vrtd hromosom markd with a dominant phnotypi mark, trmd a (rats of rominat
.!7ith
alanr, to aptur and study th proprtis of hromosoms with th nor- th xption
rnal gn ordr. Initially h usd this thniqu to dtt hromosoms arry- vry trait studid. S
ing lthal mutations' whih an nvr produ a homozygous adult. Mullr rats an nvironm
masured th lthal frquny aftr induing mutation with -rays. Mullr's ara of ativ rsar
onpt of gnti load-mutational aggag that ullr saw as harmful- van in th study of
am from ths arly primnts. Balanr hromosoms wr sus- ing and mat hoi
quntly dvlopd for ah hromosom in D. mlogstr, and alanrs Th study of sp
rmain on of th ky advantags of doing gntis in th spis. Mullr, and Dozhl
Shortly aftr Morgan movd to al Th, h hird Russian sintist Thodo. sarh. Th gnus ir
sius Dozhansky as part of his tam. Dozhansky quikly ros to prominn numr of rossal
as a synthsizer writing Geetics d the rigi of Species (197), on f barrirs an idnt
th founding ooks of th volutionary synthsis,in whih h assrtdth pri dnts, and his stud
may of a population gnti viw of volution, basd initially on rmarkably rough stimats of
littl mpirial vidn.At aout th sam tim, inflund y Sturtvant and lopatri populations
his Russian mntor Srgi htvikov, Dozhansky took up th study of wild hav now n idnt
populations of Drosophil pseudoobscur. Populations of this ommon .s7hil
th study o
spis of th mountains of wstrn North Amria ar polymorphi for hr- melogster am
mosomal invrsions. Dozhansky usd this variation as th asis for a long s- asd on th dtail
ris of xprimnts doumnting th ffts of th thr major evolutionary Sturtvant's studnts
fors: sltion,gnti drift' and gn flw. In this, h was grtly aidd y an ffts arly in dv
.lfright
xtnsiv ollaoration with Swall during th lat 1940s. shus. Ths ffort
Modrn volutionary work on Drosophil urrntly has svrl major m- ways that ar ons
phass that grw out of this arlir work. Th study of variation t th mo- .Wishaus
and shar
lular lvl was pionrd y Rihard Lwontin, on of Dozhansky's akground inform
studnts, who, starting tn 1965, xploitd th disovry of protin l- lution of dvlopmt
trophorsis by Jak Huy and othrs to dmonstrttht a larg portion of idntify th spifi 1
gnti loi in Drosopbil spis wr polymorphi. This disovry of high ral and xprimnta
Drosophila 529
|asdon th ralization that th lvlsf natural variation in most spis ld to an arimonious dbat ovr
o dtrmin th ordr and loa- whthr gnti variation is gnrally sltivly nutral, dltrious, or
i yo f t o p i s w i t h v o I u t i o n a r y mintaind y natural sltion. Lwontin hlpd to cata|ze this dat by
ns' th gntis of spis dif- ontrasting Dozhansky's viw that muh gnti vaiation was adaptiv
th phylogny of th gnus. with Mullr's mphsis on th osts of mutation and variation. This dat
iovry that th salivary glands was a major ftor in th dvlopmnt of population gnti thory. Th us
ms onsistingof thousands of of Drosopbil to study variation arris ovr to th DNA lvl, whr th s-
of rpliation without ll divi- quning of Drosophila gnoms allows unprdntd prision in th study
v magnifiation with haratr- of volutionarypattrns.
; rgions to idntifid aross Th as with whih Drosophil an rard has ld to its inrasing us
sriptionsof th banding pat- in th study of volution in th laoratory. Som of th first sophistiatd s-
alignd. ltion xprimnts on ontinuous traits wr arrid out in Drosophil
winning work on mutation. B- KnnthMathr and Fors \/.Rortson in th lat 1940s and arly 1950s.
st to ploit th fat that hro- Susquntly,sussful artifiial sltion primnts took pla on many
h a pi appars in revrs of haratristis'nompassing mophology and physiology (.g.' fundity,
whn in a htrozygous ondi- gg siz' dvlopmntalrat, lif span, rsistanto thanol)' havior (pho-
: f o u n d t h h o u l d u s n i n - totais, gotaxis, mat prfrn), and aspts of th gnti systm itslf
rt phnotypi markr, trmd a (ratsof romination, hang in varian rathr than man valu of traits).
i of hromosomswith th nor- lfith th ption of sx ratio and asymmtry, hrital variation xists for
l to dtthromosoms arry- vry trait studid. Studis of primntal volution, whr th invstigator
a homozygous adult. N4ullr ratsan nvironmnt in whih natural sltion an take pla, is also an
mutationwith -rays. Mullr's area of ativ rsarh using Drosophil. This approah has prtiular rl-
-'that Mullr saw as harmful- van in th study of lif historis and th ultimat origin of prosss lik ag-
er hromosoms w sus- ing and mat hoi.
D. melogstr, and alanrs Th study of spis diffrns, pionrd tn Drlsophila y Sturtvant,
)ntisin th spis. ullr, and Dozhansky, is anothr vry ativ ara of volutionary r-
: hird Russian sintist Thodo- sarh. Th gnus is wll suitd to th study of spiation aus of th
nsky quikly ros to pominn numr of rossal spis and th rlativ as with whih rprodutiv
rigi of Spcies (1937), on of arrirs an identifid and quantifid. Jrry oyn, on of Lwontin stu.
:sis,in whih h assrtdth pri- dnts, and his studnt H. Alln orr usd this aumulatd data to mak
'n, asdinitially on rmarkably rough stimats of th tim rquird for spition in sympatri and al-
n, influndy Sturtvant and lopatri populations. Th gns rsponsil for aspts of spis barrirs
ransky took up th study of wild hv now n idntifid in a numr of Drosophila spis pairs.
Populations of this ommon \/hil th study of Drosophil dvlopmnt dats ak to th 1920s, D.
rri ar polymorphi for hro- melogstr m a prmir modl systm for th study of dvlopmnt
riation as th asis for a long s- asd on th dtaild study of th ithorax ompl y d Lwis, on of
of th thr major volutionary Sturtvant's studnts' and th systmati mutation srn for mutations with
n this,h was gratly aidd by an .$i-
ffts arly in dvlopmnt y hristian Nusslin-Volhard and ri
rringth lat 1940s. shaus. Ths fforts ld to many of th fundamntal dvlopmntal path-
u r n t l yh s s v a lm a j o m - Ways that ar onsrvd widly among animls. Lwis, Nusslin-Volhard,
r study of variation t th mo- and \Wishaus shard a Nol Prize for this work tn 1'995. This tnsiv
3wontin' on of Dozhansky's akground information on dvlopmnt has cata|zed th study of th vo-
th disovry of protin l- lution of dvlopmnt in Drosopbil. Rsarhrs hav now n al to
lonstratthat a larg portion of idntify th spifi gnSrsponsil for many volutionary hangs in natu-
nrphi. This disovry of high rl and primntaIDrosophil populations.
s0 Drosophila
|LloGRAPY
Dozhansky'T. 1937. Geneticsd the rigin of Speles.Nw Yrk: Columia
UnivrsityPrss.
Kohlr' k. 7994. Lords of th F/y. hiago:Univrsityof hiago Prss.
Lwontin,R. . 1'974.The GeticBsis of uolutionrhge.Nw York:
E
olumi UnivrsityPrss.
Powll' I. R. |997. Progrssd Prospetsi uolutiorBiolog: e Drosophila
odl. Nw York: oxford UnivrsityPrss.
Ruin, G. ., and . B. Lwis. 2000. A rif historyo| Drosophil'sontriutions
to gnomrsarh. Si287:2216-2218.
Sturtvant,A. H. 1965. istor of Gtics.Nw York: arpr and Row.
-D.H. d B.H. E,cologicI Geetlcs
ologicl G entis, t|tr<
s t u d y i n g n a t u ls l t i o
..dmund B
btial ntry
studnts, and olltdth
tis, is dsndd from tI
don that natural sltl
...W.F. R..Wldon'
ntry
gntis ovr work lik \
gnti asis suh as th
mot|7P rt i d omiul .
logial gntiists to stu
sponse to sltion as l
hangs in trait valus r
ths disovriswr s
trm ologil glis
dmonstral gnti fo
trait variation in natur l
hr typs of invsr
thr ar dmonstrations
in th moth Biston betul
vstigations of gograph
snails, that doumnt ho
anothr hang th sl
valus to favord in d
adaptiv xplanatins fo
dmonstrations of adapt
unovrd th signatu
s n a i l s h l lp a t r r n sT. h i r
an alan on noth
tion. Ths studis rpr
Innumral thortial
tion gw from th initi
sgrgatdspiting
cologil Getis l
s t u d i s l i m i n da n y d
Spis.Nw York: lumia
tv of Drosophlla's orrtiutions
) . t,
52 ign
ould studid in ral tim. Th studis of gographi variation ondutd individual frquny.
by Ford and his ollagusomplmntd th work of othrs' suh as Gt .
of th quasi-spis,
Tursson and Jns lusn with plants (s th alphatial ntry ..Jns and dirtd voluti
lausn'' in this volum), F. B. Sumnr with mi, and Srgi htvrikov slf_rpliation.
and Thodosius Dozhansky with flis (s th alphatial ntris ..Srgi In addition to om
Srgvih htvrikov'' and ..Thodosius Dozhansky'' in this volum)' opration lading to t
who wr making similar disovris, oftn with haratrs lss amnal to svral quasi-spisr
gnti analysis. Th waning of th ook's influn gan with th disovry adjant unit. This
of iohmial vriation nd ontinud through th mrgn of a sophisti. in th snario and al
atd approah to studying ontinuously varying traits that ar mor rpr- fat that RNA squ
sntativ of th stuff of volution than disrt polymorphisms in visil lngth (about 100 nu
haratrs. Nonthlss, th qustions aout sltion and volution pr. would aumulat,d
sntd in th book' and th onvition advand that ologial and gnti ordf to od for prot
studisar nssaryto undrstandadaptiv volution, rmain drok influ- mad it possibl to 1
ns on modrn volutionary iology. spis without lngtl
funtional protins. I
|LIoGRAPHY
pndd on thir nl
Ford, . B. |975. ologilGnetis,4thd. London:hpmanand Hall. -J,' form in th rih pri
partmnts ould thn
ign,s thory wal
ign, anfrd (. t927)
hmistry podud l
anfrd Bign is a Grman iophysiist and formr dirtor of th a dant supply of amin,
Plank Institut for Biophysial hmistry in Gttingn. In 1967, toget.er srv as nrgy-rih l
with Ronald Gorg.!ryford Norrish and Gorg Portr' h won th Nol sis of short gnti t
Priz in hmistry for th study of xtrmly fast hmial rations. Bgin- slf-organization. Th
ning in th 1960s, ign's rsarhintrststurnd to iohmial qustions, and mpirial assum
and during th 1970s and 1980s h formulatd an laorat thory on th wr svrly ritiiz
mrgn of lif on arth foundd on mathmatial alulations and pri- to idas on th volu
mnts in viral populations. tion of th onpt
ign's fundamntal thortial and philosphial postulat was tht th prinipl of natu
natural sltion Was th organizing prinipl rsponsil not only for vo. ut on any physial r
lution ut also for th mrgn of lif. This laim was basd on nw ad- standing th rol of
vanmnts in th fild of molular iology. Starting in th 1960s, it lawful pross.
am possil to dsri volution asd on rpliation, mrrtation, and
|LIoGRAPH
omptition not only of organisms ut also in mixturs of slf-rpliating
moluls and nzyms atalyzing rpliation. ign's rsarh along ths ign, M. 1992.Stpst
lins gratly advand th work f iohmist Sol Spiglman and othrs in ign, ., lW. Gardin
of gntiinforma
th 1970s that dmonstrtd th volution in th tst tu of viral RNA s-
Fry, I. 2000. h mr
quns. Basd on ths xprimnts' ign dvlopd his modl for th Nw Brunswik,N
origin of lif.
ign formulatd a ky nw onpt in origin-of-lif study and virology
..quasi-spis.''
alld th Following th omptition for uilding loks in
a population of RNA squnsngagdin slf-pliation?a uniqu pattrn
of th whol population mrgd asd on th distriution of th various
mlrtants. This rsulting population, th quasi-spis,rfltdth onsn-
sus squn rprsnting an avrag of all th mutants aording to thir
ign .'3J
og'faphivariation ondutd individual frqunv' ign disovrd that, asd on th spifi dynami
) work of othrs, sh as Gt of th quasi-spis,th mharrisrnlf sltion guarntd r-rlnrandom
l th alphatial ntry ..Jns and dirtd volution toward a population apl of aurat nd fast
mi, and Srgi htvrikov slf-rpliation.
..Srgi In addition to omptition, ign also postulatd stags of molular o-
h lphatialntris
)ozhansky'' in this volum), opationlading to th mrgnof a ..hypryl,''an nsmlin whih
'ith hrtrslss amnbl to svrlquasi.spisunits partiiptd, ah naling th rliation of th
ngnwith th disovry adjnt unit. This oopration was dmd rr-rialto ovro1 a cath-22
gh th mrgnof a sophisti. in th snario and allow furthr volution. Th diffiulty stmmd from th
ging trits that ar mor rr- ft that RNA squnsin th quasi-spishad t limitd to a rtain
:rtpolymorphisms in visil lngth (about 100 Irulotids).Aov this thrshold, too many mutaions
t sltionand volution pr- would aumulat,dstroying th squns'idntitis.At th sam tim, in
d that ologial and gnti ordr to od for potins,muh longr hains wr rquird.Tlr hypyl
volution,rmain drok influ. md it possil to pull togthr th information ontnts of ah quasi-
siswithout lngthningindividul hins' thus naling th synthsisof
funtional protins' Howvr' th oopration of svral quasi-spisd-
pndd on thir nlosur within a ompartmnt, postulatd y ign to
ion:hprnand all. -l.T. fom in th rih primodil ..soLlp.''optiion mong th divrs om-
partmntsould thn prmit furthr volution.
ign's thory was asd on th optimisti ssumption that prioti
hmistry produd in th primordial soup a far-from-quilirium, aun-
l a fomr dirtor of th Ma dant supply of anrino aids, nulotids, and thi polymrs. Ths ould
n Gttirrgn.|n 1967, togthr srv s ngy-rih uilding loks and primitiv talysts for th syrrth-
;orgPortr, h won th Nol sis of short gnti squns irnd for th ruil prosss of iologial
fast hmial rations. Bgin- slf-organization. This optimism s wll as svral of ign's rhortial
-rrndto iohmialqustions' and mpirial assumptions rlating to th quasi-spisand th hypryl
Ltdan laoratthory on th wr svrlyritiizd y othr rsrhrs.Nonthlss' his ontriution
matilalulationsand pri- to idas on th volution of gneti information and to th latr formula-
tion of th onpt of th RNA world is indisputirl. So is his lim that
losophial postulat was that th prinipl of natural sltion Oprats not only on iologial systms
[ rsponsilnot only for vo- ut on any physial systms fulfilling rtain onditions and that, notwith-
is laim was asd on nw ad- standing th rol of random vnts' th mrgn of lif was a naturally
ogy. Starting in th 1960s, it lawful pross.
l on pliation' mutation, and
BIBLIOGRAPHY
in miturs of slf-rpliating
'n. ign'srsrh along ths ign,. 1992.StpstlrdsLif. ford: Ofrd UrrivrsityPrss.
ign,., W. Gardinr,P. Shustr, rrndR. Winklr-Oswatitsh. 1981.h lrigin
st Sol Spiglmanand othrs in
of gntiinformtion.Sitifimri244, no.4:78_|18.
n th tst tu of vial RNA s- Fry, I' 2000. Th mrgof Lif t rth: lListorild Scintifit',arui.
n dvlopd his modl for th Nw Brunswik,NJ: RutgrsUnivrsityPrss. -t.t.
all tims. Mmrans, and oftn additional ovrings suh as 1lwalls and
utils, dfin th insid and th outsid ordrs of all organisms, all of
whih ar mad of ithr prokaryoti or ukaryoti lls. In hmil terms,
this implis that th insid of any organism is in a stat of Watr, salt' sugar,
d that all ukaryots (organisms protin, and lipid onntration vry diffrnt from its outsid. Th nviron-
nosoms),inluding th familiar mnt yond th outr mmran is vr-prsntand oftn hanging, but it is
symioti mrgs of atria. th ubiquitous ativ mmran (whthr nakd or supplmntd y skin,
grnphotosynthti organlls ) waxy surfa, shath, or othr additional outr layr) that stalishsand
Laalld yanoatria.And - maintinsidntity of living ings.
lon anstor' this vidn on-
..powr
)' too' th oygn-using
ALL L|F FRo H ARLIs LLs
. tradvia gntis,physiology'
r i a .F o r t h l s t i l I i o n y a r s All lif forms on arth mak protins insid thir lls, inluding mmran
4' on has th sam kind of gn- protins.Protins, whih ar long hains of smallr units alld amino aids,
,gy firms. This aunrulation of ar synthsizd on intrallular tiny sphrial struturs alld riosoms. Ri.
; from on kind of batriumto bosoms look lik tiny dots vn at high magnifiation as sn with an l.
-rinttims, th way of th a- tron mirosop. mbran iohmistry and protin synthsis are univrsal
lns, and prmannt allians as fatursof lif that attst to th fat that ll lif volvd from ommon llu-
l s s o i a t i o n st o
s | t d a t r i lar anstors.
of litral takovrs, hav also Batrial gns,thos of humans as wll as all othr animals, ar omposd
liffrnt kinds of bodis, rathr of DNA; th hmial omposition of gns, lik many othr dtails of at-
ia volvdto om largr lls. rial hmistry, ar lik thos of th largr organisms. Most ateria hav at
'll nimals and plants, indd all last 3,000 diffrnt gns that dtrmin thousands of diffrnt nzym pro.
symiotimrgr. tins. Batrial lls grow and rprodu y rprodutiv division, whih is
known as fission. Th prokaryoti ll division pross is far lss omplx in
apparanthan th ..hromosomdan'' alignmnt and sparation pross
in ukaryots. Known as mitosis' th ll division pross in nlatd organ.
prsntin th fossil rord long isms always involvs movmnt insid lls on littl traks alld miro-
plants' volvd photosynthsis tubuls (sfigur on pag 536). Th insid of a ukaryoti ll is inssantly
'ovidfood to th rst of th io_ moving, as sn through a mirosop whn th ll is aliv. vn th largst
struturknown as prokaryoti. prokaryoti lls divid y fission; thy lak th mirotuls and th
o not hav an nlosd nulus mirotuul-asdmovmntstypial of nulatdlls. Som laim that th
a alld nuloid in whih th disontinuity twn atria and all othr forms of lif is th largst volu.
'iols, spindls, or mirotubuls tionary gap in th living wor]d.
ts), and prkaryoti lls lak Although larly w (i..,largr forms of lif omposd of nulatd lls)
h i h o r g a n i m o l u l s r o i _ volvd, somhow, from trial prdssors, th diffrns twn a-
shar many ut not all traits in tria and ukaryots ar signifiant. Th major diffrns twn atria
main groups of mor familiar, and th mmrs of th animal' plant, and fungal kingdoms is that all largr
Lgi(a11yasts,molds, and mush- forms of lif ar omposd of lls that haror doul mmran_oundd
ls that ar oundd y dynami nuli. For that rason thy ar ukaryots (u=tru, karyon=krnl, sd;
th tinistatriato th largst Grk). All atria hav DNA, ut thy lk oth th nular mmran
nrgynd aron mtaolism to and th tru hromosoms whr th DNA is pakagd with spial
n s .T h m t l i m a i n t n n protins.
rih, lipid-rih mmranS x- Batria, as a group' are far mor mtaolially divrs than plants' ani-
:fs as raw matrials. N4mrans mals, and fungi. Whras all th lattr rath oygn' som atria an
of ontinuity through spa and rath sulfat' aron dioid, nitrat' or vn arsnat. Batria, dspit
mranssurrounding all lls at thir small siz, mor anstrl lifstyl, and rputation as parasits and
536 dosmbioti rigin of ukarotes
How might our atrial anstors hav volvd to form th rlist nul-
atd lls? Th volutionary pross always involvs th apaity for rampant
population growth far yond th nvironmnt to sustain it (i.., ioti po-
..hks', growth
tntial),th gnrationof inhritd variation, and th on of
the population (inluding its variants) y th failur of th ioti potential to
rahd. The inhibition of maximal population growth y limination of
many' for whatvr rason' is alld natural sltion. Naturl sltion
maintains halthy rprodutiv populations f many typs of organisms' ut
it dos not produ thm in th first pl.
.!7l1,
thn, how is inhrited variation gnratd? Although random muta-
tion larly xists and has n prodigiously doumntd, inhritd variaion
is gnratd in many mor diffrnt and important ways than via random
mutation. Indd, random mutation almost rtainly did not produ nul-
atd lls from atia.Rathr, th maior mod of volutionry variarin in
atria that ld to th first nulatd lls is likly to hav n via a pross
alld symbiognesis. This pross is allid to symiosis' whih is th olog-
ial rlationship twn ings of diffrnt typs (diffrntly nanrd organ- -''!*\.
D:rrwinian imprftiorrand oddiry from wlrih pth f history rrry . Sry' D. G. l987. Ph,vlog
onstrutd.Th signifianof th karyolasigont,n organllr systm in ukayoti nuloytpl
Netu Yorh dm of !
whih a nulus is mddd, was dsrid in th arlY yars of th twntith 'Wos,
.202. )n tlr vol
ntury (Janiki 1915). inimlly th karyornastigontonsistsof nulus,i.t Sicct'sUS 99: 8742-
nular otllltor (alld a rhizoplst in arly potozoologial litrtur)'irnd
an undulipodium (th 9+0 kintosom/ntriol and its shft, th 9+2 miro-
tuular aonm). (Som prf to writ this univrsal pattrn of ninfold
thology and th stu
symmtry of th mirottrulsin th shaft m()rauratlyas [9(2)+2].) Whn
th ntriol in mitosis dvlopsinto th kintosom of th motility organll, \/hil rlirny individuals l
w ar sing th lgay of th swimmr omponnt of th arlist ukaryots. Austrian Konrad Lornz
h strutur and lravior of prototists tht lak mitohorrdri undr (1907-1988) ar rgrd
anoi orditilnsand today's motility s,vmiossin many organisms with thology as a rognizds
swimming batria' sllh othrwis odditis and puliaritis, ar undrstood trvld to Lidn in 193
in th ontxt of thir volution. h nuleus itslf is most likly produt <lf ollaortors. L<rrnzwas
gnti-lvlintgation of batrizrlsymior-rts(arhtriLrm sulfidogn and his visionary br.rlth
plus swimming uatrium) tht gan as part of tlr karyomstigont from primntal appoah to th
whih it was rlasd. As wll as bing prat
This volutionary snario fo th origin of ukaryots is asd on th in- rvrot out its mthodol
valuai dsriptionsy Harvard profssoLmul R. Clvland (1892_|971I ogists slrould attmPt to a
and Harold Kiry Jr. (1900_1952),hairman of th Dpartmntof Zoo|og at vlop? Wht ar its imn
th Univrsity of alifornia at Brkly. lvland providd dtails of ll volv? Lrnz strssdt}
motility (movrnntsof th kayomstigontszrnd othr ll parts irr mitosis) having spontan()usly
and Kiry am to undrstnd :rlonymphid volutio y rognizing th ail- himslf shokd y th d
ity of karyomastigonts to rprodu indpndntly of th nuli in mitohon. in havioristi psyholog
drizrlll lingsthat proaly nvr aquird mitohondria. Prsrvationof vlopmnt of an thogra
th pulishd work of ths sholars (inluding th unprdrtd16--rm hvis-is th stating
blak and whit films y lvland) would of intrst to thos who wish to In 797, Lornz and T
dlv furthr into th details of ll evolution. Grman zoologist Karl vt
dislvisonrnig or
havior pattrns.''Von I
BIBLIOGRAPHY
languag of hony bs.\
lv]and,L. R., and A. V. Grirstne.1964. Th finstrtltllrof th flgllat ompartiv study of th
i<ltrihprd<li:rnd its ssoiatdmioorgnisrs. Prodittgs of th
onstrut thir phvlogny
RllSoietf L-ldo, SrisB 157:668_683.
Part II.Th gnr in te Atide (waterfo
Jniki,. 1915.aminationsof parasitiflagllats.
Dues<lui, Prioni,Stphttn<lmph, lomph. on th p:rraasl morphology in ronstru
appartlls.)n th oIrstitutinof tlrnulr"rs
urndamitosis.Zitshriftfr fr th ptanlf th
Wisssbftlih Zoologie | 1Z:57_691' arly thologistsWr
iry' .' lr.1941. Dvsovinid flgllatsof trmits.Uiursitof lifori
and Tinrgn mt at a W
Publititlsitt Zlolog45, nos. 1-5.
Mrgulis,L., . . Doln' and J. H. Whitsid.200.5...lmprftions dditis''
r-r] id of a fid ation pat
in th originof th nulus.Pillbiolog1', 1: 175-191. rpatdly liitd s
L., and D. Sagan.|998. Wht ls Se?Nw York: Simonand Shustr.
l!,Iargulis' t i u l rp k i n g h e v i t '
2000.'V/htls Lif? Brkly:Univrsityof lifrniaPrss. simpl modls of th adul
argulis, L., rrdK. V. Shrvartz,1998.Fiu igdms: Illustrtduid trthc o l l p tt r i . r f i d r i n
Pbl of Life lrtb.3rd d.Nw Yrk:W. . Frmanand Company.
s i g n s t i l n u l ia n d f i s d t
Samulsson, .I.,P. R. Darvs,and G. Vidl. l999. Orgni-rvalld mirofossilsfrom
th ProtrozoiThul suprgrouP'Northwst Grnlan1. PrmbriRsrll undrstood in th on
96:1-23. volutionary ontts fat
thrllg d the Stud of Bhuiorl uolutio 541
instint was strongly ritiizd in th 1950s y Amrian psyhlogistDanil ojts of study, lik a
.Wilson's
Lhrman for ing insuffiintly attund to dvlopmntal prosss.Tinr- ward o. xtn
gn's inlusion of dvlopmnt among his four qustions rprsntdhis at- insts to xplain huma
tmpt to aommodat Lhman's ritiqu. Susquntly' many yars of o8, a fild that also is t
fruitlss ..natuf vrsus nurtur'' dats nd onflits ovr dfinitions Anothr important of
hav mad urrnt thologists lry of th onpt of instint. Nvrth- funtion known as nut
lss, th study of inhritd havioal adaptations is still n lmportant part partiular haviorl tas
of thology. th startl rspons of fi
Aras of partiular urrnt intrst and ativity within thology inlud tional haring of owls,
prdator-pry intrations and th various systems of antiprdatory alarm mols.
alls found in a wid rang of spis, th soial dominan struturs that ..ognitivthol<
Also,
also vary widly aross diffrnt spis,ompartiv studis of soial play, to dsri th rintrodl
th mmory dmands of food ahing nd rtrival, tool manuftur and volutionary study of i
us, and th xistn of ulturally transmittd traits. onduting arful phys
thology an undrstood y ontrast to altrnativ approahs to th Nxt, h turnd his att
.!ithin
study of havioral volution in iology and psyholgy. iology, - M a n y d i s a g r w i t h h i s
havioral olgy applis population-lvl modls of volution to th undr- and anthropomorphi.
standing of animal havior. Bhavioral ologists frly orrow modls in human psyhology v
from onomis and gam thory, whih rprsnt individul organisms as instrumntal in gtting
rational optimizrs and allow analysis of intrations twn nvironmnt tions aot th volutr
and havior on rprodutiv fitnssand th distriution of bhavioral traits. ranging from ognitiv
Unlik traditional thology, havioral ology has n lss onrnd with primats.
immdiat auss of th havior of individual nimals or its dvlopmnt.
|LIoGRAPHY
hanisms rsponsilfor gnratingoptimal havior ar oftn simply as-
sumd and not plaind. Nvrthlss,thr hs n signifiant rossovr Bkoff,., . Alln,and
twn th two sudisiplins, with many of th originators of havioral mpiril nd Theor
N4ITPrss.
ology having first traind as thologists.
Burkhardt, k. 2005.Px
omparativ psyhology also taks an xpliitly volutionary approah to Fondig of thobg^
animal havior. Howvr, unlik iologists, who tnd to intrstd in dif- hny'D. L., and R. M.
frnsamong individuals and among spis,psyhologists ar traind to Soil id. hiag
lok for gnral pattrns of havior and prinipls of larning' somtims Frish'K. von. 1.967.h
.!7hr hadt'ik,trans.t
rfrrd to as laws of larning. omparativ psyhologists do s dif-
G r i f f i n ,D . R . 2 0 I . n m
frns twn spis, thy often rgard ths as rvaling stps on an vo-
Univrsityof hiag
lutionary laddr to full-lown human ognition, and thy ar oftn drwn to Lhrmn,D. S. 1953.A
invstigatingtasks to mark thrsholds of ognitiv sphistiation.Thus, for Qurtrl Ruituof
ampl, thr has n muh intrst among omparativ psyhologistsin Lindaur,. 1961.om
finding out whih spis an rogniz thmslvs in mirrors, an pass a UnivrsityPrss.
Lornz' K. 1981.The Fu
fals lif attriution task, or an larn a human languag.any thologists
Tinrgn'N. 1963.on t
prfr to osrv animals in natural or naturalisti haitats. Som sintists, ir1lshll ogi 20: l
to found on oth sids of this divid, ar pluralists who livthat th in- .Watson,
J. B. 1913.Psy
tgrativstudy of animal havior rquirs oth field and laoratory invsti- 158-177.
gation; othrs ar lss umnial. W i l s o n , . o . l 9 7 5 . S o
thology has had svral important offshoots. Both Lornz nd Tinbrgn
xtndd thir approahs to th study of humn havior: Lornz in his
study of aggrssin, and Tinrgn (with his wif lisath) in studying
autisti hildrn. Th willingnss of tholoists to trat humans as sintifi
thobg nd tb Std of Bhuklrl uolution 54
y Amrian psyhologist Danil ojtsof study, lik any othr animal, also importantly forshdowd d-
vlopmntalprosss.Tinr- ward o. \/ilson'stnsion of volutionary modls of th soial havior of
ur qustionsrprsntdhis at. insts to plain human soiality, thus giving ris to th fild of soioiol-
. Susquntly,many yars of ogy, a fild that also is tightly onntd to havioral ology.
and onflits ovr dfinitions Anothr important offshoot of thology is th omparativ study of nural
onpt of instint. Nvrth- funtion known as nrrrothology. Spifi adaptations of nural systms to
ations is still an important part partiular havioral tasks hav n dsrid in a varity of spis, suh as
th startl rspons of fish, th oordinatd voalizations of frogs, th dir-
'tivity within thology inlud tional haring of owls' or th unusual snsory mhanisms of star-nosd
jystms of antiprdatory alarm mols.
oial dominan struturs that Also, ..ognitivthology'' is th lal oind y Donald Griffin (1915200)
nparativstudis of soial play' to dsrith rintrodution of issusof ognition and onsiousnssto th
rtrival,tool manufatur and volutionary study of animal havior. Griffin mad his arly rputation
:d traits. onduting arful physial analyss of th holoation apabilitis of ats.
o altrnativapproahs to th Nt' h turnd his attntion o qustions of animal mind and awarnss.
psyhology.\ithin iology, - any disagrwith his approah us thy find it xssivlyandotal
ldls of volution to th undr- and anthropomorphi. Nvrthlss, th ognitiv rvolution that took pla
:ologists frly orrow modls in human psyhology was slow in oming to th thology, nd Griffin was
p r s n ti n d i v i d u a l o r g n i s m s a s instrumntal in gtting othr sintists to tak sriously th ida that qus-
:rtionsbtwn nvironmnt tions aout th volution of mind ould addrssd y studying topis
distriutionof havioral traits. ranging from ognitiv maps in honys to intntional ommuniation in
gy has n lss onrnd with prlmats.
ual animals or its dvlopmnt.
B|LIoGRAPHY
rl havior ar oftn simply as-
. has n signifiantrossovr Bkoff,., . Alln, and G. . Burghardt,ds.2002. Th ogitiuil:
mpiricl d TheortilPrs1letiuslnimI ogitio.amridg,A:
of th originators of havioral
I Prss.
Burkhardt'R. 2005. Pttrsof Bhuior:Konrd Lornz, Niko Tibrg,d the
lliitly volutionary approah to Fodigof thrllog,Chiag<r: Univrsityof hiago Prss.
who tnd to intrstd in dif- Chny,D. L.' and R. M. Syfart.2007. Bbootphsis: T uolutionof
is' psyhologists ar traind to Soil id. hiago:Univrsityof hiago Prss.
r i n i p l so f l a r n i n g ,s o m t i m s Frish,K. on. 1967. The De Lgugnd rittioof Bs.L. .
hadwik,trans.amridg,MA: HrvardUnivrsityPrss.
arativ psyhologists do s dif-
Griffin, D. R. 2001. iml inds: Bod ogitio to osiosess.hiago:
tsas rvalingstpson an vo- Univrsityof hiago Prss.
ion, and thy ar oftn drawn to Lhrmn,D. S. 1953.A ritiquof Konrad Lornz'sthoryof instint.e
gnitiv sophistiation. Thus, fr Qurterl Ruieluof Biolog 28: 337_363.
1g omparativpsyhologists in Lindaur,\4. 1961.ommuitiomogSoilBs.amridg,A: Harvard
UnivrsityPrss.
mslvsin mirrors, an ass a
Lornz,K. 1981. b Foudtionsof thology.Nw York: Sringr-Vrlag.
man languag.Many thologists Tinrgn,N. 1963. on th aims and mthodsof thology.Zitshriftfiir
l i s t i h i t t s .S o m s i n t i s t s , Tierpsologie 2: 414 63,
.\X/atson,
'luralistswho livthat th in- J. B. 1913. Psyhologyas th havioristviws it. PshologilRuie2:
, o r h6 l d a n d | o r a t o yi n v s t i - t58-177.
.Wilson,
. o. |975. Soiobiolog.amridg,A: Harvard Univrsityo,.,':-..o.
rots' Both Lorenz and Tinrgn
human havior: Lornz in his
his wif lisath) in studying
ists to trat humans as sintifi
544 ugis
tr and tmpramnt \,
ugnis
matrial (th natur of
ugnis is fundmntally th grrralization tO humans of th pratis of at irth. H lso assum
plant nd animal rdrs'who popagtth st and ull th worst of thei mnts, and that thos l
stoks. Th ida that suh mthods might applid to humans has a long produing at a shokil
history, raing ak at last to th anint Grks. Thus, in Plato's Repub- don to Prvnt it, ivil
li (1945,157-158), Sorats tlls Glauon that..anything lik unrgulatd tak harg of thir ow
unions would a profanation in a stat whos itizns lad th good life''' Th ida that huma:
and Glauln knowldgs ht h is arful to 1i1tonly th st of his nW-nor wr dat
sporting dogs and gm irds so as to ilvoid th othrwis rapid dtrioration losphr Jhn Stuart l
of his stok. man natur has takn
Apart from a fw fforts to systmatially shap human hrdity, s in th plial. | Priipls o,
program adoptd by th utopian ommunity at onida, Nw York' in th ..of all th vulgar mod
1860s, ugnis rmained largly onptual until th turn of th twntith moral influns on th
ntury. It ws only thn tht <lrganizationsxprssly dvotd to this aim th divrsitis of ond.
wr stalishdin th Unitd Stats,Britain, and Grmany. Thr ar multi- ill was swimming a6
pl rasols why th nd to implmnt ugnis on a larg sal am to laim that natur trurr
sm urgnt around this tim, ut prhaps th nrost important Was a nw al studis of inhritan
worry aout th possiility of iologial dgnration. Darwin, who wrot in
In his oz the origi of Spcis (1859), harls Darwin argud that im- agr with Franis Ga
provmnt in plants and animals rsultd from a fir omptitiv struggl, due only a small ff
.What
a pross napsulatdin th famous xprssion ..th survival of th fittst,'' tis ar innt.''
oind by Hrrt Spnr in 1866 nd latr adoptd y Darwin' Although anity in whih it ap;
Darwin did not disuss his own spisin the rigin' many of his ontmpo- popl that th Pros
raris wondd aout its impliations for humans. ln partiular, thy frttd that th mentally and :
ovr th possiility that a rlaxation of th struggl mant that human But whil Glton so
progrss would slowd, haltd, or vn rvrsd. Darwin himslf was with th risis, and th
dply disturd y this prospt' arrd in he Dscet of (1871,168) h than disouragingth
pulilv xprssdhis onrns out th ountrsltivffts<lfharity, ..p
what Galton alld
vaintions gainst smallpo, th uilding of asylums for th sik and in- any squamishnsso
san' and othr aoutrmnts of ivilizd Soity' noting that as a sult' middl-lass popl rr
..th
wk mms of ivilisd soitispropagat thir kind. No on who hrd to th prvailin1
has attnddto th rdingof dmsti animals will dout that this must th stat should kr
highly injurious to th ra of mn.'' govrnmnt into th il
Darwin's ousin, Franis Galton, ws among th first to xplor th soial om an antistatistwo
impliations of th thory of volution y natual sltion.Galton notd in am a mattr for lg
his mmois that h had n inspird y rading t rigi to pursu a th inrasing apta
long-standing intrst in th topis of hrdity ..and th possil improv- whih smd to ols
mnt of th Human Ra.'' In ..Hrditary Talnt nd haratr'' (1865), prvious to th nviro
pandd to th ook Hereditr Geius (1'869), Galton showd tht mn lin in p<lpulationqu
who had ahivd distintion in sin. litratur. and th law wr mor unfit for srvi in th
likly tlran mmrs of th puli at larg to hav had mal rlativs who Unitd Stats, th low
wr thmslvsminnt. In a raft of latr works, Glton tndd this r- and astrn urop ol
sarh and dvlopd what h saw as its orollris. Galton thought that his I smd to onfirm f
inquiris provd byond dout that all human ailitis and traits of hara- rs who wr flm
ugics 545
In Britain, th land f its irth, ugnisrmind mostly a mattr of prop- |869. Hrditri,
aganda. Howvr, in North Amrir, Sandinavia' Grmany, astrn urop, L l n d r l n :N l . l l l l i l ] l l l .
and Japn, among othr plas, laws wr passd prmitting th ompulsory \98. rtlricst
Gillhm' N. !. 2001' l
strilization of th flmindd, riminals, sual dviats, and othrs
Birth of ugis. >
dmd unfit to prorat. |n th 1927 as of Bh u. Bll, te U.S. Suprm K v [ s ,D . l . 1 , 9 9 5 . l t h
Court uphld th onstitutionality of ompulsory strilization in dision rdit. Rprin <J
.!7ndll
signd y hif Justi olivr olms' a politial progrssiv. fa- Kli, $7. 201,.Btliltlig
mously Wrot: ,.It is ttr for all th world if, instad of waiting to xut rn of th tttttr1,,
Prss.
dgnratoffspring for rim, or to lt thm starv for thir imility, soi-
M i l l ' J . s . [ 1 8 3 5 ]1 9 7 7 . i
ty an prvnt thos who ar manifstly unfit from ontinuing thir kind.'' In Sturt ill. Vll. 18' l
th Unitd Stats, ugni argumnts wr also invokd in support of rstrit- Llnivrsity of Tlrtlttt
ing immigration frorn Southrn and astrn uop, argumnts that playd a [ | 8 4 8 1 1 9 6 5 .I '
rol (aithough how larg a on is datd)in passagof th national_origins pplictkls to St
quota systm adoptd in 1924. But ugni zal rhd its apex (or nadir) in Joh Sturt il/. Vols
Paul, D. B. 2003. Dlvin,
Gmany, whr troitis gainst nrntal patints' Jws, and othrs wr jus-
..raial hygin.'' Radik, eds., Th l
tifid y th ostnsibl nd for a program of ambridg Univrsiti
As th as of olms suggsts'sllpport for ugnison appald to di- Plttl. l 945. h RpttllI
vrsonstitunisand was invokd in suppt of disparatagnds:rh d- Univrsit1.Pss.
..flov'' as wll as traditional marriag, Trrrda, .' and P. J. V/in
fns of paifisrrras wll aS War'
RiI Ntioli rr
and ass to irtlr ontrol information and dvis as wll as thir rprs-
ntral uopn Un
sion. In its hyday in th 1920s and 1930s, ugnis garnrd support not
just from soial and politial onsrvtivsut also from Faian nd vn
arxin soialists and rformrs of many strips. In tinr' ugnism to
vo|ution: Th
quatd with its worst instntiations'and aov all, with Nazism. As a r-
sult, th word is today usually rsrvd for pratis that th spakr or Julian uly had
writr dtsts.Thus ritis of prnatal diagnosis dm it ugniswhil its ad- ollat vast amounts of
voats strnuously rsist th lal. And lgislation barring fist-ousin mar- lutil: Th ldern S\,r
riags is not rgardd as ugnis,notwithstanding its ompulsory hartr. rat n nylopdir
Sltivrding is still pratid, and in som forms it is widly pprovd, ing his survy togtl-r
ut on lgay of ugnis'assoiationwith atroitisis a distrtd disours Chishd among ths
in whih w shrink from alling ths pratis y thir rightful nam. strssing sltion'sirn
spially in Britain' as
ILIoGRAPY lution.
Brorg,G., and N. Roll-Hansn,ds.2005.ugisdth WIfreStte: Tlr rrylopdiap
SteriliztioPoli in Dmrk, Sdn,Norta,ltd Finlnd.Rv. paprak votd svralhaptrst
d. ast Lansing:ihignStatLlnivrsityPrss. m a i n t a i n v r i a t i o r - rI.I
,1'927.
Buk u. Bll. 274 U.S. 200. gnom lvl' suh as p
arlson, . A. 2001. The Ufit:A Histor of Bd lde, old SpringHaror, NY:
ogy' Huly ppriat
old SpringHaror LaboratoryPrss.
his for.rson organisrrrs
Darwin, . [1859]|964. th rigi of Spis.Introdutiorr y . Mayr.
amridg,A: HrvardUnivsityPrss. inftlrration aorrtlo
'h
[1871]198l. Dseto|-n,nd Seletil
i Rltioro S' stttturdhis suv:v.
Introdutionby J' T. Bonnr and R' M. May. Printor.r, NJ: Printon I i o n .H s r r s s dt i
UnivrsityPrss. ation. lins wr his s;
.With
1958. Th Autobiogrphof hrlsDnui, 1809_1882: rigil
a dad for. Ths
missionsRestord.N. Barlow' d. Nw York: arourt Bra.
Gltorr,F. 1865.Hrditar,v tlntnd hratr.
mill'slgzi|2: v y s , r v i t h H u [ y t 1i r
1 5 7 _ 1 6 6' 1 8 - 2 7 . Th hif virtu of ths
vlution: Th Modrn Svnthsis 547
aind mostly a mattr of prop- |869. Hrditr Gis: Iquir itltl lts Ltt,s d osqs'
Lavia,Grmany' strn urop, Llndon: Marillr-r.
rssdprmitting th ompulsory 1'908. mor is of 1' Lif. Londt.ln Mthun.
Gillham, N. W.200l . l-if of Sir Fris Ito: Frlm fri 1llortio to th
s, sual dviats' and othrs
Birth lf ugzis. oford: oford Univrsity Prss.
f Buk u. Bll, te U.S. Suprm
Kvls, D. J. ] 995' ] th Nm of gis: Glttis d the Uss of Hum
rlsoy strilization in a dision Hrdit. Rprint d. amridg, MA: Harvard Univrsity Prss.
s,a politial progrssiv.H fa- Klin, W. 200,l'. Bildig Btter R: Gder, Sulit, d ugis from tb
if, instad of waiting to xut Tur lf th tttr to th Bb Bool. Brkly: Univrsity of alifornia
starvfor thir imility, soi- Prss.
Mill, J. S. l183.5l 1977. ti|tsation'In .|.M. Roson, d', olleted'V/orks of lohn
t from ontinuing thir kind.', In
Sturt ill. Vl. 1 8, sss rl Politis d Soit1,,Pt' I, 117-148. Toronto:
lo invokd in support of rstrit- Univrsity of Toronto Prss.
iurop, argumnts that playd a [1848l 1965. Prii1llsof Pllitil oom, ith Som of Thir
l passagof th national-origins pplitills to Srlil Phillso1l.ln J. . Roson, d., olltd Works of
al rahdits ap (or nadir) in Joh Sturt ll/. Vols. 2_3. Toronto: Univrsity of Toronto Prss.
Paul, D. B. 2003. Drwin, soial Darwinism, and ugnis.In J. odg and G.
i n t s J, w s , n d o t h sw j u s -
..raialhygin.'' Radik, ds., h mbrirlg om1lio tl Dri,214_29. anrridg:
amridg Univrsit Prss.
lr ugnison appald to di- Plato. 1945. Th l|publi of Plto. F. . ornford, trns. London: Oford
ort of disparatagndas:th d- Univrsity Prss.
,,Blood
a s w ] l a s t r a d i t i o n Im r r i a g , Turda, ., and P. J. \7indling' ds. 2007. d Homld'': ugertisnd
dvisas wll as thir rprs- Ril Ntiolism i trl nd Soutrn L|rop, 1900_1940. Budapst:
Cntral uropan Univrsity Prss. -D,B.P.
ugenisgarnrd support not
ut also from Faian and vn
rips.In tim, ugnisam to
uolutio: Th odr Sthsis (Julian Huley)
aovall, with Nazism. As a r-
r pratis that th spakr or Julian Huly had ptional skills as a ommrial writr. knw how to
sis dmit ugniswhil its ad- ollat vast amounts of informtion and how to xtrat ssntialpoints. o-
; l a t i o n a r r i n gf i s t - o u s i nm a - lutil: Th odern Sthesis (1942) is prft ampl. Huly didd to
nding its ompulsory harater. rat an nylopdiaof rnt dvlopmntsin volutionary studis.Hold.
m forms it is widly approvd, ing his survy togthr Wr a small numbr of arfully hosn thrads.
ttroitisis distortd disours hrishd among ths was natural sltion. Although h mad othr points'
s y thir rightful nam. strssingsltion'simportan ws Huxly's ntral aim. This ook srvd,
spially in Britain, as a training manual for th ..synthti''thory f vo-
lution.
icsd tb WelfrStt: Th nylopdiaapproah in uolutiol? movd systmatially.Huxly d-
,tu,nd Fild' Rv. paprk votd svralhaptrs to gntiphnomna,spiallythos that rat and
'rss.
maintain variation. H thn shiftd to dsri auss of variation at the
.!(iith
gnom lvl, suh as polyploidy. a kground in dvlopmntaliol-
d lde. o|d SpringHaror, NY:
ogy, Huxly appriatd th importan of ths phnomna. Shifting from
s.Introdutiony . ayr. his fous on organisms and ologial phnomna, uly nt onsidrd
information aout 1oalpopulations, varitis'and suspis.Thr thms
'lliuin RltilIlS, struturdhis survy:vriation, prosssof divrgn'and prosssof iso-
Printon'NJ: Printon lation. strssdth inrportn of polytypi spis and ggraphial vari-
ation. lins wr his spial intrst,a phnomna h had hlpd populariz
i' 1809_1882:With riginl
a dad for. Ths stions of uolutirln read simply as litratur sur-
:k:HarourtBra.
milln.sgzi|2: vys' With Huly trying to Ompil informtion s is don in a txtook.
Th hif virtu of thsstionsis thir radth.Hul mad tnsivus
548 volution:Th odrn Synthsis
of knowldg gland from his ontats' who spannd oth urop and th IBLIoGRAP|
nglish-spakingworld.
uly, J. |\942' 196
uolutiln Was no mr survey. Huxly plad natural sltion and adap.
Alln and Unwin.
tation solidly at th ntr of his study. Sltion dominats all volutionary
prosss'h insistd;it was th xplanation of first rsort. To sur, Hu-
ly idntifid a wid rang of oth iologial prosss(.g.,drift, isolation, volutionary ol
hyridization, and hanging mutation rats). frquntly gav thm ky volutionay ompu
rols in rtain onfind topis. Nvrthlss,sltionrmaind his most po- from iologial vo
tnt volutionary mhanism. For Huly' adapting to wid rang of sl- Taylor 1999). an
tion prssurs was lif's most frqunt pattrn. of possiilitis for
Huly onsidrd himslf a philosophr as muh as a iologist. Th final human programmr!
haptrs of uoltio onsidr volutionary trnds and progrss. uxly pr- Biologil voluti
sntd svral spifi (and ojtiv, h thought) ritria to masur progrss: diffiult prolms.
gratr ontrol ovr th nvironment' gratr indpendn from th nviron- normous numr
mnt, and gratr ffiiny. Th volutionary advantags of ths qualitis surviv and to rpro
smd ovious to Huxly. Natural sltion drov toward ths qualitis. Th origins of v
This onntion was ky. uly prsntd his approah as an improvmnt ( 1 9 5 0 ) s u g g s t i o no f
ovr his prdssors,who rquird suprnatural drivrs and guiding fors. volution.'' This os
His shm did not. Ovrall, humans srvd as Huly's pinnal of progrss, solutions to prol
with our intllign (onsisting of tru sph, onptual thought, and om- 1950s, svral grou
plx motions) apping volution's ours. Huly hld out for furthr approah.
progrss' too' suggstinghumans had a moral imprativ to ful that pross. In th lat 1960s,
uolutio nds with uly onsidring ways humans might at ithr to has- dpndntly introdu
tn or to str thir own volutionary futur. gnti algorithms (l
For svralgnrationsof iologists,uolutio providd an importnt his- and volutionary pr(
torial narrativ for volutionary studis.HuIy prsntdharls Darwin of suh approahs i
as a gnius who disovrd in natural sltion on of lif's ntrl prinipls. volutinary algoritl
This gnius was lipsd in th yars |880_1920, supposdly owing to Dar- Th simplst vrsi
win's failur to undrstand inhritan. Th ris of gntis solved th pro- odd solutions to a
lm. Its rnt growth, and th dvlopmnt of othr spialitis sin, vastly al vlu to ah in
improvd our undrstanding of partiula pis to vlution's puzzl. This th population to r(
narrativ mad Huly's rol sm natural. Th tim was rip, h said, for rossovr' and muta
somon to ollat th many sparatndavorsinto a omprhnsiv'syn- gnti algorithm it
thti whol ' uolutio was uxly's attmpt at that synthsis. uxly's ria is mt.
ovrall mssag was that Darwinism had rturnd in improvd form, a volutionary om
phonix rising from turn.of-th-ntury ashs. ntifi appliations,
uoltion dlivrd no rilliant nw insight. Instad, it providd a omp- root havior, mor
tnt survy of rnt dvlopmnts in volutionary studis. Within that r- suprsoni jt dsign
sarh ommunity' Huxly promotd natural sltion as an organizing An intrsting vri
thm. Latr gnrations followd th sam path and pointd to uolutil mig,was introdu
for its nthusiastisupport in that dirtion. viwd as a gnti
Huxly pulishd a sond dition of ullutio in 196. ollagus pu. truly utal ol
lishd a third dition, with a sond nw introdution, in \974.In ah, th dmonstratdin a v:
main haptrs rmaind ssntialIyunhangd. Rnt trnds in .
manuftur of phy
uolutior ompting d Gnti lgorithms 549
Thr ar two ways in whih a trnd in som fatur of organisms might hav
valu.Th fatur ould valal to us, or it ould valud y th volu-
ld than anint ons. ompar tionary pross' so to spak, in th sns of ing psrvd. From a sintifi
omparison is apt in that rtain standpointth first altrnativis problmati. In sinour prsonal or ul-
ar to th fist fossil orgnisms tural (rligious,soial) valus shold not rlvant or at last not ntral.
.h
notionof n ordring among Indd,thy ar a distration. For xampl, if th atual dfining dirtion-
trrangd living forms on a linar ality in evolution wr inrasing nrgy usag' an infatuation with som
:alld th sl tLrr or gteat faturw valu, suh as intllign' might aus us to ovrlook it. of
rftr Arisrotl, th hain was un- ours' w might intrstd in th volutionay sours of our valus
isms and thir rankings did not rathrthan trnds gnrally. For xampl, it might that intllignis fa-
Jan-Baptist d Lamark addd vord in volution by natural sltion, that w ar th produt of that
sms movd up th hain as thy trnd,and that w valu intllignon aount of its advantagsfor us. But
i viw hallngd th ordring it- hr too th valuativ omponnt of our intrst is sid th point, pt
ranhing and divrgn rathr as original motivation. Thus it is hard to s how progrss' if it is dfind as
; rognizd progrss' writing fa- a trnd in fatursvalual to us, ould a sintifially usful notion.
.t vagu yt ill-dfind sntimnt' Th sond altrnativ is to intrprt valual to man that whih is pr.
552 uolutior rogress
B
dld, vn rntly' by onfusion out
(A) A hypothtiallong-trmtrend in advanmnt'that is, prgrss.Lif
nt trnd thory it is not nough to say bginswith a singlspis,with som lw lvl o{ advanmni(thsigl
lar, o must spify whih of svral vrtialblak lin sgmntat th ottom lft).As tim passs,nw spis
n. maximttm. r minimum-aus arise,and som spisomtint,with hangouringin th origin f
diffrnt intrprtations. A trnd in th vrynW spis.Hr all hngsar inrss,so th trnd is said to
:grof advanmnt among all spis stronglydrivn. (B) Sam as A, with dotted lines addd to show th trndsin
gur, A shows a hypothtial long-term th maximum and minimum. () A wkl drivn trnd in whih inrass
pdominat,ut a numr of dasslso our. (D) A passivtrnd in
s,a singl linag,with som low lvl
'spisaris' and som spis om whih inrssnd drassar qually frqunr,ut lowr limit on hng
(thvrtialdottd lin) forslineagsto diffusto th ight.Stet for
: origin of vry nw spis.Th mn d i s us s i o n .
vl of:rdvnnrntof all spis in -
ras in that avrag,that is, movmnt
that a trnd has orrd' without any Disussion of progrss has not n onrnd with th istn of a
n th man' trnd, prhaps aus:1trnd is takn for grantd (dspitou inability to
l th dgrof advanmnt ahivd y masurit). Rathr, it has n aout th pattrn of hang, th undrlying
im,irr othr words, a ris in th highst mhanism. In figur , natural seltion strongly favors dvanemnr' so
: figur,B is th sam as A, ut annotatd that all hangs ar inrass.Th man, maimum, and minimum all in-
h dottd lin on th right.) Maima ar ras' Figur is th sam pt that th nt sltiv advantag is wak,
gnrallythought to rpsnt th most mning that sltion Somtims favors drass.(h lassi ampls ar
f s o , w o u l d p r s ntth m i m u m i n a th volution of rtain rdud forms, lik parsits.)Th sult is that lss
would b th last spison th right at advand spis prsist or ar rplaed y drassfrom aov' and th
.ndin th minimum is an inras in th minimum stays roughly th sam. h trnds in A and ar sid to
vandspis(th dottd lin on th lft driun, or mor prisly,A is strngly divn, whras is wakly drivn.
:rstausapparntly th lowst lvl Figur D is diffrnt in that thr is no ovrall sltiv advantag to inras.
|triahav istd, and indd hav dom- Inrassand drassour qually oftn' so th only ndnyis t sprad
histry of lif. (In othr words, it is lar symmtrially, oth up and down. Howevr' thr is a oundary, a lowr
)prsntationof th history of lif.) limit n dgr of advanmnt,that loks th sprad of th group on th
5 54 uolutionr rlgress
lft. This oundary an thought of s th lowst lvl of advanmnton- shold mo fit thl
..in tJ
sistnt with bing liv. Th rsult is that tir mn and maximum inras Darwin's trrns,
whil th minimurn stays th sam (as in th wkly drivn as).Th trnd is also knw, natul sl
said to e pssiu. on gologial timsal
Th urrnt ontrovrsy ovr mhanism gan in 1996 with ook on du just onstant h;
progrss y StphnJay Gould in whih h raisd th passiv mhanism to nss. On th othr
th lvl of plausiility. At issu is th rol of sltion in progrss, that is, adaptations, suiting tl
whth th (prsumd)trnd in th history of lif is wakly divn y sl- vord ovr thos mor
tion or mrly passiv.(A strong|ydivn mhanism is ruld out austh offer advantags on ti
tru minimum is thought to hav rmaind stal: atria prsist.) Gould Thr ar som iolo
h:rs bn widly nrisintrptd as laiming that volution is random, ut th suh as rtain llr"rl
irpliation of th passiv mhanisris not randomnss.Within ah volv- pathwys. But w do
ing linag sltion dos oprat, and organisms ar adapting. Rathr, th rigorous way, or how
implition is that volving linagsas a group r govrnd y many diffr- that is lar now is thz
nt sltiv fors, omplly onfigurd' so that sltion favors inrass ar possibl in prinip
and drassqually oftn. Th pross an undrstood s purly dtr- Th most widly r
ministi, lik th diffusion of a gs. As a possil asis ior
Unfotunatly,as will sn,thr ar hardly ny data availal,so from onntion with progr
a sintifi prsptiv th dat aout mhnism is taking plir in a nar usd in volutionay st
vauum. Still, thory has produd som insights.Fo ampl, it is now lar for progrss, suprfii
that in th dbat ovr mhanism two ommon osrvations ar irrelvant. sutly onnoting advar
Th first is th prsistnof nonprogrssiv spis,th stal minimrrm. This nition maks this usag
osrvation is not hlpful aus tlr minimum is prditd to rmain un- on an say anythinga
hangd in oth ass, passiv and wakly drivrr.Likwis, th is in th tain a thnial dfinit
maximum and th urrnt istn lf som highly advand spis,lik hu- srssof omplity. l
n-tans,ar not hlpful. A rising mimum is ptd in all ass. typs, numr of physi
dvlopmnt (sfigur
don, and with on -
A RND IN WA?
n dmntd.Nor
any andid:rtsfor a long-trnrtrnd hav n proposd: (1) fitnss;(2) in hang in ths vri
omplity; (3) aility to sns' ontrol' or rspond to th nvironmnt; (4) som vidn within
ody siz; (5) intllign;(6) vrsatility or volvability' maning th aility hs bn a trnd in nur
ttl hang in voltion; (7) th rvrs of volvaility, lso alld dgr of Th xption is <
ntrnhmnt;(8) nrgy intnsivnsso rat ofnrgy usag;and (9) dpth nstd, to som t
and sophistiationof mhanismsof inhritan'For th most pat advoats within whols' ovr
of all ths andidats hav takn a as-uilding pproah, with th rsult rs of lvls hav v
tlrat th litratur offrs a numr of lvr and somtims-powrfulargu- atria. Aout 2 bili
mnts in favo of ah. Unfortntly, som of th most intrsting-lik ioti rlationship to
fitnss, ompliry, nrgy intnsivnss, and aility to ontrol th protozoans suh as l
nvironmnt-ar diffiult to oprationaliz. Thus thr r hardl any lls, ut us th
data on th sal of lif as a whol, for any trrrd statisti, for ny of ths nxt lvl up. Thn
andidat varials. joind to produ th
For som of ths variabls, many wuld want to ount a long.trm trnd, ga), th anstor of
if on ould doumntd, as progrss.on varil is fitnss,ndrstood lias to muskrats. Fin
as aility to srviv and rprodu. [t might sm that Darwin,s prinipl of individuals joind to
natural sltion virtually guarants a trnd in fitnss. Latr organisms alld a ryozoirn. V
uolutior Progrss t-t i
,Wstlvl of advanmnt on- should mor fit than arlir ons' on th whol, having atn thm, in
i m a na n d m a i m u r n i n r s Darwin's trms'..in th ra for lif'' (Darwin 1859,363). But as Darwin
,aklydrivn as).Th trnd is also knw, ntural sltion produs adaptation to loal nvironmnts, and
on gologialtimsalsthshang dramatially.Thus sltionmight po-
gan in 1996 wit. a ook on du just onstant hang with no nt improvmnt' no nt inras in fit-
isdth pssiv mhnism to nss. On th othr hand, on long timsals organisms with gnral
lf sltion in progrss, that is, daptations, suiting thm to a widr varity of nvironmnts' ould fa-
lf lif is wakly drivn y sl- vord ovr thos mor narrowly adaptd. Ar thr gnral adaptations that
:hanismis ruld out arrsth offr advantagson timsalsof hundrds of millions or illions of yars?
stal:atria prsist.) Gould Thr ar som iologial mhanisms that may hav prsistd that long,
lat volutin is random, but th suh as rtain llular transport mhanisms and parts of rtain mtoli
.s7ithin
andomnss. ah volv- pathways. But w do not know how to idntify gnral adaptations in any
risms ar adapting. Rathr, th rigorous way' or how to tst whthr thy hav bom mor pvalnt. All
lp ar govrnd y many diffr- that is lar now is that oth altrnativs, a long-trm trnd and its asene,
l that sltion favors inrass ar possil in prinipl, and thrforthis is an mpirial issu.
undrstood as purly dtr- Th most widly rognizd andidt for a long-trm trnd is omplxity.
As a possibl asis for progrss it is rally a stalth andidat aus a dirt
rdly any data availal, so from onntion with progrss is rarly aknowldgd. But th way omplxity is
ranism is taking pla in a nar usd in volutionary studis suggststhat it funtions as a kind of od word
hts. For ampl, it is now lar for progrss, suprfiially valu fr and thrfor sintifi sounding ut still
non osrvations ar irrlvant. sutly onnoting advanmnt. Th asn of a widly known thnial dfi-
is'th stal minimum. This nition maks this usag problmati. Noody knows what omplity is, so
l u m i s p d i t dt o r m i n u n - on an say anything at all about it. Th litratur of rnt dads dos on-
lrivn. Likwis, th ris in th tain a thnial dfinition' or rathr svral dfinitions for ah of th svral
righly advand spis' lik hu- snssof omplxity. Ths inlud omplity in th sns of numr of part
ptdin all ass. typs,numbr of physiologial or havioral prosss'and numr of stps in
dvlopmnt (sfigur on pag 556). Howvr' littl mpirial work has n
don, and with on xption' no trnd that spans th history of lif has yt
bn doumentd. Nor do w know vry muh aout th mhanisms involvd
: bn proposd: (1) fitnss; (2) in hang in ths varials, passiv vrsus drivn. (On a smallr sal thr is
:spondto th nvironmnt; (4) som vidn within th animals, ovr th past 500 million yars' tht thr
:volvaility, maning th aility has bn a trnd in numr of part typs and that it has n passiv.)
olvaility, also alld dgr of Th ption is omplity in th sns of nstdnss.organisms ar
of nrgyusag;and (9) dpth nstd, to som xtnt' lik hins os, with multipl lvls of parts
n.For th most part advoats within whols. Ovr th history of lif, rganisms with vr gratr num'
ilding approah, with th rsult rs of lvls hav volvd. Th first living things 3.5 billion yars ago wr
and somtims-powrfulargu- atria. Aout 2 illion yars ago som atria joind togthr in a sym-
r of th most intrsting-lik bioti riationship to produ th first ukaryoti lls, similr to modrn
and aility to ontrol th protozoans suh as mob' Thy w suprfiially similar to batrial
z. Thus thr ar hardly any lls, ut aus thy volvd as olonis of atria, thy oupid th
trnd statisti,for any of ths nxt lvl up. Thn aout 600 million yars ago lons of ukaryoti lls
joind to produ th first multillular individual (proaly a kind of al-
vant to ount a long-trm trnd, ga),th anstor of all modrn multiIlulrs' from mushrooms to magno-
r variabl is fitnss, undrstood lias to muskrats. Finally' aout 480 million yars ago lons of multillular
smthat Darwin's prinipl of individuals joind to form th first olonial animl' a oral-lik animal
:nd in fitnss. Latr organisms alld a ryozoan. odrn olonial organisms inlud ryozoans, orals'
5.'6 uoltionr Progrss
Myr. . 1988.Tord,
u lluti li st.rn
M S h a 'D . W . 1 9 9 1 .
d hiloslph6, n
1 9 9 8 .P o s s i lla
hypothss... u
RihrdsR . . J . | 9 9 2 . h (
d ldologil R
hiago Prss.
R u s ' . | 9 9 6 . , , n u 7' ,
ambridg,A:
Simpson,G. G. 1949.Th
Prss.
S p n rH' . l 8 5 7 . P r o g r
Stins'G. L. |969, h
North arolina Prs
Van Valn,L. 1973.A n
Vrmij,G. J. 1987.u
A lassi ampl of why dfinitions of progressin trms of movs to Prss.
omplity ar prolmati.Th uppr skltonis of a lion and th lower
skltonis of a whale. By any measurmntth akonof th whale is th
simpler' and yt adaptivly the whl's akon is what is nddfor a martn
volutionary psy
mammal, just s th lion's akon is what is ndedfor a trrstrialprdator.
Whals dsndedfrom land ttrapods(four-lggdanimls)sO thir S o m h v i o r |s i
bkonsmark a sltion-drivn move from omplityto simpliity.(From study of human hav
MShea 199|. |5.\ In this road sns,v
q u i r y a r d f i n dn o t l
ut y th kinds of qu
fr from paradigms' W
th soial insts, and many vrtrats,from fish to humans. ! do not
ologis with whih th;
know if vn highr lvls-olonis of olonis-hav n rahd. Thr
a fild of inquiry, volu
ar no lar-t ass. Humans sem to ssoiat at many lvls' ut only
I i t i o no f h D s
wakly at th highr ons. Intrstingly,if humans hav not tuly rahd
i n d Aimls in
th olony-of-olonis lvl, nd if no othr organism has ithr, this raiss
t i u l t d p a a d i g m s
a novl possiility: th only varial in whih w an doumnt a trnd,
d i s t i n t t h o r i s o u t
omplxity in th sns of nstdnss,in fat dos show a trnd, ut that
human havior and m
trnd might hav ndd, prhps 480 million yars go. Prhaps th ag of
Th first of ths pa
progrss is ovr.
th 1970s. Th or id
BIBLIOGRAPHY n a t u r a Ia n d s u |s I t
many spis hoos m.
Ayala,F. J,1974. Th onptof iologialprogrss. In F. J. Ayala and
T. Dobzhansky,ds',Studisi th Philostlphof Biolog, 39_354.London: ml ourtship display
\4amillan. sltion will tailor m
Goudg,T . ^. 1961. Th Asent of Lif. Toronto:Univrsityof Toronto Prss. w a . s o i o i o I o g i s ras1
Gould' S. J. 1988.On rpiaingth idaof progrsswith an oprafionlnotionof rsn in a populati
dirtionality.In . H. Nitki, ed.,uolutilr
Progrss,319-38.hiago: thory. Aordingiy' hr
Univrsityof hiago Prss.
nations of how huma
1996. Full Hose: Th Sprdlf IlefrlmPIto to Dri'
Nw York: Hrmony Books. anothr-to plain hr
Huxly'I. S.1942. uoltio:Th odrn Snthesis. London:Alln nd Unwin. h i s , f o r m a n i f s t i n gl
uiy,T. . 1893.uolutilnd tbis.London:amillan. th ntrl thorti
uolutior Psholog 557
in trmsof movs to
Prss. -D.W.Nr.
is of a lion and th low
akbonof th whl is the
i s w h t i s n d dt o m a r i n volutionary psyhology
reddfor a trrstrialprdato.
4gdanimals)so thir Som havioral sintists dfine volutionary psyhology as simply th
omplitvto simpliity.(From study of human havior and psyhology from an volutionary prsptiv.
In this oad sns,volutionary psyhology is fild of inquiry. Filds of in-
quiry ar dfind not y speifi thoris aout th phnomna thy study,
but y th kinds of qustionsthy pos aout thm. Filds of inquiry thus dif-
fr fronr paradigms, whih r dfind y th spifi thoris and nrthod-
lm fish to humans. !7 do not ologis with whih thy answr th qustions that dfin a fild of inquiry. As
ris-hav n rahd. Thr a fild of inquiry, volutionaypsyhology gan with Charls Darwin's pu-
iat at many lvls, ut only liation of Tbe Desct of in ] 871 and Tb pressilll rlf the motions
tumans hav not truly rahd in d ils tn 7872.It was not until a ntury li1tthat larly ar-
o r g a n i s mh a s i t h r ,t h i s r a i s s tiulatd pradigms gan to emrg in volutionay psyhology, ah with
ih w an doumnt a trnd, distint thoris about' and spifi mthods for studying, th volution.of
t dos show a trnd, ut that humn hvior urndmntillity.
1 yars ago. Prhaps th ag of Th first of thes paradigms was human soioiology, whih mgd in
th 1970s. Th or ida of soioiology was that havior has volvd undr
natural and sul sltionIusts org:rniform has. For ampl, fmalsof
many spishoos mats sdon th quality of mal or-rtship displays. If
:ss.In F. J. Ayala and
mal ourtship displays vary in quality, and that variation is hritabl,sual
, of Biollg, 39-54. London:
sltion will tailor mal otltship havior to fmal prfrn.In this
Univrsityof Toronto Prss. way, soioiologistsargud' a form of havior an an ildaptation' whos
ss with n opfational notion of prsnin a population is to plaind y th prinipls f volutionary
r Progrss,319_338. Chiago: thory. Ardingly, humn soioiology sought to offr volutionary pla-
nations of how humns ar havioraily adptd to soial lif with on
fro Plto to Dri.
anothr-to xplzrin human havioral adapttions for dominan hirar.
. ' s i sL. o n d t l n :A I l n a n d U n w i n . h i s , f o r m n i s t i n ga r t d d a l i n g w i t h a g g r s s i o n , n d o m a t i n g . l n d d ,
n;amillan' th ntrirl thortial prolr-rof humirn soioiology Was to plain th
5.'8 uolutitr Psholog
ILloGRAPHY
Barrtt'L.' R. Dunr, and J. Lytt.2002. Hum uoltionrPsholog.
Printon'NJ: PrintonUnivrsityPrss.
Bullr, D. J. 2005. Adptig ids: uolutiorPschologd th Prsistt
Qustior tlltNtur.mridg,N{A: IT Pss.
-h
Buss' D. |\^.2007. uolutior,Psholog: Nu.lSiof th id.3rd d.
Boston:Allyn and Bon.
Laland, K. N., and c. R. Brown. 2002.Ssd Noss:uolutionr
Prsptius lIItBhuiour.oxford: oxford UnivsityPrss.
.!ilson, Adaptiv landsps.Tl
. o. \978. um Ntre. amridg,]vlA:Harvard UnivrsityPrss.
-D.I.B. th tops of hills f adap
volutionary mt;rphor<
pssiblthat Wiglrta
..volution in N1ndlianPopulations'' (Swall Wright) aldan, ..A thmat
tstablPopuIation
Th lassi papr ..volution in ndlin Popultions'' y Amrian popu- 28 (191),17-,|,42'w|
fitnss,drawn y tlr g
l a t i o n g n t i i s tS w a l l W r i g t w a s p u l i s h d t n G e t i si n 1 9 3 1 . I n i t , a n d
Provin points out that t
in ..Th Rols of Mutation, Inrding,rossrdingand Sltionin vo- information aout th
lution,'' a shortr papr givn the following ya at an intrntional gntis ultimatly,th landsap
..daptivland-
ongss'in whih h introdud his famous mtaphor of an
..shifting
sap'' (s figur), \/rightprsntdis alan thory of volu-
tion.'' His fllow population gntiist aross th Atlanti' R. A. Fishr
(1930), hzrd just givn his no-Darwinin aount of volutionary hang, would working agai
..gntidrift''' lso at
on that saw suh hang oling s th rsult of trlutatinsinto a larg pop-
ulation, and sltiln thn liminating all ut th st. Wright had workd that rnathmatis sho
for a numr of yars at th U.S. Dpartmnt of Agriultur. From studisof may wll ountr th r
th rding of attl, h was orrvind that th st way to gt hang is by without (or vn dsp
isolating small goups, working within thm to gt th st of all possil slvs of grat valu
haratristis, and thn finally rintroduing th highst quality to th whol lutionary hang.
hrd (Provine 1986). his is a vry non-D
.Wright's undr th influn of
shiftirrg alan thory followd th sam pattrn: isolation of
groups' improvmnt within th group, and thn mingling with th whol. In natural sltion. (Do
his ongrsspaper, !(right rprsntdth prossas somthingourring on Species, first pulishd
a landsap,with th st (th fittst)ing at th tops of pks nd th worst tins gv mo rdn
down in th vallys. H saw volution as ing a mttr of moving from on was doing on hromt
pak to nothr highr pk. Th hallng was how to gt down th sid of 1999.) In part th infl
th first pak so that on ould start liming up th othr. Bussltion omplaind that natur
,,uolution
i ndeli Pooltios'' 561
uoltiorPsholog.
shologd tb Persistent
[T Prss.
of tbind.3rd d.
Silt,e
(oss: uolutior
lford UnivrsityPrss.
daptiv landsaps.This pitur, showing how th fittr organisms lim to
'A:rvaro
U"'*.YJT.; thtops of hills of adptivfitnss'illustratsprhapsth most famous
volutionarymtphorof th twntithntuy(from lfright 1932).It rs
possilthat Wrigt am to tlr mtaphorfrm rdingan artily B. S.
J.
Hldn,..A athem:rtialThory lf Artifiial and Ntural Sltion'Prt VIII.
;'' (SwllWright) tstalPopulations,'' Proedigsof th mbridg Philosophil Soit
rulatiorrs''y Amrian popu- 28 (I9|' \7_142, whih ontainsa pitorial reprsntation of paksof
fitnss,drawn y th gntiist. H. !7addington. Wright,s iograph.!7illiam
in Getisin 1931. In it, and
Provinpoints out tht th landsapis odnsinga hug amount of
rdingand Sltion in vo-
infomationaout te fftsof diffrntgnsnd that onsquntlv,
r at an intnationag l ntis ultirnatly,
th landspmy not ntirlolrrnt.SProvin l98o.
rtaphorof an ..adaptiv land-
[ting alan thory of volu-
ss th Atlanti, R. A. Fishr
:ount of volutionary hang, would working against this' !right introdud th ontrovrsial notion of
..gnti
of mutations into a larg pop- dri{t,'' also at tims known as th ..Swall Wright fft.'' H argud
th st. \7right had wokd that mathmatisshows that, in small groups, th han fftsof rding
lf Agriultu.From studies of may wll ountr th ffts of sltion, so nw faturs an l.l produd
r bst wy to gt hng is y without (or vn dspit) natllral sltioIr.Tlrs nw fatus may thm-
to gt th st of all possil slvs of grat valu, spiallyin th whol, and an ld to frth vo.
h highst qulity to th whol lutionary hang.
This is a vry non-Darwinin thory, although latr vrsions, partiularly
:h sam pattrn: isolation of undr th influn of Thodosius Dozhansky' gav a muh largr rol to
n mingling with th whol. In natural sltion. (Dozhansky's mastrwork, Getics d tb rigi of
jssas solthing ouing on Speies,first pulished irl 197, gav strong mphasis to \/right.Latr di-
,htops of paks and th worst tions gave mor rdnto natural sltion,thanks tl studis Dozhansky
] a mattr of moving from on was doing on lrromosomal variations of fuit flis in ntur. S Rus
ls how to gt down th sid of 1999.)|n part th influn on Wright m from nri Brgson, who had
up th othr. Baus sltion omplaind that natural sltionnnot yild innovations. Br.rta f ar greater
562 uolutio of Lguge
influn was Flrrt Spnr (partiularly as passd on y !right's tahr' opn-ndd varity oi r
L. J. Hendrson). Populations r sn in a kind of balan, homognous; r o f m a n i n g , l spsh o
when somtlrigdisrupts th balirn,ausing frgmntationand htrogn- nit st of maningful
iry, th whol strivs to rturn to homognity' t a highr lvl' This is a f l x i v n s s i,n w h i |
lassi as of Spnriandynami quilirium. gua8'nd prvarieti
In rnt yasl th shifting aln thory has n vry svrly riti- sags. Although langu
izd, although rrstvolutionistsstill think that thr is som rol for gn- Kanzi, show subtl sig
ti drift (sCon, Barton nd Tr'rrlli1'997).At th molulr lvl, nath t h i r s i g n do o n l P u t
th ffts of natural sltion, drift is thought to b vry importnt (Kimura muniations rmain
1983) and is th asis of th major wy of alulating signifiantdats in vo- By th tim humal.r}
..gnti ..Molular volu. on hr parnts and ask
lution (through loks''). (S th main ssay
tiorr'' y Franiso J. Ayala in this volum.) y a r - o l d h i m p a n z ,
munirionon th orr
B|LIoGRAPHY tl.rirlanguag skills to
Brgson'H. 1911.rtiuuollltil.Nw York:Holt. produing th full rang
oyn,J. A., N. H. Barton'and . Turlli.1997.Prsptiv: A ritiquof Swali hominins volv lngu
.Wright's
shiftinglanthoryof volution.uolutkl51' no. 3: 64-671. from th anstorsof tl
)ozhansky, T. [|97] 1951,Getisd th rigi of Spis'rd d. Nw York:
Shllirrshav rrt
olumiaUnivrsityPrss.
Fish'R. A. 1930' he GticIThor of Nturl Sltion.oford: Oford win's tim. Nonthls
UnivrsityPss. m o u s l y a n n d a I I t - ' r
Hndrson,L' J. 1917.Th rdr lfNtre.mridg,MA: HarvardUnivrsity languag ausof its
Prss. nams that hav bn g
,b -rlr1'
Kimura, . 1983. Nutrl of oleulr ullutio.amridg: sis, for xampl, sl-t
amridgUnivrsityPss. ..moo,
sounds suh as
Provin,!. B. 1986.SuIIWrightttltdurllutilrl,Birllog.hiago:Univrsity
lf hiago Prss. attiuts th nrg,
Rus, . 1996.ond to : Tb oeptof Progrssi uolutilrBiolog. atd with physial l;ro
Camridg,MA: Harvard UnivrsityPrss. originatd from moti
1999.ster of stris:Is uoltion SliIostrutil?mridg' h y p o t h s sd o n o t l n d
A: Harvd UrrivrsityPrss.
Notions out wh
Snr,L7.1862, FirstPriipls.London:Willimsand Nrgat.
Wright,s. 193l. volutionin \,lnd|ian populatins'Gntis16:97-159. today as thy wr a
192.T|.lrolsof mutation,inrding,rossding and sltionin millions of yars old;
volution.Prodigsof th Sith ltrtiologressrlf Gtis1: Among th lattr. so
356-366. . -.R. arisn bfor th
..ultu
was hratrizdy
als, prsonal :rdom
volution of languag Uppr Palolithi tooln
Although many animals ommuniat using voalizations' only humans us to ston' and thir too
ompl languag, asd on a disrt numr of aritrary its of Sounds mor spilizd for p
and gsturs' to omprhnd and produ a smingly infinit numr of agr that Uppr Palo
novl mssags.Listrringto sph nd rading r snsory aspts of ln- 40,000 yars ago' thy
guag' whil talking, writing, and ody languag ar motor aspts.In th o l o g i s t s l i v t h l r t t h
|960s, hrls Hoktt listd 15 univrsal dsign faturs that distinguish sumption of linguisti
.W
human languag from ommuniation systms in othr animals. oktt n plad on th
viwd thr fattrrsas partiularly importnt for lruman languag:(1) dis- std, that numrol1s
plzrmnt,or bing al to ommIriatabout mattrs that ar rmot in points, vidn of tr
tirn and sp; (2) produtivity, whih is tlr ability to ommuniat an o u r r dm u h r l i r '
uolutio of Lgug 56
passdon y \/right'stahr, opn-nddvarity of novl mssags;and (3) duality, in whih a small num-
.ind of alan, homognous; rof meaninglssphonologial lmntsmay omind to rat an infi-
fragmntation and htrogn- nit st of maningful uttrans.Two othr intstingdsign faturs ar
ity, at a highr lvl' This is a flivnss,in whih on an us languag to olmuniate aout ln-
1. guag,and prvariation,why languagan sd to dlivr flsms.
y has n vry svrly riti. sags.Although langug_traindaps suh as th wll.known onoo,
h t t h r i s s o m r o l f o g n - Kanzi, show sutl signs of utilizing som of Hoktt's dsign fturs in
At th molular lvl, nath thir signd or Omputr-failitatd ommunitions with popl, thir om-
t to vry important (Kimur munitions rmain xtmly simpl dspit dads of intns shooling.
u l a r i n gs i g n i f i a n d
t tsin vo- By th tim a human hild is four years old, sh is likly to fastn a ady y
ain ssay
..olular volu- on hr pants nd ask, ..\/hrdid I om fronr?'' Th highly traind four-
yar-oldhimpanz,on th othr hnd, is likly to sign a spontanousom-
nruitionon th ordr of ..ooki,ooki, hurry.'' Aps, in ft, nvr us
thirlanguagskills to ask qustions.Furrhr, thir larynxs ar inapl of
Holt. produingth full rang of human sphsounds. How and whn, thn, did
Prsptiv: A ritiqu f Swll hominins volv languag during th 5 to 7 rillion yars sin thy divrgd
u o l u t i o 5 1 ' n o . 3 : 6 4 _ 6 7 \ . from th anstorsof thir losstgntiousins, th himpanzs?
,igi of Spcis. 3rd d. Nw York:
Sholars hav n fasinatd by this qustion sin t last harls Dar-
rl Slectio' Oxford: oxford
win's tim. Nonthlss,in 1866 th influntial Linguisti Soity of Paris fa-
mously annd all onrmuniationsdaling witlr tlr origin and volution of
r b r i . l g . A : e r r . n r dI I n i v s i t y languagausof its spulativnatur, whili is valdin th amusing
' namsthat hav r-r givn to various hypothss.Th ..ow-wow'' hypth-
r u,,Itil,C.lrrlhridg: sis, for ampl, suggStsthat languag gan as imitations of natural
sounds suh as ..rnoo''' ,,ozz,'' nd ..rash'';th ..yo-hav-ho''hypothsis
r Bi,,log.Chig<-l:
Univrsiy
attriuts th mrgn of languag to rhythmi hants nd gunts assoi-
'rogrssi ulltior Biolog. atdwith physial laor; and th ..pooh-pooh'' hypothsis suggstslanguag
originatdfrom motional ris or lamations. Although intrsting,ths
Soil ostrution! Carrrridg, hypothssdo not lnd tmslvsto sintifitsting.
Notions aout whn nd lrow languag first aros r as ontrovrsial
ms nd Norgar.
tody as thy wr il ntury ago. Sonl rsrhrsthink tht larrguagis
n s .G l i s | 9 7 _ | 5 9 .
: r o s s r d i ngn d s I t i o ni n millions of yars old; othrs liv it originatd a mr 40,000 yars ago.
l ogrssof Getis'-,.*. Among th lattr, s<rmarhologists think that languag ould not hav
arisn for th ..ulturl plosion'' of th Uppr Palolithi in urop that
was haratrized av paintings,portal art, living sits,dliraturi-
ls, prsonal adornmnts' hunting and fishing thnology, and signs of trad.
Uppr Palolithi toolnrkrs usd and ngravd ivory and bon in addition
voalizations,only humanS us to ston' and thir tools wr ttr mad than rlirons and frquntly
r of arbitrzrryits of sonds mor spilized ior prtiuiar tasks. Althor'rghmost anthropologists wold
a smingly irrfinit nmr of agrthat Uppr Palolithi innovations indiat that languaghad zrrisny
ling ar snsory aspts of lan- 40'000 yars ago' thy disagraout whn it iniially mrgd.Som arh-
]uagar motor aspts. In th ologistsliv that th prior matrial rord is too simpl to justify an as-
dsign faturs that distinguish sumption of linguisti haviors. Othrs think that too muh mphasis has
'Wstrn
ms in othr animals. Hoktt n plad on th uropan Uppr Palolithi; thy suggst, in-
nt for human langug: (1) dis- stad' that numrous potntial signs of languag inluding ard spar
) o u t m t t st h a t a m o t i n points, vidn of trad ntworks, and th r'rsof rd pigmnt (h)
t h i I i t vt o o m m u n i a t n ourrdmuh rlir (prhapsas long ago.1s 10,000 yars) in th Afrian
564 ultil<lfI'gg
rord of th Middl Ston Ag. Rathr thar-rsing |anguag as having vrrtuallyinvdd lang
mrgd suddnly and rlativly rntly,thy livit volvd mor gradu- t r g to n a t u r l 5 l t
ally ovr muh longr priods of tim. x p l i i t l y r j h n o t i
othr sintistshypotlrsizthat langug lrigirrrdstill urrlir.Skltal all systrns,lthor-rglrt
iologists who study th volution of th parts of th skull ald nk that f- . . t o p - d t l w n ' 's h o l o f t
ilitat sph livthat th laryn and throat had volvd to thir modrn mans volvd th ailit1
forms y 150,000 yars go' thus alllwing th rapid vlalizationsthat har- infinit numr oi rul-
aterize modrn languags.Thy lso:rknowldg that sph rrdnot ntail nuity hy;lothsis,''on th
th full rang of ontmporry voalizations nd that rudirrllrtarylanguag of inrmntal volutior
may hav istd at lst a million yars ago. and ss its roots rflt
Palonurologists,who study th volution of th rain, liv that lan. wild primats hav in t
gug mrgd vn arlir, around or for 2 rnillion yilrs ago. Thv man) xtnd th disuss
s t u d y a s t s o f t h i r r s i d so f f o s s i l s k u l l s ( n d o a s t s t) h t r p r o d u d t a i l s alls twn rnothr t
from th outsid part of th rain (or rral ort) that rvr imprintd Nl disussion of lan1
on th walls of th rains whn hominins (or othr animals) wr aliv. ing th wll-known gs
Thy also lrart th inras in rni:rl apaity (as a sur<;etfor rairr on th dp psyholtlg
siz) that ourrd durir-rgth ours of hon-rinin vOlLltion. Altlrtlugh guag' iniudir-rgmoth
spaking dpnds on ativity in vrirlus parts of th rain, I}roa'ssph til, and gsturl omp
aa on th lft sid of th frontal los is spially important for oordi- w also us flailing han
nating th intrit movmnts of th tongu' mouth) and larynx tht r physil ontt rr,ithp
nssary for sph. Broa's ara is assoiatd with a spifi pattn of mats also tlmirrsu
onvolutions that dos not appr in th rins of our losst gnti siorrLrllyolfatoy signa
ousins' th grat aps. This prt of th brain is somtims rprodud on diffrnis that, in ad
ndoasts. ints rvald on ndoilsts from f<lssilhominins suggst th:rt primilt produs voa
th frontal lo sph ar:rmay alrady hav gurrdvlopirrgy 1.9 mil- abov. Alvotslf g
lion yas gl. Studis f ston tools from th sam tim lso suggstthat v r l t u a l l yt l I l g . u gi
hominins wr in th pross of oming right-handd, whih aps as a 6nd narrowly (.g.,mz
population 21rnot. It is important to not, howevr, that rains had not yt tht rodu visuzrl,
nlargd to nywhr n:lr thir molrn sizs, and that th vi]nfronr M 1 ' o w n i d s. r r i n
ndoasts rgarding Broa's ara is diffiult to inrrprt. Th suggstion of livthat th oots o{
rudimntary languag t this arly point in tim should thrfor viwd rsnts a kind f sign
as intrsting ut spul:rtiv. mrgn of th first 1
Languag and populatior-r-lvl righr-hnddnssoth dpnd lravilyon th himprrzmothrs us
lft sid of th rain. Nonhuman pimatshv r:rinsthat a lso asynrnrtri. lnothrs dlivr e ons
al (or latralizd), to som dgr, ut thy do not gin to pproah human ..mothrs,''or tnusi
lvls of nurologial asmmty. In ddition to lirnguag funtions, th lft fatttrshlp aisr
hmisphrs of humans r diffrntilly invlvd irr skilld movmnts' suh gugs.himpanz m
as thos of th right har-rd, as wll as analytial,tilrr-squning prosss.Th tir infants, who rm
right hmisphr,n th othr hand, lsat mor gloal purstlits,suh as vi- unaidd' to thir hairy
suospatial and mntal imaging, ognizing fas, appriating humor, and voltltion of walking u
haring and produing mr.rsi.It also ontrols th lft hand. Although it is th thy do rtain a grasp
ltt sid f th rain that spks, it is tlr rntrsilright hmisphrthat pro- t h i h l p l s si n f l l l l t ' i l
vids th ton of voi, or prosody, lvhih olors our uttrans.In this sns, Un]rths irumsta
languag dpnds on th right as wll as th lft hrnisphr. adaptiv for assurin
Som sholars' howvr, think that prosody hd nothing to do with th movd from thir rnot
mrgnoi lnguagal-rd,instad,dvlopds ll kir-rdof uditor1'frill tht kirrd lf dismodr
uolutil of Lgug .'6.'
tan sing languag as having vntually invadd lnguag.Thy also assrt that musial skills wr not a
r iivit volvd more gradu- targt of naturl sltionin th sme way that linguisti ailitis w, and
pliitly rjtth notion that languagould hav volvd from primatlik
originatdstill arlir. Skltal all systms' although thy aknowldg that ton of voi likly did. This
s of th skull and nk that fa- ..top-down''
shool of thought thrfor fouss on undrstnding how hu.
lt had vllvdto thir modrn mans volvd th aility to link symoli rfrntialsounds into a potntially
rapid voalizationsthat har- infinit numr of rul-govrndmaningful sntns.An opposing ..onti-
dgthat sphnd not ntail nuity hypothsis,''on th othr hand, posits that languagmrgds a rsult
and that rudimntary languag of inrmntalvolution from th primat all systmsof our erly anestors
and ssits roots rfltdin th ominations of prosodi alls that various
l o f t h r a i n , l i v t h a t l a n - wild primats hav in thir rprtoirs.Som ontinr-ritists(.g.,John Nw-
r 2 millin yars ago. Thy mn) tnd th disussion vn furthr into th past y fousing on ontat
coasts)that rprodu dtails alls btwn mothr nd infant mammals.
'al ort) that wr imprintd No disussion f languagvolution would mplt without mntion.
; ( o r o t h r a n i m a l s )w r a l i v . ing th wll-known gstural.originshypothsis,whih rognizsand uilds
lity (s a surrogat for ain on th dp psyhollgiallinks twn thought, sph,and gstur.Lan-
hominin volution. Although guag' inluding mothr-infant intrations' nompasssvoal, visual, ta-
ts of th rain, Broa's sph til, and gstual omponnts. Humans do not just ..talk'' with our vois;
spiallyimportant for oordi- w also us flailing hands, faial prssions,ody postur' and (somtims)
l' mouth' and laryn that ar physial ontat with popl or ojts to onvy maning. Nonhuman pri-
rtd with spifi pattrn of mats also omin suh gsturswith motional voaliztions, and oa.
r i l l s o o u r | o s s tg n i sionally lfatory signals, in thir blnd of multinrodal ommuniation. Th
in is somtimsrprodud n diffrn is that, in addition t motionl alls and inttlnations, th human
n f o s s i l h o m i n i n s s u g g s tt h t primat produs voliztions that hav th linguisti proprtis outlind
b g u nd v l o p i n g y 1 . 9 m i l - aov. Advoats of gstural origins think that th natural sltion that ld
h sam tim lso suggst that vntuallyto languaginitially targtdgsturalommuniations, thy d-
right-handd,whih aps as a find narowly (..g., manual gsturs)or mor roadly (rtiulatory gsturs
owvr'tht brains had not yt that produ visual, aousti,voal, or tatil signals).
s, and tht th vidn from My own idas ar in kping with th gstural hypothsis,writ roadly. I
to intrprt.Th suggstion of livtht th roltsof languagar tmlyanint and that prosody rp-
im should thrfor viwd rsnts kind of signatur that is lft ovr from vnts that prdd th
mrgnof th first languag.Although th ody langugthat human and
lnssoth dpnd havily on th himpnz mothrs us with thir infants is virtually idntial, only human
- , r a i n sr h a t r l s o s y m m r i - mothrs dlivr a onstant stram of ay talk, whih is vry prosodi. Suh
o not ginto approah human ..mothrs'''rrrusial
sph,is univrsal among humans, and its eousti
to languag funtions, th lft fturshlp ais round th world larn th strutursof thir nativ lan-
lvd in skilld movmnts, suh guags.himpanz mothrs dirt fw, if any, spial voalizations toward
, tim-squningprosss.Th thir infants, who rmain physially atthd to thir mothrs y linging'
moglobal pursuits' suh as vi. unaidd, to thir hairy hsts.Bausof dvlopmntalfftsrlatd to th
fas, appriating humor, and volution f walking upright, human infants hav lost this aility, although
th lft hnd. Although it is th thy do rtain a grasping rfl. arly hominin mothrs must hav arrid
sial right hmisphr that pro- thir hlplss infants in thir arms' at last until ay slings wr invntd.
J 1 5o L l rU t t r a n sl n. t h i s s n s , Undr thsirumstans'soothing nratrnalvoalizations would hav n
ft hmisphr. adaptiv for rassuringnary infants tht wr, of nssity,priodially r-
dy had nothing to do with th movd from thir mothrs' mras.Ton of voi may thus hav volvd as
d as a kind of uditry frill that a kind of dismdid tnsion of mothrs' radling arms. Taking a u
566 uolutio of Lgg
|BLIoGRAPY 8 n r a t h i s v a r i t i n
this variation. Th stud
Armstrong,D. F.' !. . Stko,nd S. . Wilox. 1995. Gstured the Nture of quirs intgrating on<
Lguge.amridg:amridgUnivrsityPrss.
with an undrstanding
Falk, D. 24a,Brinde:Nt'uDiscrlurisbout Hum rigis d Bri
uolutio,Rvisdnd panddd. Gainsvill:UnivrsityPrssof Florida. A t u a r i a l s t u d i s 1
2004. Priinguistivolutionin arly hominins:Whnmothrs? n l y s i si n h u m e n s .l n
Bhuiorld Bri Sis 27:491-541. t n so f h u m n m o r t e
209.Fidig ur Tongues:others,Ifntsnd the rigis of Lngug. r i s k o f d a t h . o I o g i
Nw York: Basi Books. d a t h r a t s ,f r t l m w h i
Hoktt,, 1966.Th prlmf univrsals in langug.In J. Grnrg,
d.,
suh as th nt rpod
Uniuerslslf Lguge,1-29. amridg,MA: MIT rss.
Holdn, . 1998.No lastword on languagorigins'Si282: t455-I458. futur ptd rpo
-h 'Word: haratristis suh as
Knnally,. 27. First Th Serchfor th origis of Lgug.Nw
Y o r k :V i k i n g . varials ar dirtly rt
Laitman,J. T.' J. S. Ridnrg, and . J. Gannon.1992.Fossilskullsand hominid n r p o d u t i v r t )r
voal trats:Nw approahsto hartingth volutionof human sph.In
individuals that produr
J. !ind t al', ds.,I,ngugrigi: ultidisiplirpprlh,395_407'
Dordrht,Th Nthrlands: Kluwr admiPulishrs. th dirt Iink tw
asataka,N. 2003. he onset of Lnguge.amridg:amridgUnivrsity shaps th divsity of
Prss. | s r l s o n t r a il
l\4Brarty'S., and A. S. Brooks 2000. Th rvolutiontht wasn't:A nw ropi of importn r
intrprtationof th origin of modrnhuman havior.Jourl of um spis.
u o l u t i o 3 9 : 4 5 3 _ 5 6 3 .
o d r n l i f - h i s t o r yl
Nwman,J. D. 2004. othrsy any othrnam:othr-infantommuniation
in non-homininmammals.Behuiorlnd Brin Sics27: 51'9_520. o n t o l i n t h 1 9 5 0 s .]
.William nithologist who infus
Pinkr' S. 1994. b Lgglstint.Nw York: orrow and ompany.
Savag-Rumaugh, S.' S. G. Shankr,nd T. J. Taylor.2\.Apes,Lnguge,d H i s m o s t i n f l u n t i a l a
the Hum il. Nw York: oxford Univrsity Prss tht irds ly in a nst
maximizs th torl o
known as th ..Lak l
uolutiolt of Lif Histlris 567
parnts must alan th numr f hathlingswith thir aility to rar thm funtion annot .
to indpndn. If parnts attmpt to ar mor offspring than thy an sup- offs, and how hav
port' som or all of th young may di. Likwis, prnts prduing fwr yl influn survil
offspring than thy an rar will fldg fwr than thy ould, and will lik. th ntral qustion i
wis hav lowr rprodutiv suss than if thy had produd a largr sptivs of Lak anl
luth. lif historis that hs
primntal studis on many spisof irds, in whih luths ar rtifi- ativ today.
ially nlargd,hav found that luth sizs produd y irds ar gnrally Modern lif-histor
smallr than th Lak luth Siz (.g.,Vandr.Wrf 1992). This disrpany th nvironmnt' d
twn th Lak and th osrvd luth sizs stimulted a grat divrsity of strain lif-history val
xprimntal and thortial study that in turn gnratd many adaptiv hy- an volv rapidly w
pothss' mainly drivd from optimization analyss (.g.,..optimlity mod- dution of prdator
ls',; Orzak and Sor 1994) that am ommonpla y th 1970s and onto nw host sp
1980s. For ampl, modls prdit and xprimntal studis hv shown 2007). Seltiv prd
that urrnt yar rprodutiv ffort ffts futur rprodutiv suss of has ld to th voluti
parnts; that is, thr is a trad-off twn urrnt and futur rprodutiv tns of gnrations(I
ffort (s rviws in Godfray et a|. 1991 and Rlff 1992\. \/hn this and tion favors rprldu
othr trad-offs (suh as that btwn luth siz and gg siz; Smith and shard volutionary l
Frtwll 1'974) are inorporatd into modls of luth siz, w n plain isms, gnti varitio
muh of th srvd dviation twn rl luth sizs and Lak's pr. volution of lif-hist
ditd luth sizs. tionary biologists to
Following on Lak's introdutin of Darwinian thlnking into th study of ation, dvlopmnta
lif-historyvariation, Lamont ol (1954) introdud a mathmatialfram. so that aus-and-f
work to th study of lif.history ology y dmonstrating hlw hangs in d- Th Darwinian pr
mographi varials influn th rat of inras of a population (Roff and othr filds of
7992). ost famously, h raisd a simpl mathmatial onundrum known as n and lif span in
ol's parado: an rgnism that livs forvr, produing offspring pr flund y an organi
yar, lavs no mor dsndants' all ls ing equal, than on that livs for rprodution. In mos
just on yar (an annual organism),ut produsz + 1 offspring.Th rason is lins with ag. Thur
tht nly on offspring is rquird to rpla th parnt in our annual organ. lss than dos morta
ism, aftr whih oth th immortal and nnul organism produ th sam ar undr wakr sl
numr of offspring. Givn th impossiility of irnmlrtlity, why do not all this dlin in th int
organisms volv toward th shortst, simplst lif history? any iologists, Snsn.oupld
.Williams,
inluding Willim D. Hamilton, Gorg and ri harnov traits that fvor su
(harnov and Shffr |97), hav takld this onundrum, and its solution ost to ultimat long
ws sminal to th formalization of Darwinian thinking in modrn olgy. gns hav n idn
Th solution to ol's parado rquirs rognition that ah lif-history d- only on th old an r
ision auss a shift in th proility f aruing rtain nfits,and rtain mulation of dltrio
osts, in trms of survival and rprdution.For xampl' if rprodutivsu- N4or powrful mat
ss inrass with ag (.g., du to nfits of parntal prin) and th mor insightful pri
risk of motality twn rprodutiv priods is low, thn sltion may fa- lif.history variation l
vor rprodution for multipl sasons. In <lntrast,whn th risk of mortality in genrating nW spe
is high, seltionmay favor alloatingmor rsoursto urrnt rprodution. latd into th applid
Alloation trad-offs suh as ths onstrain how lif historis volv. R- volution influns tl
sours' inluding tim, nrgy, and ssntial nutrints' must dividd tainability of agriult
among growth, maintnan,and rprodution. Rsours alloatd to on and disasrlatsto
uolutiolt of Life Histories 569
with thir aility to r2rthm funtion annot lloatd to othr funtions. Undrstanding ths trad-
' offspringthan thy an sup. offs, nd how havioral or dvlopmntal disions at on stag of th lif
vis'parnts pr<lduirrgfw ylinflunsurvival and rprodution at othr stags(and totl fitnss),is
han thy ould, and will lik- th ntral qustion in th study of lif-history volution. Th Darwinian pr-
tlry had produd a largr sptivsof Lk nd ol ouglrt nw lvav of thinking into th study of
lif historis that has ld to myriad insights and a fild of inquiry that rs vry
ls, irrwhih luths ar artifi- ativtoday.
'odudy birds ar gnrally odrn lif-history analyssaim to undrstand how volutionary history,
Wrf 1992).This disrpany th nvironmnt, dvlopmnt, and inhritan intrat to rat and on-
stimultda grat divrsity of strain lif-history v:rriatior:r.An irnportant rnt insight is that lif histois
r gnrtdmany adaptiv hy- an volv rapidly whn nvironmnts hang (.g.,th xprimntal intro-
..optimlity mod-
ralyss(.g., dution of prdtos, olmrizll xploitation y humans, and pansion
|mmonplay th 1970s and onto nw hst spis y parasits of plants and animals) (arroll t al.
rimntlstudis lrav shown 2007). Sltiv prdatior-ror lrarvsting of lrg individuals, as in fishris,
uturrprodutiv suss of has ld to th volution of sustantially rdud ag and size at maturity ovr
rrntand futr-rrrprodutiv tnsof gnrirtions(Huthings nd Frasr 2008); this lrpns ussl-
,i Ro{{ .'991-\1.\l/h..--t. r:.d tion favors rprodution for animals ar atn r harvstd. Thus, whil
l siz and gg siz; Smith and shardvolutionary lristor},tndsto rat similritis among rltd organ-
of lutlr siz, w an xplain isms, gnti variation for lif-history trits is aundant and allows th rapid
ll luth sizs and Lak's pr- voiution of lif-history stratgis'Suh ongoing dptation allows volu-
tionary iologists to wigh th rlative influnsof phylogny' gnti vari.
nin thinking into th study of ation, dvlopmntal systms, and th nvironmnt ir-rshaping lif historis,
rodud 21mathmatil fram_ so that us-nd-fftrlationslripsom larr.
monstratinghow hangs in d- Th Darwinian persptivon lif historis an ring insights to mdiin
nrasf a population (Rff and othr filds of iology, inluding publi halth onrns suh as sns-
nmatialonundrum known as nand lif span in aging human soitis.Th volution of lif span is in-
r'r' prtlduing iz offsping p f]undv arr oganism'srprodutivvalu, th masur of futur ptd
rg qual,than on that livs fr rprodution. In most organisms on rding starts rprodutiv valu d-
s/7+l ffsprng'h ason is lins with ag. Thus mrtality lat in lif affts total rprodution muh
th parnt in our annual organ- lss than dos mortality arly in lif, so taits ffting mortlity lat in lif
ual organism produ th sam ar undr wkr sltionthan ar traits afftingmortality arly in lif. It is
f immortality, why do not all this dlin irr the irrtnsityof sltionwith ag that llows th volution of
st lif history? Many iologists, snsn. oupld with th sltivdvantag of short gnration tims'
Williars, and ri Charnov tritsthat fvo sussfulrprodution ariy in iif will fvordvn at a
his onundrum, and its solution ost to ultimate longvity (an ampl of antagonisti pliotropy); indd,
n thinking in modrn ology. gnshav n idntifid tht hav ths ffts. Traits with ngtiv ffts
nitionthat ah lif-history d- only on th old an volv in populations through gntidrift (i..,th au-
-ringrtain nfits, an] rtain multion of dltriousmutirtions).
jor
x a m p l 'i f p r o d u t i v s u - or powrful mathmatial tools, nw thnologis (.g.,gnomis), and
of pantrlxpin)and th moinsightfulxprimnti.ttion pmittingiologiststo ttrundrstand
Js is low, thn sltion may fa- lif-history variation within and mong spis and th rol of suh variation
trst, whn th isk of mortality in gnratingnw spis.Th rsulting insights ar inrasinglying trans-
:sours tO urrnt rproduti<rn. latd into tlr applid sins whr iologists ar studying how lif.history
,in how lif historis volv. R- volution influns th onsrvation of ioti rsours,th yild and th sus-
rial nutrints, must dividd tainailityof agriultur,and how th volutilnof ontogny, rprodution,
ion. Rsoursalloatd to on and disasrlats to human halth.
570 uolution of Se
lLIoGRAPY
s-slnd to hirv
aroll,s. P.' A. P. Hndry,D. N. Rznik,and . W. Fo.
2007.volutronon lonirl fmal trans
ologialtim-sals.Functiolcolog 21: 87-9.
harnov, . L., and W. A4. Shaffer. 1973.LIi-historyonsquns mle trnsmits onl1
of natural
sltion:ol's rsultrvisitd.Amri Ntrlist|oi| hlf of hr parnt
79]..-79.
ol, L. . |954. Th populatiollonsquns of lif historyp.no.". Qurtrl mal' Thus s arri
Ruicuof Biolog29: |0_137. hug thortial dis
Godfra,H. . J., L. Partridge,and P. H. Harvy.1991.luth
siz.l Ruie erl alld ..th p;
of olog d Sstmtics22: 409429,
Huthings,J. A.' nd D. J. Frasr.2008. Th naturf fishris- spurrd svral lad
and farming-
indudvolution.olulr colog 17:294-13. highligliting th imp
Lak, D. 1947.Tsignifian of lut..i,.. Ibis89: 3o9-52. lutionary iology'' (
ozak, S. H.' and . So.1994.optimlitymodlsand fits of sxuality' it rv
th tstof 1aptatlon.
meriNturlist143:361-380.
twi as fit, on avr
Roff' D' ^. |992. h Fuolutio of Lif Historis:Thor
lnlsis.Nw Yok: quird to Ovrom1
hapmnand Hall.
Smith,C. ., and S. D. Frtwll, 1974.Th optimal alan viduals an havthi
twnth srzand
numrof offspring.Ameri Nturlist 108: 499-506. in a world that hal
Vndr.!rf, . 1992.Lk,sluth siz hypothsis:An aminatiorrof ally, s an out()
th vidn
usingmta-ana|ysis. olog 73:1699_1705. -s.P.' d .\Y.F. (1) kills a lrg rl
ft o podution
dividuals wit]r rtir
vlution of s () hangs rirdially
Hr,you se,it taksall th runningyou n do, to kp tys ar killd and l
in th sanr tion for s, th ph
pla.
_h Rd world, ffting mosl
Qun,in Lwis aroll's
Throughth Looking Glss(1872) ? In rnt dd
disas (Hamilton 20
Th ida tht inf
of th many manings of th word se*' th most fundamntal mak s dvntg
rfrs to th
prodution of nw gnoms by th romination th spts of an org
of pristing gnoms.
any batria and a fw viruss ar apabl of oasional. th thorists' dsrip
limid s' ut
th-most thoroughly sxual raturesar th ukryots, Bs parsirs an
whih iniud ani-
mals' plants' and othr organisms with nuli in quikly ompard to
thir lls. Humans ar
fairly typial ukaryots in ing oligatly sul: gnotyps. on striki
th only wy to mk
nw human is to split your gnom in half and to gt tions should ourin-
anothr lrali from som
othr uman who has don th same. Dspit th ar ommon in on
ntral rl that sx plays
in organismal lif yis, iologists do not undrstand Qun hypothsishas
what its funtion is.
Th first biologist to suggst a plausil hypothsis is of hosts and para
for tlr dvantag of
sxuality was August.V7ismann(1ss9). H proposd fls, ut sintistsd
that s inrasdth
vaiability within a population and thus inrasd th to .
fftivnssof natural
sltion. This is an unusual sort of advantag; typially, Th othr phnon-r
natural sltion
promots traits that onfr an advantag to th individual ful mutation. With hi
.!(i'ismann' that ars th trait.
howvr, saw sx as a fatur of a populatin that ronrbirrant offsprin
onfrrd an
advantag to that population. .sismann'seplanation This is orriy tru if rat'
was influntiai nd
wnt largly unhallngd until th 1960s, whn Gorg individul arryig <lr
. Williams o-
jtd to its rlian on th ruttion thory
8roup sltin and insistd tht sx should hv
short-trm, individual-lvl dvantg lik ny other harmful mutations [
trait. On th onrrarv.
msur' t stilnat
uolutio of S .t71
to hav a high nough rat of harmful mutation to mak sx usful, ut th altrnativ forrs of a
rat sms to too low in Worms and flis. An important dtail is that s hangs unfold in volu
only hlps to inrase avrag fitnss if hzrrmful allls intrat synrgistially, mor thorolrghly or ar
that is, if thy bom poportinatly nror hamful as mor of thm u- dation of svlras
mulat in a gnom. A waknss of th mutation thory is that it dos not T r i n d i n s t t i s t i 1l
sm to aount for ologial pattrns in th distriution of asual linags rivd his dotoral d
as wll as th Rd Qun hypothsis ds. who was a studnt of S
\Jhna gnom rpliats, it tnds to dtriorat. utations our and work on th samplirrg<
thy tnd to harmful. Th nvironmnt also tnds to gt Wors from ya hav no ffts on th fit
to yar: your nemis gt ttr at attaking you, spially parasits and opd y Motoo Kimur
..Motoo Kimura'' nd ..
pthogns with short gnration tims that volv rapidly to adapt to your
gnotyp. Ths two prosss tnd to mak mintirining stati gnom biohmial and r-nol
oth diffiult (du to mutation) as wll as a ad ida (du tl parasits).S s p i s .O n o f t h r i t i t
sms to an adaptation to dal with on or anothr of ths prolms, or al distriution of alll
prhaps to dal with oth of thm at on. Thr has n a rnt trnd to- triutions xptd und'
ward omining th Rd Qun and mtrtational mdls and rognizing wns drivd ths
tat sx has n-rultipladvantags.vn Gorg Williams, whos qust for an aout th nattlr of mut
individual-lvl advantag of sx launhd th modrn study of its volu. to alld, not Only al
tion, hs ondd that its spis-lvladvantagsmight ruial aftr all pirial data' but also p
(Williams 1992|. Asxual spissm to b muh mor pron tl xtintion thortial advans wt
than sual spisH . n , m o s t s u v i v i l l gs p i sa r s x u a l , n d m a n y , i n - utral thory nd the
luding mammals, sm tO hv faturs that prvnt thm fom vr - allls among a st of n
oming asual. Although th prolm of sx has rntly bn rdud and gn flow.
from a mystry to a mr ontrovrsy' sintistsstill nd a good dal mor Thr of wns's otl
infrrnation to undrstnd xtly how sx hlps irrdividual oganisms to i n r p r i n g h g l r i
hav mor sussfuloffspring and how it prvnts populations from going alll vailal in a popl
trnt. pndd on th prisst
nti load lays bnath
B|LIoGRAPY th so-alld ldn's
Bll, G. 7982. b strpitf Ntur.Brkly:Univrsityof lifornia Prss. ost of so many unfit g
amilton,!. D. 2001.Nrrot,tlRldsof GeLnd.ol,2,h uolutioof Se, indd nutral. wns's
oxford: Oford UnivrsiryPrss. ntiststhought aout th
Judson,o.2.002. Dr. Tti'sSduito Il rti.Nw York: tropolitan for th fftiv popult
Books.
ar rally ontributing t
K l n d r a s h vA,. s . 2 0 0 1 .S n d U . T r e d si n G e t i s1 7 : 7 5 _ 7 7 '
as of onfusion, spt
Ridly, \'4, 1994. Th Rd Que: Sd tb uolutio of Hum Na,re. Nw
York: amillan. popultion siz, a quest
\Wismnn, A. 1889. sssuplttHerditd Kidrd BiologilProblems, his <lllgusdrivd n
Ofod:lrndonPss. and human disass,m
Williams, G. . 7992.Ntrl Sltil: Domis, Luls,d hIleges.oford: asd on th nonrando
Oxford UnivrsitvPrss. -B.B.N.
wns has a lgnda
strutor.H hs inspir
'Warrn Australia, prinlarily at
wns, J. (b. 1937) Univrsity of Pnnsylva
Although thy may sm sotri at first glan' many of th stirtistial as- tistil Soity of Austr
pts of ppulation gntis'suh as th prisdistrir'rtionof fitnssamong of th Royal Soity.
ues 57
)n to mak s usful' ut th altrnativ forms of a gn' play vital rols in dtrmining how gnti
\n important dtail is that sx hangs unfold in volution. Fw sintists hav unovrd ths vital rols
I al1lsintrat synrgistia1ly, mor thoroughly or arefully than \WarrnJ. wens, whos work is th foun-
rarmful as mor of thm au- dtion of svral aras of rsarhin volutionary population gntis.
Ltionthory is that it dos not Traind in statistial gntis at th Australian Ntional Univrsity, wns
jistribution of asual linags rivdhis dotoral dgrtn 1964 undr th suprvisin of P. A. P. Moran,
who was a studnt of Sir Ronald Fishr. wns is known primarily for his
riorat.utations our and work on th sampling distriution of nutral allls, whih ar allls that
o tndsto gt wors from yar hav no ffts on th fitnss of thir arrirs. Th nutral alll thory dvl-
opd by otoo Kimura and Tomoko Ohta (se th alphatial ntris
i you, spially parasits and ..Motoo
volv rapidly to adapt to your Kimura'' and ..Tomoko Ohta'' in this volum) wddd th study of
maintaining a stati gnom iohmial and molular polymorphisms to pattrns of variation among
lad ida (du to parasits). Sx spis.On of th ritial tsts of this thory ompard th osrvd statisti-
anothrof ths prolms, or ai distriution of alll numers and frqunis at a singl gn to th dis-
rr has n rnt trnd to- triutionsptd undr nutrality. In a srisof paprs in th arly \970s,
:ionlmodls and rognizing wns drivd ths ptd distriutions undr a varity of assumptions
aout th natur of mutations. Th wns Sampling Distriution, as it am
;.!illiams,whos qust for an
;h modrn study of its volu- to alld, not only allowd th nutral thory to vluatd against m-
rtags might ruial aftr ll pirial dt, ut also providd th foundation on whih svral additional
muhmor pron to xtintion thortial dvans wr uilt. Ths inludd various rfinmnts of th
r i s s u l ' a n d m a n y ' i n - nutral thory and th dvlopmnt of mthods for using th distriution of
at prvnt thm from vr - alllsamong a st of nighoring populations to stimatrats of migration
s has rntly n rdud and gn flow.
:istsstill nd a good dal mor Thr of wns's othr ontriutions stand out. First, h showd how
hlpsindividual organisms to intrprting th gnti load, th diffrn twn th fitnss of th st
r v n t sp o p u l a t i o n sf r o m g o i n g alllavailal in a population and th avragfitnssof all othr allls,d-
pndd on th pris statistial distriution of fitnss. Th onpt of th g-
nti load lays nath svral ontrovrsis in volutionary gntis, from
th so-lld Haldan's dilmma (how an an volving population ar th
ost of so many unfit gnotyps)to th liklihood that molular variation is
:y:Univrsityof alifornia Prss.
,d.o|.2,Tbuolutionof Se. inddnutral. wns's ritial study of th gntiload hangd th way si-
ntiststhought about thsissus.Sond,wns drivd pris prssions
retio.Nw York: tropolitan for th fftiv population siz, whih is a masur of how many individuls
ar rally ontriuting to th nt gnration. This work larifid svral ar-
tis17,.75-77 ,
as of onfusion, spially surrounding th fft of atastrophi drass in
,tolutioof m Na're. Nw
population siz, a qustion of intrestin onsrvation iology. Third, h and
'idredBiologilProblms. his ollagus drivd nW mthods for dtting assoiations twn gns
and human disass,mthods that solv long-standing statistial prolms
, Luels,d hllges.oford: asdon th nonandom sampls that mrg from mdial rords.
-B.B.N.
w n s h a s a I g n d y p u t a t i o n s s u p r l t u ra n d l a s s r o o m i n -
strutor. H has inspid gnrations of studnts throughout his arr in
Australia, primrily at LaTro Univrsity, and in th Unitd Stats at the
Univrsityof nnsylvania.H rivdth . J. Pitman dal from th Sta-
|an,many of th statistial as- tistialSoity of Australia ln 1996 and' in 1999,ewas ltd as a fllow
is distriution of fitnss among of th Royal Soity.
574 obiolog
|LIoGRAPHY
wns,!. J. 2004. tbmtilPopultiGetis.2ndd. Nw York: Springr.
.V.
wns' J., and G. R. Grant. 200|. Sttistilthodsi Bioinformfis.Nw
York: Springr. -IT
obiology
With th advnt of spa ploration, two filds of rsarh-origin.of-lif
studis nd th sarh for tratrrstrial lif-wr mldd into a singl nw
disiplin. In 1959, Nol Piz_winning miroiologist Joshua Ldrrg
oind th trm eobiolog for this nw intrdisiplinary fild, offing th
rational that on ndd to know th onditions undr whih lif,s origin
had n possil on arth in ordr to know what kind of hmial ondi-
tions to look for on othr plants. Furthrm<lr' h argud that strilization
of sparaftwas ruial for thy landd on othr plants,lst ontamina-
tion of thos worlds y arth miros or vn organi moluls from arth
rndr it impossil forvr to know with rtainty what kind f nativ or- i1
ganis or organisms had n thr originally. At first lrgly an Amrian ;.tl i,
;i:lif&
sin austh National Aronautis and Spa Administration (NASA)
I',
litr'
l-
Was th singl largstsour of funding, work in thsaras had th potntial
:t;
.s0.ar,
,iS
to b politiized th old as did all th sin rlatd to spa -
ploration. The Sovit Union was highly srtiv,for xampl, aout what
strilization standards,if any' had n applid to its intrplantarymissions
targtdfor impat or soft landing on th moon' Vnus' and ars. Similarly,
th Unitd Stats was highly srtiv aout how its prtis in strilization
was rlatd to lassifidgrm warfar rsarh.Gradually, howvr, intrna.
tional onfrns sponsord y th ommitt for Spa Rsarh
(osPAR) dvlopdintrnational sparaftstrilizationstandards,vntu-
ally inorportd into te 1967 Outr Spa Trty. Intrnationl origin-of-
lif onfrnsand a wid rang of NASA-sponsord mtingson th oigin
of lif, organi moluls in spa,Pramrian palofossils'nd othr topis
also gradually onsolidatd an intrnationalommunity of sintistsworking
on pis f th puzzl as disparat as iohmistry,gohmistry,plantary
gology, and radio astronomy. In 1968 th first oiology journal gan
..ll}':Rti
puliation (now titld rigis of Life d uolutio of tbe Biospber).In ^RI' {.R
As l,l'il-\ '
1972 exotologists foundd thir first profssional soiety, th Intrnational N0 I
:lds of rsarh-origin-of-lif
.wr mldd into a singl nw
roiologistJoshua Ldrrg
disiplinary fild' offring th
tions undr whih lif's origin
what kind of hmial ondi.
'r,h argud that strilization
1 o t h rp l n t s 'l s t o n t a m i n a -
n organimoluls from arth
. r a i n t yw h t k i n d o f n a t i v o r -
v . A t f i r s t l r g l y n A m i n
SpaAdministration (NASA)
in thsarashad th potntial
th sinrlatd to spa x-
tiv,for xampl, aout what
. . dr o i t s i n t r p l a n t a r ym i s s i o n s
on' Vnus' and Mars. Similarly,
row its xprtis in strilization
h ' G r a d u e l l .h o w v r .i n t r n a -
,mmitt for Spa Rsarh
stri1izationstandards, vntu-
Traty.Intrnational origin-of-
o n s o r dm t i n g so n t h o r i g i n
a n p l o f o s s i I sa.n d o t h r t o p i s
:ommunityof sintistsworking
:mistry,gohmistry' p1antary
first xoiology iournal gan ''|lli
uolutio of th Bilspber).|
AR|i.RAl}; '\l.tlRs lN n1Irll llY AR l lril,\.o R!is'il''- Us' ANI)
s ',il.\. :\S Nf} |l1. \Ytt.l' t} (}vRlil) wltl ']n^llRs ()R PL,R. l Is
N{) l'ss Rn.\s(}NI}l,F: 0 ql,ll(ts' tNs'\D () A llANI}, t,
;sionalsoity,th Intrnational (iR(}(1t' {'ll }i^('I.l:s ()R R.ln{)s('ls l,| oRANs,'
) L ) .o n l y w i t h t h t h n o l o g yo f
Beingsfrom outr spa hav long n th stuff of fantasy and nightmr.This
would it om possil to d- .!lls,..Th
drawing,y !7illiam R. Ligh, appardin H. G. Things That Live
o undrstand(or tst) what uni- on ars'', osmopolitgzie,Marh 1908, 35_343.It appardjust
lutionaryhistory. ..anals''
at th tim whn thrws muh popular disussionout whthrth
aadmilaoratory rsarh as on rs Wrth produtsof ral intlligns.Not how oftn suh piturs
:nts to fl on sparaft, suh as assumtht volutionis progrssivnd nonontingnt,with humanlikforms
Lsraft found no lar vidn th invitaloutomof th pross,anywhrand vrywher.
:noughdata to suggst that if lif
found dp nough low th
ays.In addition,th dvlopmnt
576 obiolog
of radio astronomy thniqus to sarh for signals from intllignt tratr. in th dirtion of iol.
rstrials (STI) was part of oiology fronr th outst, although ongrss faily quikly ut ..nith
ut off NASA funding for STI wok in 1992. B th 1980s and 1990s, nw stats(2000, 109).Il.s
astronomial thniqus wr dvlopd to sarh for plants of stars ynd arly in lif's history wl
th solar systm' with th first suh plant ing dttd in 1995. Laora- ating as oth an inforr
tory work has pusud numrous goals, inluding study of th organi om- lism.
pounds fond i metorits, turnd moon oks, and (via sparaft)
omts' and study of anint roks for fossilizd miros nd/or organi |BLIoGRAPHY
moluls that might ssoiatd with on-living organisms. This work Dik' S., and J. . Strik.2
has ld to pushing bak to 3 illion yars or mor th oldst known fossil mi- strobiolog.Nu, Br
ros from arth, narrowing th tim window fo th pross of hmial Fry, I. 2000. The mrg
uruit'l,,. Nw BruIl
volution to hav ourrd. Howvr, it has simultanously rva[dthat
Grhurm,L. 1987,Si
arth is ing bomardd y organi moluls formd in hug irmounts in Nw York: olurnr
intrstllar spa' so that th original prursors of lif on arth nd not all zn, R. 2005. Gesis:
hav bn synthsizd hr from purly inorgani starting matrials. oiol- Hnry Prss.
ogy rsarh has also spun off numrous othr important disovris: Lynn L d r gJ ., l 9 6 0 . h i
argulis's thory, now univrsally aptd, that ukyoti lls aros via 940.
.Wos's L v l o kI, . 1 9 7 9 . G i :
formation of srial symbioti olonis of pokaryoti lls; Carl dis-
Prss.
ovry of th Arhaa' a grop of atria as diffrnt from othr tria as Mllgu|is.L. |970.origin
thy ar from ukaryots; Jams Lovlok's Gaia hypothsis, whih laims Prss.
that all lif on arth ats as a oordinated homostati systm maintining i I l r 's . I 9 5 3 .A p r o d t r
onditions within th limits lif n tolrat; th thory that a hug stroid onditions.Sie|1
o p i n , A . l 9 3 8 . T h o r i g
impat at th nd of th rtaous priod was rsponsilfor th mss - 'W.s,
. 1987.Btrialt
tintion that inludd th dinosaurs; and th ..nular wintr'' hypothsis, Wolf,A. 2002. Grmsin s
pditing that a similar mass tintion ould follow a nular War of any imagination,1958-19
signifiant siz, and spurring on muh of global limat modling sin.
Th brodning of th rsarh agnda rsultd in th xoiology fild ing
rnamd astroiology in 1996.
Following th Soviet biohmist Alksandr Oparin, origin-of-lif rsearh
assumd in th 1950s that arth's primitiv atmosphr must hav n
hmially rduing' that is, it ontaind no fr oygn' arbon mostly s
mthan, and nitrogn as ammonia. Th 1953 Ury-Millr xprimnt
showd that Spontnous synthsis of amino aids and othr rlvant io-
moluls ould our rapidly undr suh onditions, vn assuming rzrndom
hmistry. By th lat 1970s, howvr, xoiology rsarhsuggstdarth's
arly atmosphr was mor likly to hav n hmially nutral (or pr-
haps vn slightly oidizing), undr whih onditions th synthsisof organi
moluls is drastially rdud. Altrnativ pathways of hmial volution
ar undr invstigation, inluding synthsis of organi polymrs using lays
as tmpltsand talysts;high-tmpratLlr, high-pssurhmistry suh as
might our at undrsa hydrothmal vnts, whr many rsarhrs now
think lif may have first originatd; and omining mtorit-drivd organ-
is to s what kinds of liflik protolls will produd. It is now widly
blivd that th hmistry tht ll frm inorgani prursors to tlr first
biopolymrs ours nonrandomly' mor rapidly, and spontanouslyundr
widr rans of onditions than was livdin th 1950s and 1960s' Biasd
obiolog 577
]nalsfrom intllignt xtratr- in th dirtion of iologially rlvant moluls' lif proaly originatd
:h outst,althoug ongrss fairly quikly ut ..nithy han nor dsign,''as sinhistorian Iris Fry
By th 1980sand 1990s, nw stts(2000, 109). Rsarh has also foused on th RNA world, period
h for plantsof stars byond arly in lif's history whn RNA istd for protins ut was apal of
ing dttdin |995. Laora- ating as oth n information-arying rnolul and a atalyst of mtao-
t i n gs t u d yo f t h o r g a n i m - lism.
t roks, and (via sparaft)
BlBLIoGRAPY
i z d m i r o s n d / o r o g a n i
-living organisms' This work Dik' S.' and J. . Strik.2004. The Liuig Uniurse:NASA d th Deuelopmentof
r th oldst known fossil mi- strobiolog.Nlv Brunswik'NJ: RutgrsUniversityPrss.
w for th pross of hmial Fry, I. 2000. Th mrgof Life ln the rth: A Historil d Scientifi
ouruitll.Nw Brtrnswik,NJ: RutgrsUnivrsityPrss.
simuitanouslyvald tht
Grham'L. |987. Sitl,Philosophd llm Bhuiori tb Souit IJnion.
s f o m di n h u g a m o n t s i n Nw York: olumbiUnivrsityPrss.
,rsof lif on arth nd not all Hazn,R. 2005. Gsis:he SitificQust flr Lif's rigins. Nw York: Josph
ri startingmatrils.xobiol- nr Prss.
:r important disovris: Lynn Ldrrg, J. 1960.oiology:Approahsto lif byondth arth.Si|32:
93400.
t h a t u k a y o t i l l s a o s v i a
Lovlok,J. 1979. Gi: Ntu l-ooh t Lif o I,rtb.London: Oford University
aryoti lls; arl Wos's dis_ Prss.
Iiffrntfom oth atri s argulis' L. 1970. rigi of ukroti elis.Nw avn' T: Yal Univrsity
aia hypothsis,whih laims Prss.
l m o s t r i s y s t m r n i n a i n i n g illr,S. 19.5.A produtiorr of minoiridsurrdrpossiblprimitivarth
h thory that a hug astroid onditins.Sie||7 : 528*529.
oparin,A. 1938.Tlrc rigi of Lfe. Nw York: Mamilln.
rs rsponsilfor th mass x- .Wos,
..nularwintr'' hypothsis, . 19 87. Btrial volution.i cro bi ol lgi l Ruit't-t
s 5 1': 221'-271'.
\(olf,A. 2002. Grms in spa:Amrin lif sintists,spapoliy, and puli
d follow a nular war of any imagination, 19 58-1964.lsis 93: 183-205. -/..s.
l a l l i m a t m o d I i n g s i n .
ld in th oiology fild ing
s78
Fisher 579
and physis at a varity of puli shools. |n 1917 h mrrid Ruth iln ulturl, d edi
Gatton Guinnss, with whom h would parnt ight hildrn. On of Fishr,s agriultural rsar
si daughtrs,Joan Fishr Bo, authord th most wll.known iography of fild's onptual f
him in 1978. lightd mainly y l
At Rothamstd xprimntal Station' twn I9I9 and 1933, Fishr good ompany in -
mad substantial ontriutions to statistisand gntis.In statistis,h in- omplishmnts did
trodud th onpt of liklihood (Fishr 1921). Th liklihood of a param- Royal Soity in 19
tr is proportional to th proaility of the data and it givs a funtion that 1955.In 1943 Fish
usually has a singl maximum valu, that is, th maximum liklihood. And ridg Univrsity; i
th nxt yar Fishr introdud a nW onption of sttistis,th aim of lg' amridg; a
whih was th rdution of data and th prolmati of whih Was th sp- lizath II. Numl
ifiation of th kind of population from whih th data am, stimation, and Aftr a luminous
distriution. Soorr thraftr, Fishr pulishd SttistiI ethods for R- of 72 onJu| 29, \|
serh Worhers (1925), in whih h artiulated mthods for th dsign and Fishr has en ds
valuation of primnts. Th 1925 dition was th first of many. In gntis, volatil tmpr. Fis
Fishr's first papr (in 1922| adumratd a mathmatial synthsisof Drwin- to his viws, drov
ian natural sltion with th ntly rdisovrdlaws of ndlian hrdity. writings hav bn
That, and othr work (.g.,on th volution of dminan), would ulminat mor fftivly or
in The Getil Theor of Ntrl Sletil (1930), on of th prinipal tion and his gnius
txts' along with thos of J. B. S. Haldan (1892-1964) and Swall Wright
(1889_1988),omplting th roniliationof Darwinism and Mendlism and BIBLIOGRAP
stalishing th fild of thortial populatin gntis and its appliation to B o x ,J . F . 1 9 7 8 .R . .
ugnis.Rmarkably,Th Gnticl Theor of Nturl Seletk was ditatd Fishr,R. A. 1921.o
y Fishr to his wif, Ruth, during th vnings.Th book was rvisd and ris- frm small sar
1922,n t
sud in 1958. In 1929 Ftsher was ltd to th Royal Soity.
diburgh42:3
Fishr lft Rothamstd to tak th Galton hair of ugnis at Univrsity 1925.Sttist
ollg, London, in 1933. Karl Parson (1857_196) was rtiring from th 1 9 3 0[ 1 9 s 8
hair' whih was thn dividd into two, so that oth Fishr and Parson's York: Dovr Pu
son, gon, ould b appointd. Fishr ontrolld th gntisstion; Par- 1935.h D.
'
son ontrolld th statistisStion.(Fishr had turnd down Parson'soffr Fishr,R. A., and F.
dil Rsr
of hif statistiianat Galton Laoratoris for th position at Rothamstd in
791'9.).Whnh took th Galton profssorshipt Univrsity Collg' h was
thoroughly mird in ontrovrsy ovr th foundtions of statistis with Karl
Fishs
Parson and his followrs. Fishr's rvolutionary work in statistisam at
around th tim that Parson's own wok was showing waknsss; un- Th word fisbes ls a
frindly omptition cata|zed th ontrovrsy, and Parson would tak ill hold. Most popl 1
flings toward Fishr, who had his own ill flings toward Parson' to his fs is usd, but thil
grav. Th ontrovrsy in sttistiswith Parson would not th only on- ths too ar fishs
trovrsy in whih Fishr was involvd. Bginning t 1929, Fishr and Amri- moving y ithr (
an physiologial gntiist Swall \/rightwould ngag in an animus-filld and/or unpaird far
ontrovrsy ovr th gnti asis of volutionary hang that would last un- gn (and rlassnl
til Fishr's deat ln 7962. gills'' gos a long ''
In 1935 Fishr's The Desig of perimezls appard and, lik SttistiI onsidrd fishs(L
ethods for Resrh Workrs' would pandd and rissud many living typs of fish
tims. Ths two works, and Fishr's Sttistil bls for Biologil, gri- of known fossil for
Fishs ''81
n 1917 h marrid Ruth iln ultrl, d dil Reserh (with Frank Yats, 1938) volutionizd
nt ighthildrn. On of Fishr's agriultural rsarh. Fishr's work in statistis was rvolutionary at th
r most Wll.known iography of fild's onptual foundations. orover, Fishr work in gntis, high-
lightd mainly Th Geticl Thor of Nturl Seltiott' would, with
twn 1979 and 1933, Fishr good ompany in Haldan and Wright, rvolutioniz iology. Fishr's a-
and gntis.In statistis, h in- omplishmntsdid not go unrognizd. ws awardd th mdal of th
) 2 l ) . T h l i k l i h t l o do f a p r m - Royal Soity in 1938, th Dwin mdal in \948, and th oply mdal in
data and it givs a funtion that 1955. In 1943 Fishr was appointed Balfour Profssor of Gntis at am-
;, th taximurn liklihod. And ridg Univrsity; in 7957 he m prsidnt of Gonvill nd Caius ol-
Lptionof statistis,th aim of lg, Camridg; and in |952 h was ratd Knight Bahlor y Qun
llmtiof whi was th sp_ lizabth II. Numrous othr honors wr stowd upon him'
:h th dta am' stimatiln,and Aftr luminous ar as statistiianand iologist, Fishr did at th g
hed Sttistil ethods for Re- of 72 on Ju| 29, \962, in Adlaid, Australia, whr h had rtird i \9 59 .
ated mthods for th dsigrr and Fishr has n dsridas harmig and Warm among frinds, ut with a
W a st h f i r s to f r a n y .I n g n t i s ' volatil tmpr. Fishr's tmpr' ombind with an unwavring ommitmnt
rathmatialsynthsis of Darwin- to his viws, drov him to hatd ontrovrsis with othr sintists. Fishr's
vrdlaws of ndlian hrdity. writings hav n dsrid as diffiult; muh of wlrat h ontriutd was
of dorrrinan).would ulminat mor fftivlyonvyd y othrs mor apal of simplifying his prsnta-
ion (19\, on of th prinipal tion and his gnius.
(1892-1964) ad Swall Wright
lLIoGRAPY
f Darrvinism and ndlism and
-b
on gntisand its appliation to Bo,J. F. 1978. R, . Fishr: Lif of Sietist.Nw Yrk: \Wily.
Fishr,R. ^. |921. on th ,.prlblrlr''f a offiintof orrlrtin
ddud
l lf Nturl Sltion was ditatd
fronr small smpl.rt<l113_32.
gs.Th ook ws rvisd and ris-
|922. on th dominanratio. Prledigs lf th Rol Soitof
th Royal Soity. d i n b u r g h4 2 : 3 2 | - 4 1 ,
n hai of trgnisat Univrsity -]'925.
Sttisticlt|lodsflr RserhW<lrkrs. L<lndon:olivr and Boyd.
857-196) was rtiring from th 1930 [1958l.h GeetilThorof Nturl Sltio.2ndd.Nw
;o that oth Fishr and Parson's York: DovrPubliarions.
trolld th gntis stion; Par- 1'935.Te Dsig rlf primertts.London: Olivr and Bod.
Fishr'R. A., and F. Yats.\938' Sttistilblesflr Biologil' griultrl,d
had turnd down Parson's offr -R..s.
edil Reserh. London: Olivr and Boyd.
for th position t Rotharnstd in
ship at Univrsity ollg' h was
oundations of statistis with Karl Fishs
ionary work in statistis am at
.k ws showing waknsss; un- Th word fishs is aout s slippry as th oganisms to whih it rfrs r to
.rsy, and Parson would tak ill hold. ost popl gnrallygt a rathr similar mntal imag whn th trm
ll flingstowrd Parsorl. to his fs is usd, ut this rrly inluds thinking of ls or vn sharks, although
,rsonwuld not th only on- thstoo r fishs.Dfining a fish as ..a wtr-dwlling' akond animal
inning in 1929,Fiser and Amri- moving by ithr (or oth) latral ody undulations or osilltions of paird
would rrgagin an animus_filld ad/orunpaird fan-lik ody appndagsalld fins and whih otains oxy-
.ionaryhngthat would last un- gn (ilnd rlassnmrous dissolvd wasts)vi spializdstuturslld
gills,' gos a long Way toward inluding all organisms that r sintifially
ts appardand, |Ike Sttistil onsiderdfishs(Hlfmrrt al. 79971'Polg t al. 2005). So divrs ar th
b xpandd rrd rissud many living typs of fishs,howvr, rht this dfinition is not lzrrut. Irrlusiorr
istil Tbles for Biologil, Agri- of known fossil forms furthr strths yond auray almlst any attmpt
'82 ishs
:ountfor th nar impossiil- nutral uoyany rlativly sir to ahiv. Although dmonstrating similar
t straightforwadnoun. sponssof th vrtrt ody to th natural sltion dmands of living in
s, alit vidntially drivd Watr' th sparat volutionary lads of th Chondrihthys and ost-
g normously divrgd in al_ ihthys hav oftn usd vry diffrnt mhanisms to ahiv a ommon
url sltion.Sond, fishes daptation' For xarpl' to ovrom th fft of gravtt on thir dnsr-
to individuals f a
r singr.rlar than-wtr odis, hondrihthys hav volvd lag' oil.rih livrs that
] th most iodivrs of living provid uoyany. Ostihthys hav modifid an axiliary spiratory out-
g) fishsnumr no lss than pokting of th sophagus into a gas-filld uoyany laddr. Suh diffr-
of al[ -tantvrtrts;thy nsdmonstrat th random natur of adaptation asd on han gnom
l arth's surfa ovrd by variation.
2olotlf t pln.ssura o- Th xtant hondrihthys r flsh atrs and gnrally larg organisms. A
nd stuinnvironmnts. typil shark is aout 2 mtrs long. vidn from analysis of th shrk
ttr'and oftn in sutrrann gnom indiats that thy hv volvd at a muh slowr rat than hav ad.
physiologial,and havioral vand Osteihthys. This may aount for th rlativly small numr of
to th lvidly vari:r1ondi- spisin istn today. Living ostihthys ar highly varial in dit and
siz, ut thy ar gnrally sallr than sharks' with som ing lss than a
plus an<lthrusually, if rlu- ntimtrin adlt maimum lngth. Thir ditary radth and small siz al-
r. A dzn' prhatrlsonsider- low thm to thriv in many mor haitats than any othr lad of fishs, sp-
lwn lnl frorrrfossils' som of ially in fshwatr nvironmnts. Forty.on prnt of all fish spis liv in
l ony fragmntsfrom th or- frshwatrhaitats, whih ar highly dividd and isolatd on from anothr'
in roks from th arly am- whil 58% of living spis our in th sa, whih (in spit of its normous
living lingsar th fishlik tnt) is mor ontinuous hitat than th piolrtinntl fshwtrs. In-
rt ut akonlsshagfislrs dd som frshwatr ony fishs show astoundingly high rats of voltion.
:s; th jawlss and truly vrt- Th ihlids of South Amria, Afria, and Suthrn India ar spially sp-
, l u t4 0 l i v i n g s p i s ;t h e t i - ios, with litrally hundrds of spis having volvd in situ in ah of th
h y s )w, ith at last 880 living svralgologially young Afrian Rift Vally laks. Baus thy hav xtn-
)xtant spis, th ony {isl.rs siv parrrtalar of ggs and young' ihlid offspring an om haviorally
numr of living speis som. imprintd on thir Parnts nd show a prfrn for individals similar to
rrrdlmprs shr a ommon thir parnts in hoosing thir own mats (assotativ mating), rsulting in
t thy lk jaws and paird fins. rapid grrtiisolation and sltion lading to nw spis.
tthloatdaound thir oral Th random pattrn of volution without dirtion or lar sltivvalus
n th midlins of thi l_lik is also aundantly dmonstratd y fishs. Almost ll ostihthys long to
rondrilrthysand Ostihtl-rys th vast group known as Tlosts, inluding ls, hrrings, salmons, minnows,
l m d y h r d r . t h n - o nm
in- atfishs,lntrn and othr dp-safishs,ods, anglrfishs,basss,phs,
Litnornrous thr-dimnsional almost ll oral rf fishs' and so on. Thr ar 25,000 or mor spis.
ls of oth groups slrows similar Tlostsfirst appad r.rt200 million yars ago in th mid-Tiassi. Thir
two ladsto th natural sl- uniqu uniting fatur is struturally rthr mino shift in th intrnal sup-
tto mov, to fd, and to avoid port of a portion of th tail (audal) fin that dos nor appar to giv thm any
: to assumvrying angls of at- lar adaptiv advantag ov othr ontmporary fishs. Nvrthlss, this
rovr th fishs' odis. Jaws on lin:rghas n th gnsisof most fishsliving today, illustrating th ran-
m th st of th lrad sklton dom and han natur of volution.
s, and tlr lral-ranhialavity
xpndd to rat sution
B|LIoGRAPHY
nto th mouth. As fins and hy-
:d agilitv, havy trnal armor FishBas:A Gloal In{ormationSystmon Fishs.http://www.fishas.org/hom
o l < l st I t t l g t h rr,n k i n g ,n a .htm.
584 Fith
al to idntify spifi variabl positions in th influnz squn that 1.96, a post h hll
ould b usd to hlp prdit thos strains that will aus futur pidmis. of All Souls ollgt
This old departur from th luidation of past rvnts)th status quo for al- ivd its Darwin Nz
most ll of volutionary iology, to th prdition of futur volr.ttionrytra- Ford am to th
itoris stands s a tstmnt to forward thinking arrd prsin1that hs st in th Lpidpt
makd l/altrFith as a trulv xDtionalmolular volutiollisr' thr of a populatio:
Their prsonal s
BlLIoGRAPHY ality rords going
Blin,I. 1953.Th tlghogd tb o.Londn:Widnfldnd Niolson. th papr pulishd
'W. aility of th popult
Bush,R. .' . A. Bndr,K. Suarao,N. J. o, llr-rd . Fith' 1999.
Prditinghvolutionof hunrr-r influnza. Sin286: 792|-1925, ltion, whih allolr
Fith, W. \4. 197.Distinguishinghomologousfrom analgousprotins.Sstmti h maintaind his i
Z o o l o g 1 9 : 9 9 - 1 1 3 . on ths animals ir
1'971,a. Rate f hangof onomitantlyvariblodons.lourI of
stsllrfor th sr
olulr uoltiolt |: 84-96.
1971.Toward dfiningth oursof volrrtion:inimun-r:lrrng for a Ford's earlistw
spifitrtopology.Sstmti Zoolog 2:406416' ly of a gn that af.
2000. Homology:A prsonalviw on somof th prolms.Trdsi G- cheureui. Thy sh
tis16 227_21. gn' but also n t}
Fith,W. M., R. M. Buslr.. A. Bndr,and N. J. Co. |997.Long trnrtrndsin
growth. Although I
th volutionof () HA1 humanirrflunza typA. Prldigs o|.tb
Ntionl Adryof SieesL]S 94: 7772,-7718. gns' te Gmmr
Fith,W. \4., and . Margoliash. 1967.nstrutionof phylognti trs.Sie Ford is prhaps
\55 279-284. was th first to off
Fith, !7. ., and . N1rkwitz.1970.An improvdmthodfor dtern-rining odon pard for Julian Ht
variabilityin gtrnd its appli:rtion
to th rtof fiationof mut:rtinsin
r ' o l u t i o n .B i o h t i , . dG t i s4 : > 7 9 - 5 9 3 . -r.../. Ford's ologiI
rn toBthr in th
spis in suh pro1
rurrnt mutatlon
Ford, dmund Briso (1901-1988) whih on morph i
. B. Ford, usually alld Hny y lris olla.usnd frinds, is st known and polymorphis
as th hif proponnt of th rsarhprogram known as ologial gntis. nti polymorphisr
Intndd to throw light on th rol of gnti divrsity in th adaptation of among loi ontrit
..su
organisms to thir nvironmnts, eologial gntis gins with osrva. what h alld
tions of variation in ntural populations. Brding studisthn dtrminth phnotyps' Batsi
mannr of inhritan of th diffrnt varitions. Cmparativ analyss of ampls of suprgn
diffrnt haitats and orrlations of nvironmntal and gti varials Although not gi
lad to hypothssout th rol of variation in volutionary adaptation. Fi- R. A. Fishr in th ]
nally' primntal manipulation of nvironmntal varials rsults in th th Sarlt igr m
onfirmation or rjtion of th hypothss.Ford's rsarhphilosophy is st Ford was al to tr
out in lris sminal ook, ol-lgilGentis. yarly nsuss.Fi
Ford spnt his rrtiresintifi arr at Oford Univrsity' H matriu- quny wr too g1
latd as an undrgraduatin 192, reeivinga BA in zology i |924 and a ontntion at tht
BS in \927. tle ws appointd Dpartmntal Dmonstrator in Zoo|og of fftivpopula
(1927|, thn Univrsity Dmonstrator (1929) and Lturr at Univrsity ol. that th original an
lg (1933). I 199, h ar Radr in Gntis, th first xpliit p- Ford was also in
pointmnt in that sujtin oford. H stalishdth Gnti l-aoratoris populations. For
in 1951 and rivd an appointmnt as Profssor of ologial Gntis in th hindwings of ,V
Ford 587
in th influnza squn that 196, a post h hld until his rtirrntin \969. }-ord was lrda Fllow
rt will aus futltr pidmis. of All Souls ollg in 1958. H was a ilow of th Royl Soity nd r-
ast vnts' th status quo for al- eivdits Darwin dal tn 1954.
tion o{ futur volutionary tra- Ford am to th study of gntis and volution through his liflong intr-
rinking and prsin that has st in th Lpidoptra. Bfor attnding Oford, h gan a study with his fa-
rolulr volutinist. thr of a population of lit (phdrs) urii, th Marsh Fitillary.
Thi prsonal osrvationsmrad th yrs from 1917 to 1935, and lo-
lity rords going k to 1881 mke this an unusually lngthy study. In
th papr pulislrdwith his fathr in 1930, Ford attriutdth unusual vari-
on: Widnfldnd Niolson.
.W. ability of th population during priods of aundan to th rlaxation of s-
'x, and . Fith. 1999.
\. Sic286: 1921-7925. ltion,whih allowd suoptimal phnotypsto surviv.Throughout his lif
.om anlogusprotins. Sstmti h maintaind his irrtrstin moths and ttrflis,prrlishingtwo volums
on ths animals in oliins's Nw Naturalist sris. Ilts Bltttrfljes ws a
v:rriblodons.Jourl of
stsllrfor th sris.
Ford's arlistwork at Oford was a ollaorativ study with Julian Hu-
lution:inimum hang for a
):406416. ly of a gen that afftd th rat of darkning of tlr y olor in Gmmrlls
.reds
m of th prolms. in Ge- chureui. Thy showd that th irt dpnds not only on th Mndlian
gn,[.lutlso on th siz of th y, th tmpratur'and th rat of odily
| . o . 1 9 9 7 . L o n g t r m t r n d si n growth. Although not th first to ttiut volr'rtionaryimportan to rat
t,vpA. Prodigs lf th
gns'th Gmmrus studisprovidd an important arly ampl.
7.-7718.
:tirrof phylognti tres. Sien Ford is prhaps st known fr his wok on gnti polymorphism. H
Was th first to offer a ritial dfinition of th phnomnon in a papr pr-
vd r-rrhod for dtrmining odon pad for Julian Huxl,s The Ntu S1'5ngmtis(1940), rfind as follows in
rttlf filrion tlf rnut.rtionsin Ford,ologil Gtis (|964,84). ..Gntipolymorphisrrris th our-
-r.K.l.
rn togthr in th sam loality of two or mor disontinuous forms of a
spisin suh proportions that th rarst of thr annot maintaind y
rurrnt mtation.'' H distinguishd twn transint polymorphism, in
whih on morp is in th pross of rplaing anothr, and stal or al-
)a8usand frinds, is st known nd polymorphism, maintaind by stailizing sltion.H notd thL1tg-
21In known as ologial gntis. nti polymorphism oftn irrvolvd larg phnotypi ffts, los linkag
lti divrsity in th adaptation of among loi ontriuting to thos ffts, and strong sltion, lading to
..suprgns.''
al gntisgins with osrva- what h alld H intrprtdth ABo lood grous' Rhsus
dingstudisthn dtrmin th phnotyps, Batsian mimiry in uttrflis, and htrostyly in plants as x-
iations.omparativ analyss of ampls of suprgns'a position orn out y susquntrsarh.
rnmntal and gnti varials Although not givn to mathmatial nrodling, Ford ollortd with
ln in volutionaryadaptation. Fi- R. A. Fishr tn th |947 study of olor polymorphismin Pi dlmiul,
)nmntal varials rsults in th th Sarlt Tigr moth. Sin htrozygots at this lous ar distinguishal,
. Ford's rsarhphilosophy is st Ford was al to trak gn frquny and popultion siz in a long srisof
yarly nsuss.Fishr,s analysis stalishd that th hangs in gn fr.
o f o r d U n i v r s i t 1. H m t r i u - quny wr too gt to rought aout y gr.rti drift lon, a point of
rg a BA in zoology i |924 and a ontntion at that tim. Although hallngd y Swall \right on th asis
rntl Dmonstrator in Zoo|og of fftivpopulation sizs,susquentwork on this olony has stalishd
9) and Lturr at Univrsity ol- t h t t h o r i g i n a l a n a l y s i sW s o r t .
in Gntis, th first pliit ap. Ford was also intrstdin th fftsof polygns on variation in natural
alishdth Gnti LaL.loratoris populations' For many yars h studid variation in th pattns of spots on
rofssorof oloial Gntis in th hindwings of ill ;jrti,th Madow Brown uttrfly. Tlr pattrns
.'88 Flrm and Funtilr
wr mark:rblyonstant ovr wid aras of urop, ut thy sho,dgrat thn th fr-rrrtionstht '
hzrngsovr slight distarrsin sor.rthwstngland arrd th Isls of Sillv' of ftlrm. Russll rfrr
Spot pattrns on th lrg islands tndd tl similar, whil tlros from T h p r t l - { u n i r lpno s i t i r
smallr islands diffrd rdially. Ford attriutd tl"ronstany lf pattrn r i o r o h | sD e r w i r l
ovr tim to stailizing sltion and th diffrn btwn larg .rnd small natural tlrologians.Aft
islnds to th fft of avrirgingovr divrs haitts orr th larg islands. implrtanof naturl st
In addition to his gnti wok' Ford mad orrtriutionsirr svral othr in th i830 dat t
filds. pulishd on th hmistry of uttrfly pigllrnts'on th arhaol- Hilair (olman 1964
ogy of southwst nglnd' and on th ontnts of hurhs in th Oford forms of nimal odis
distrit. h d t b s r v d ,t h i r
Prhaps Forl'sgrastirrrilton th str'rdyof vari:lti<lnin natrrr:llpopula_ tlrat animirls slrardbst
tions has <.lmaout tlrrough his intations 'uvithhis studr-rts and ol- D a r w i l r r p t l r r dO l l t l ) i
lagus.H influndth work of Philip M. Shppard,yil A. lark, A..}. on th siJ of |.untion
ain, H. B. D. Kttlwll, Laurn M. Cook, Linolrr P. Browr, John R. G. funtion-was th aus(
Turnr' Ronilld A. Fishr, and many othrs.Ford ws prhaps too inlind to
It is gnrall1,akn<lu
it only his frinds and ollgus,ut, s Brya l:rrk (199.5,l62) maks
two gat laws: Unitl.
lar in his Royal Socit Biogrpbicl moir, ..still ologil Gee-
of typ is rnant that
tis . . . is nssary far for srious wlrkrs in th fild, still it is th lrst
in orgar-riings of t
and last ompromising sttmntof th .sltionist'viw, and still it is a joy
thir haits of lif. ol
to rad.''
d s n t . h p * s
y th illustrious ul,
|BLIoGRAPHY
ltion.For ntural s
lark,B. . l995. dmundBrisoFord. Bilgrphil tlirsof Felkuslfth parts of ah ing t<
RrylSoitl,of l'odl41z147_168,
having irdgltdthm
rd,. R., . B. Ford, lrndK. G. ]vlWhirter.1964'volutionrystrrdis on
illiurtl:Th Islsof Silly,1958_59.Hrdit1'19:471_488. ing aidd in sotn .
F . i s h r , R . A . , a n d . B . F o r1d9. 4 7 . T s p a d oaf g r r i l n t u oln d i t i o n s i n a th dirt tion tlf th
olony of th mth Plidomiul L-.Hrdit 1: |4_174, suitd t th sv
Fod,. B. 194.5.Buttr|lis, London:llrs. onditilns tlf ist
1964 ||9751.clllgit..l caetis.4th1.Londn:haprnirnnd ll. inlrritan of forlll
-]..
1 8 - 5 92, 0 6 )
Latr-nintnth-ntu
Forrn nd Function (dward s. Russll)
doutd th powr rlf r
dward S' Rr"rssll's For d Ft.mtilis uniqr.rlr,alulrsollr in th iormal, o somtilllsL
history of r-rimlmorphology. It is a livIy xploratin of th most imporrant o d i so f a n i m | s( s
morphologil thlristsfom th nintnththrough th arly twntith n- Form tld Futirl t
turis. How ould a rport on long-dad mophologists livly? Th livli. tts and rpr:oduti<
llss oms frorr-rth fat thr Russll organizsth thoristsaround ntral tratiss.((il1.lis of Rr
dt thi1tprsistdthroughout th nilrtnthlltuy and prsistsvn to- rrintnth ntury lrlrr.
day. Th dat is in th titl. Form nd Futio ould just as wll hv nals y lirtr historians
n titld orm ursus Futio. To put th mattr simply, whih am first, t n t i O ne t i t s p u l i . r
form or funtion? Why do odis of anirrralshav th ftlrms thy d<l?Do th morph<lllgists nd sorn
forms ist bzrusthy allo''v funtions to svl?[f s, frrntior-rlisnr
is sond itlr-ry,ws tht
favord arrd form is a v.prldut of funtion. or ar organismi forms or d u r i n g t l l r l yt w n t i
strutursa produt lf autonomlus fom-grlrtingprlsss?If th lattr, (sth main ssay ..Th
tlm and Futrtin .t89
Lrpr'ut thy showd grat thn tlr funtions that ody arts srv ar th y.produts' llot th auss
..pu morphology.''
of frr. Rrrssll rfrrd to th prl-f<rrmpsitin s
,l.rndnd tlr Isls of Silly.
, similar, whil thos from Th pro-funtion plsitiontook various fornrs during th nintnthntury.
d th onsteny of pattrn Prior to hals Darwin, funrilnalistsWrtlologists,most of whom wr
n twnlarg Lrnd small ntural tlroltrgians.Aftr Dilrwin' most furtionlistsWr livsin th
itatson th larg islands. importanof naturl sltionor Lamarkism. Th ontrast was illustratd
ontriutilnsin svral othr in th 1830 dat twn Gorgs uvir irnd tinn Goffroy Saint-
|y igmnts,on th arhol- H i l a i r ( o l m a n l 9 6 4 ; A p p l 1 9 8 7 ; R u s 2 0 0 3 ) . u v i r l r i m dt h a t t h
s f hurhs in th oxford forms of animal odis and body parts wr ditatd y th funtions tht
had to srvd, tlrir so.alld ondirions of istrr.Goffroy laimd
f variationin naturl popula- that animals shard astrt ldy plr-rs, and lr insistd on th unity of typ.
s with his studnts and ol- Darwin rportd on this dat in o th rigi of Spis,ut am down
p p a r d , r ' ' r i l A . l a k , A . J . on th sid oi furrtion.Naturl sltion-a pr()SStht shps form ttl suit
,inolrrP. Browr, Jt>hnR. G. funtiln-was th aus of volutionary hng.
d was prhapstoo irrlind to It is gn|l,vakn<lrvldgltlrat all organi Lringshr,n formd on
a r r l r k ( I 9 9 5 , 1 6 2 ) m a k s tw() grat laws: Urrity of Tp, and th onditions of istn.By unity
,,stt||cologil Ge-
ir, of typ is rnt that fundarnntlirgrmntin strutur,wlrih W s
th hld, still it is th last in lrgani ings of tlr stll lass, nd rvl.rihis quit indprrdntof
i o r l i s rv. i w , n d s t i l l i t i s j o thir haits of lif. On my thlry,trnity of typ is plaind y unity of
dsnt.Th prssilnof onditilrrslf istn'so oftn insistd on
th illtrstriousuvir, is fully rrrrady th prinipl of natural s-
ltion. For natural sltion ats y ith now adapting th varying
l Mtirs lf FIlo's of th pats of alr ing ro its lrgni and inorgr-rionditions of lif; or y
hving adptd tm durirrg long-pst pildsof tim: th adaptations
4. volutionarystudis on eing aidd in som ass y us and disus, ing slightly afftd y
Irtlit1,1'9 47 |-488.
th dirt ation lf tlr trtrlonditios o lif, nd ig irr all ss
g r l l i n n t t l l o t l t l i i t l n isl l
|it1,1: 11_174' sujtdto th svral larvs of gIOWth. n, in fat, th law of th
nditiolls of istn is th higlrr law; as it inluds, tlrrough rh
hrllan nd il.
-onr.1tln: inhritalr tlf fonlr 1ptatiolls,r:rt rf Urriry of Tvp. (Darwin
-1.til.
1859,206)
59
594 The Gldpgos rhipelgo
mokingird that h knw am from diffrnt islnds and that smddif- David Lak, who throu
frnt physially. thought als<raout th tortoiss tht th arhiplago's Ga16pagosjust ftlr\x
govrnor had dsridas distintivto thir rsptivislands.As h trid to siv olltion of th h
put in ord th nrany small linhs tht h had lltd, h saw that thy Amrirr usunr oi N
too rvalddiffrnson diffrnt islands. ould thsall rnrvaritis Lk at first put .rll sig
or Wr thy somthing mor, in fat and in thory? War yars on his {inding
Whn Darwin turnd to ngland' h gav his spirnns to spialists to was no dout mulr infl
atalog. At this point Darwin flt oligd to slip ovr tO n volutionary po_ onlmist rnst a.vr. I,:
sition. Th irds partiularly wr undoutdly diffrnt spis, and this thal
prisly at th tir-l-r
ould plaind naturally only on tlr supposition tht original foundrs th synhsis Sstmli
hd om to th arhiplago and hangd and volvd as thy and thir su- D r i's F ihs (1947 )
ssorsmovd from islnd ro island. A ruial part of this thinking was tht rlught on y strrrggl
volutionav hang involvs division of populations and susqtrntspi- Lak's ook has now gi.
tion. Darwin thrfor always thought of volution's history as ing funda- linkd Dawin with th
mntally on of splitting and divrging. Its pattrn was oral-lik (his first nat fft of making
mtaphor) o a tr of lif (th mtaphor tht h mad f:rmousin O the ori- md Drwin an r,ol
gin of Spcis).Hr lr diffrd fro arlir volutionists wo thought of mokingirds (sSLrllt
volrrtion as ssntiallya mov up a singl lin, with variations along minor Aftr Lk,th Gli1
sid branhs. i n t h n o - D a r w i n i a ns y
Although th Gal6pagos took somwhat of a sondary rol as his think- e t n i l t U r o f t h ' l t
ing nraturd, Darwin always usd islarrd iogogaphy as a ky support of lorving British thinkinE
volutionism, frquntly irknowldging th importan of his visit to th fator. In Amria, in lir
Gal6pagos. Praly in part aus of this signifian,th grat Swiss- tur lf organisms' thr
Amrian ihthyologist Louis Agassiz, th most important post-orlgin sin- olog,vof th islnds
tist wlro nvr irptdany forrr-rof volutionism, visitd th Galipagos lat was plord and strtlr
in lif in 1B7Z' Peraps unsurprisingly, h flt that is spial rationist vi. (196I), who ma.la lon
sion of lif's history was vindiatd y th Gal6pagos and its inhaitants. xistnof altrnativ
Agssiz thought thr was insuffiinttirn to produ so varid display on Th British Wr unll
islnds gologially so young (sLarson 2001, spiallyhaptr 4). ow- until th 1970s' Th lrr
vr' no on ally followd in his path' Soon th Gal:ipagosArhiplago was t o g t h rw i t h e s t l s
stalishdas a priz as for volutio, although, ln lin with th gnral otnon of th lss
post-rigi disrditing of natural sltion, most ,ould hav agrd with 199]r:B. R. Grant and I
Agassiz that sonr othr mlranism was nddto prodr-rth aninlals and 2 0 0 7 ; W i n r 1 9 9 4 ) .V
plants as thy ar found today. ringd and watlrd anr
Suh ws th stat of affirs through th 1930s, wl,rnDarwinian sltion tions nd physial han
was mlded with Mndlian gntis and modrn volutionary thory. No- fts rought on b dro
Darwinism, or th synthti thory of volution, was orn. Howvr, thr Works onsantly' nl l
was no an imnrdiat rush to ring th Gal6pagos undr th nt of naturl s- nt. orovr' tllr s
ltion. Thr was a gnral fling, spially among Amrians' that small availailitis ar lr sign
differns twn groups r rarly adaptiv. Th population gntiist S- omptition twndi
wall Wright (19\, 792) gavethortialaking to this prjudiy arguing Although th Glipa1
that mny diffrns ar du to so-alld gnti drift, th randolrnssthat is arl1,samn whl took
.!7
rought y th vagaris of rding. Tlr diffrns twn Glipagos or- II who uilt nn airl
ganisms wr oftn put down to just suh a aus.No on dnid that sltion is now ralization y s
was important ovrall, ut not in th kinls of dtails rvald y th islands. pOrtnt natural phrron
'Ih
ky figr'rrin hanging tlris opinion WS an nglish shooltahr, forts ar ing md ttl
Tb Gldpgos Arhiplgo t9t
:nt islndsand that smd dif- David Lk, who through th notrrgnrtrt rfJulian Huly lvnt to th
Gal5pagos jrrstfoWorld lWar II to stlld,Ytlr irds. H thn gan ir nras_
tortoissthat th arhiplago's
rsptivislands.As h trid to siv olltion of th irds-finhs partiularly-that was ownd y tlr
had olltd,h saw that thy Amrian usutl of Natural History in Nw York. \riting up his findings,
] o u l d r h s a l l m r v i t i s Lak at first put ll signifint hng down to drift. Rflting during th
,hory? Wr yars on his findings, Lak swung ntirly to a sltivxplanation. H
his spir-rrns to spialists to Wasno dout muh irrflundby th Grman-orn, Amrian-rsiding t.
slip ov to an volutionary po- onomist rnst ar. Lak roomd with ayr and his family in Nw Yok,
dly diffrnt spis' and this prislyt th tirrltlrrrtlayr ws finishing his pro-sltionontriution to
lposition that original founds th synthsis Sstttt"ttisd tle rigi of Speies (1942). Lk irrgui in
d volvdas rhy and thir su- Dri's Fitlbs\1947) that all diffllslrtwnth spisar ad:rptiv,
al prt of this thinking was that rought on y struggl twn groups for sours.Although th titl of
u l a t i o n sl r n d s u s q u n rs p i a - Lak's ook has now givn th popular nn] to th finhs and has publily
llution's history as ing funda- linkd Darwin with th island inhaitants,it also had th somwhat unfortu.
pttrn was oral-lik (his first nat fft of making popl think that it ws th finhs lusivly tht
h mad famous in o tbe ori- mad Darwin an volutionist. At lst, if n()t mor important, Wr th
:r volutilnistswho thought of mokingirds (sSrrllowa 1982,).
n , w i t h v r i a t i o n s l o n g m i n o r Aftr Lak, th Glzipilgosirds Wr on oi tlr prim pis of vidn
in th no-Darwinirrsynthsis.Howvr, ontrovrsyorrtinudovr th -
lf a sondarrol as his think- at ntur of t sltivfor that lrad otlght on diffns.Lak, fol.
rgogphyas a ky support of lowing British tir-rking,had mad omptition twn groups th k1'
: impotan of his visit to th fator.In Amri, in lin with gnalthinkig in that ountry aout th rr.
is signifian, th grt Swiss- tur of organisrs,thr was mor nthusiasl for a position that mad tlr
ost impotantpost-orryi sin- ology f th islnds mor signifiant.This viw of sltion and its ffts
lnism,visitd th Gal6pagos lat was plord and strongly dfndd y alifornia iologist Rort Bowman
: l r t h t h i s s p i l r a t i o n i s tv i - (1967),whl mad long visit to th islnds in 19.52_195.H found that th
Gal5pagos and its inhabitants. istnof ltrnirtivfold supplis Was th ky firtor in sltivhng.
o produso varid a display on Th British wr ltt-tit-nglrssd,
and tlrus mrtrs rrrrindat an impss
) 0 1 ' s p i a l l y h a p t r 4 ) . H o w - Llntiith \970s. Tlr htls:rn]-and-wif tm f Ptr and Rosmary Grant'
r th GalripagosArhiplago waS togthrrvith ir sussionof oworks ilnd studnts,gan what lras sirr
though, in lin with th gnral omon of th lassi long.trm studis of tlatural sltion(Grant l986,
l' most would hav agrd with 1997;B. R. Grnt nd P. R. Grarrt1989;P. R' Grant and B. R. Grant 1995,
ldd to produ th animals and 2007; !inr 7994)..sorkingon th islt D;lhn Major (s figur), th1'
ringd and wathd and rordd vry ird, noting th numr of flutua.
L930s,whn Darwinian sltion tions and physial lrangsfronr on yar tO th nxt' spiallynoting th f_
odrn volutionary thory. No- ftsrought on y drought and y rord rinfzrll.Thy found that seltion
ltion' was orrr. Howvr' thr works onstntly' irrrltlrat hngs an our rapidly fror on yar tO th
>agosundr th rrt of ntural s- nxt. orovr) thr stllsto truth in th lz.rirthzrtfood supplis and
lly among Amrians, that small vailabilitisirr a signifiant fatol. in sltir,hang, ut not luding
v. Tlr population gntiist s- omptitioll tlvndiffrnt groups.
k i n gt o t h i s p r j u d i y a r g u i n g Although tlr Glipagos has n attrd,vhuman oupany' from th
ntidri{t,t randomnss that is arly samn wlro took th tortoiss,through th Amrin fors in \orld
ffrnsrwnGaldpagos or- War II who uilt alr airas thr' to th prsnt uadorian fishrmn,thr
tus.No on dnid that sltion is now ralization y sintists and govrnmntl authoritis of what an im-
f dtilsvzrldy th islands. portant natural phnomnon th Gal6pagos Arhiplago rprsnts.Srious f_
L was an nglish shooltahr, forts ar ing md to prott and onsrv th islnds and thir inhitants.
596 h Glipglsr..hipelgl
lvill, . 1987.Th
Pros Pis,18.J
of Hrnluil
ds.vanston,II,:
S u l l o w a yF, . J . 1 9 8 2 .
th Histor of Bkl
Winr,J. 1994.ThF
York: Knopf.
Wright, S. 191.volu
1 9 2 . h r l
vlutiorl.Pro
Galton, Franis
Franis Galton was
to pursu a rr ir
study mathmatis'
Darwin and Galton
Th islt Daphn jor in th Glipagls Arhiplago,whr Ptr and win (1731-1802), a
Rsmry Grant did thir dirds-longstudy of Drwin's finlrs.Th would a ruil
volni origins oi th islt, hill with th top lown off anl ratr formd fat, it was Galton's
within, ar larly visibl.It has n possilto work out rlationshrps
th ida f improvin
twntlr dniznsof th islnds from th squntialordr in whih th
h oind th trm e
islands wr formd, y volrriativity' s th rth'spltsslippd aound
its surf. Sevrl ,Yarsaft
Galton organizd an
uropan ploratior
latituds,longituds
Now thy ar on of th most popular of dstinations for tourists. Darwin's ation of numriala
visit to th island, rathr than lvill's, has provn to hav th longr- him. H pulishdhi
lasting fft. in 1852 and was awa
BIBLIOGRAPHY aus of his pris,
ook aout his jlur
Bowman, R. 1961. orpholrlgiI Diffrtitio d dpttil i th Gldpgos
book for amatur an
Fihs. Brkly:Univrsitr,of lifor-riPrss'
Grant, B. R., and . R. Grant. 1989. uolutionr D,mics of Nturl Popul- triumph. It wnt thr
ti.l:,hLrg tus Fih of tb Glipgos, hiago: Univrsity of hiago Phonix Prss rissu
Prss. Galton Was an ativ
Grant, P. R. 1986. lklg d utllutilof Drtui's Fihs' Printn, NJ: yars. FIis ommnt
Pritrton Univrsity rss.
mattrs (.g.,latitud
199l. Ntural sltionnd Darwin's finhes.Sitifimri, Otor:
82-87.
intrstd in mtor
Grnt, P. R., nd B. R. Grnt' 1995. Prditing mirvlutionar rsponssto hartristiof a hi1
dirtional sltionon hitalvriation. u oluti on 49 : 241 -25 1. Th part of Galtor
27, Hl d Wh Spis ltipl: Th Rditil of Drtui's Fichs. aftr h had rad Dal
Printon' NJ: Prirrton Univrsity Prss. and ultivatl plants
Lak, D. |947. Dru,'itis Fihs.amridg: anrridg UrrivrsityPrss.
able to Galton that t
Larson, . J. 2001. uoltirl's Workshp: od d Sin ln t Glipgos
Islds. Nw York: Basi Books. to dmonstrt that .
ayr, . |942, Sstttttisd th rigi of Spis. Nw Ytlrk: olumr hritd. H attmpt
Univsitv Prss. Talnt and hat
Glton 597
gzine (1865)' and in th ook that followd, Hrditr Gicts(1869). In th Unitd Stats,p
In both th artil and th ook Galton trid to dmonstrat that wht h r- involuntary strilizati
.h
frrd t as ..tlntand haratr'' wr inhritd. ok trad pdigrs dfiint or
..fblmi
of judgs, statsmn'and othrs, Galton's asi assumption was that if h followd with intrsl
.$hn
pikd an minnt judg, for xampl, tht judg'slosstmal rlativs(.8., th Nazis am
fathr nd son) had a gratrhn f birrgdistinguishdthan thos furthr that rsultd in th str
.World .!Var
rmovd (.g.,grndfathr' grandson). !7omn wr not inludd, proaly II, ugni
.!hil
rfltingoth Vitorian prjudi and ml dominan in Vitorian soity. urop. ugni
Galton onludd tht his rsults suppotd his hypothsis.Although othrs rditd with implrtal
pointd out that nvironmnt (.g.,th fathr'sgood position might nsura printing, mtorolgy,
plum jo for his son) <luldalso ndrli th orlation' Galton attmptd
|LIoGRAPY
t o d i s m i s sr h i s o n t n t i o n .
Familiar with th ll urv y th tim h wrote Hreditr Geius, Ga1- Brooks, NL.24.tr
ton alr.rltd hypothtial normal distriution for th estimatd 15 million Glto. Nw Yok:
Bulmr' |vl.20o3.Frn
mais in th Unitd Kingdom aording to thir natural ailitis.H also was
J o h n sH o p k i n sU n r v
anious to ollt rea| data that h ould analyz statistially, partiularly Forrst,D.W. |974. Frt
with rgard to inhritan.Baus this was not asy to do with human - York:Tp[ingrPuh
ings, h didd to us swt pas as a modl bausthey wre asy to grow Galton,F. 1852.Rnt
and supposdly slf-frtilizing. In his primnts h found that sd diamtr Jourl of th Ro
was normally distriutd, ut th sd diamtr fonr rogny of lag.sdd | 8 5 3 .T r p i Sl
|855 . b rt o1
and small-sddplants driftd toward th man f th population as a whol.
outris.London:
H alld this ..rgrssionto th man,'' a statistial proprty that has n r- 1865.rdita
patdly dmonstratd sir-r(.g., in th as of diffrnt lirsss of mutual 1'8_327.
funds, suh as ons spializing in growth vrsus intrntionl stoks). Galton 1869. Herditr
also disovrd that whn h graphd th diamtrs of parntal sds vrsus l889. NturlIr
21.Th rt
progny sds,th points fll on a stright lin. had drawrr th first rgrs-
outris.London:
sion lin and from it h alulatd th first rgrssion offiint. Latr, Galton G i l l h a m 'N . W . 2 0 0 1 .
otaind th hunran data h so dsird whn h orgnizd an anthropomtri Birth of ugzlis.N
laborator at th 1884 Itrnationl Halth xhiition lrld irr South Knsing- K v l s ,D . 1 9 9 . 5l. t h ]
ton' London. Th laortory ropnd in th Sin Gallris of th South Prss.
Knsington Musum aftr th hiition losd. ollting quantitativ dat Kiihl, s. 1'994'Th Nzi
Paul, D. B. 1995.otr
from oth pants and lrildrn, he was now abl to drnonstratgrssin
H i g h l n d sN. J : H u n
to th man for masurabl human haratristis (.g., hight). Y/hn h Parson,K.1914-1930.
plottd forarm lngth against hight h disovrd anothr irnpotnt statis- C m r i d g : m r
tial onpt' orrlation (i..' tll mn hav long forarms)' lulatd R i l l v .P . R . l 9 9 l . S
th first orrlation offiint, myriads of whih hav n omputd sin. tll UitdSttcs.B.
Galton lso am intstdin fingrprints and thir lassifiation; his work
was ntral to th dvlopmnt of fingrprinting as a fornsi thniqu.
Galton olltd many of his most important firrdings in Nturl Iheri-
Gam thory
tnc (\889). H also aquird his first protg6es'inluding Krl Prson. A Th study of nimal
fin nrathmatiian,Parson Wnt far yond Galton in formulating statis- t w ni n t u i t i o n a r r d
tial thory. anwhil, Galton was promoting ugnis nd th ida was invstigtion. fforts t
gaining tration irr the prvailing Soial Drwinist nvironmnt at th nd of al tools alld gams
th nintnth ntury. Positiv ugnis nouragd th sltiv rprodu- A gam in this sns
tion of th supposdly fit, whil ngativ ugnis aimd to liminat th - ing whn th outom
prodution of thos dmd unfit. Unfortunatly, ngzrtivugrrisprvaild. oths. hus gam l
Gme heor 599
'd, Hrditr G eius (1869). In th Unitd Stats, passge of state ugni strilization laws rsulted in th
dmonstrattht what h r- involuntarystrilization of thousands of individuals rgardd to mntally
..flmindd.''
d.Th ook trad pdigrs defiintor ugni dvlopmnts in th Unitd Stats wr
;i assumption was that if h followd with intrst in urop and lswhr' partiularly in Grmany.
!hn th Nazis m to powr thy pssd n involntry strilization law
;'slosstmal rlativs (.B.,
stinguishdthan thos furthr that rsultd in th strilization of hundrds of thousands of individuals. Aftr
l Wrnot inludd' probably !orld !ar II' ugni strilization gradually ndd in th Unitd Stats and
r m i n a n i n V i t o r i a n s o i t y . urop.rWhilugnisis Galton's most wll_known lgy, h also must b
s hypothsis.Although othrs rditd with important ahivmnts in filds s divrs as statistis' fingr-
lgood position might nsur a printing, mtorology' and xploration.
o r r l r i o nG, a l t o n a t t m p t d BlLIoGRAPHY
rote rditr Gis, Gal'- Brooks'I\^.2004.trmsures:The Drk Visiosnd Bright Idsof Fris
Glto. Nw York: BlomsburPulishing.
n for th stimtd15 million
Bulmr,. 2003. Fris Glto: Pioerrfrditvd Biontr.Bltimr:
r natural ailitis. LI also was
Johnsopkins UnivrsityPrss.
alyz statistially, partiularly Forst,D.w.1974. FrltisGltl:Th Lif d'V/<lrhof Vit:torinGius.New
lot asy to do with humn b- York:TplingrPulishingCo.
)austhy wr asy to grow Glton,F. 1852. Rntpditioninto th intrior of South-IWstrn Afria.
Jornl of tb Rol Ceogr1lhilSliet22:140-76.
rts h fund that sd diamtr
1853, rlpilSouthfri, London:John urray.
r frm progny of lag_sdd 1'855.Thrt of rul:r, Shiftsnd otriusuilblin Wild
Lnof tlrpopulation as a whol. otris.London:John urray.
;tial proprty that has n r- 1865.rditrytalntnd hartr ' mill'sgzi|2: 157-166,
{ dif{rntlsss of mutal t8-27.
us intrnationl stoks). Galton 1869. HrditrGeius' Londn: amillan.
1889.Nturl lherit.London: amillan.
mtrsof parntal sds vrsus
20|. b Art of Trul,or, Sbiftsd otritlsuilblei WiId
. H had drwn tlr fist rgrs_ outris.London: Phoni Pss.
ssionoffiint.Latr, Galton Gillham,N. !. 2001. Lif of Sir Fris Glto: Fr-lmfrin plortio to the
l orgnizdn inthropomtri Birth of gzis.Nw York: oxford l-Inivrsityrss.
hiitionlrld in South Knsing- Kvls,D' 1995.In the Nme of ugics.mridg,A: Harvard Univrsity
Prss.
l Sin Gallris of th South
Knhl, s. 1994. Th Nzi otloz.Nw York: Oford UnivrsityPrss.
d. llting qllantitativ data Paul,D. B. 199.5.otrolligHutn rdit, 186.\to th Prst.Atlnti
a l t o d m o n s r r r r g r s s i o n Highlalrds,NJ: HumanitisPrss.
ristis(.g.,hight). Whn h Parson,K. 1914_1930,The Lif, Lttrsd Lb<lursof Fris Glto, vo|s.
|vrdanothr impotnt statis- amridg:amridgUnivrsityPrss.
long forarrs).H alulatd Rilly'P. R. 1991.Th SurgiISolutio: istorof Iuo|utr Steriliztitl
i
the Uitd S/a/es.Baltimor: Johns opkins Univrsity Prss. -N.vG.
rih hv n omputd sin.
rnd thir lassifiation; his work
ng s a fornsithniqu.
Gam thory
'ant findings in Ntrl lheri-
g6s,inluding Karl Parson. A Th study of animal havior thrivs n paradoxs, affling inonsistnis
d Galton in formulating sratis- btwen intuition and vidn that ngg attntion nd stimulat furthr
ting ugnisand th ida was invstigation. fforts to rsolv suh parados rly inrasingly on analyti-
linist rrvironmntat th nd of al tools alld gams.
ourgd th sltiv rprodu- A gam in this snsis a mathmatial modl of stratgiintration, aris-
nis aimd to liminat th r- ing whn th outom of an individual's ations dpnds on ations takn y
ly' ngativugnis prvaild. othrs. Thus a gam has thr ky omponnts. Fist, thr ar at last two
Gme Thor
Fishr's spousal of this ugnis thsis in this part of th ook, h intnds th Having argud that
disussion to takn as an insparal tnsion of th prding part. Lwntin advans an
Tbe Getil Thor of Nturl Seletio is a point of dpartur in volu. tis is invitale. The
tionary thought, rsponsil in part for th origintion of thortial popula- rsult of a dpr,irr
tion gntis and what is ommonly alld th modrn synthti thory of and nutralist position
.!right's
volution. Getil Theor was followd y Swall and J. B. S. aility; thir fondnss
aldan's major works in 191 and 1,92,rsptivly. Fishr's viws on th pting r rjtingv
rol of natural sltionin volution ar widly aotdtodv. mrad hang and
ativly maintaind
|LIoGRAPHY
hld onvitions rgar
Fishr,R. A. 1930 [1958].h Geticlhorof NturlSltio.2nd d.Nw his ltrophorti dat
York: Dovr Publiations.Rlasdin a variorum dition by oford Univrsity inspird its transform
rssin 1999, editedy J. H. Bnntt.
Tbe Geti Bsis o1
Haldan,J. B. s. 192.The ussof uoltion.London:Longmans.
.Wright, duing volutionary g
s. 1931.volutionin ndlianpopulations.Gtis16:97-159.
-R..s. arful analysis of th .
apturs how gnti 1
Th Geneti Bsis of Euoltionr bge Lwontin, informatio
ody of data that wou
(Rihard Lwntin)
and rfind. In light of
In1974, Rihard Lwontin, a profssor at Hrvard Univrsity, pulishd an rlationship btwn t]
important assssmnt of th stat of volutionary gntis. Lwontin's 1974 tion, Lwontin argud
ook,The Geti Bsis of uolutior hge, onsidrd th trmndous plad with modls th:
hangs within evolutionary gntis wrought y th introdution of th- Indd, Tbe Gntic B
niqus from molular iology and th susqunt dvlopmnt of th nu- th ral prolms of
tral thory of molular volution. Lwontin's ook is notworthy for its standing th intrativ
analysis of th natur of volutionary gntis and th dynamis of sintifi
ontfovrsy and ontrovrsy rsolution. lLIoGRAPHY
Lwontin and his ollagu, J. L. Huy, hd touhd off a molular rev- Ditrih'. R. 1994.Th
lution in 1966 whn thy introdud th biohmial thniqu of l- lourl of th Histo
trophorsis as a mans of rsolving gnti diffrns at th molular lvl. ubby, J. L., and R. . l
htrozygosity in na
Lwontin livd that this nw ltrophorti analysis of gnti variaility
Drosophil psudoo
rsolvd th arlir disput twn th lassial and balan positions ovr
Kimura,. 1983.TeN
variability and sltion. Th high lvls of variaility dettd with l- ambridgUnivrs
trophorsis smd to support th alan position and th importan of Lwontin,R. . 1974.t
aland polymorphisms in volution. The lassil position's lif was that olumbia Univrsit
most mutations wr dltriousand most wr sltionpurifying. Thy strug- Lwontin'R. .' ndJ. L
htrozyglsityin n
gld to plain ltrophortivariaility until otoo Kimura and othr i-
htrozygosityin na
ologists gan to argu that most of th dttd polymorphism was in fat 542595-609.
nutral, nithr sltdfor nor against. In b Geeti Bsis of uoltior
hge, Lwontin mphasizd th ontinuity twn th arlir lassial-
alan ontrovrsy and th thn-raging nutralist-sltionist ontrovrsy.
Geetics nd the I
For him, th nutralist position was rally just th lassial position rought in
(ThodosiusDoz
lin with nwr moleular rsults. Whil this intrprtation of th ontrovrsy
maks a god as fo ontinuity in volutionary gntiS, it nglts th With grat harisma a
growing ontriution of biohmial rsarh to th dvlopmnt of molu- am a rallying poin
lar volution. gi of Spies (1'937\l
Gntisand th origin of Spis 60
part of th ook' h intnds th Having argud that on ontrovrsy has n transformd into anothr'
rsion of th prding part. Lwontin advans an plantion for why ontrovrsy in volutionary gn-
is a point of dpartur in volu- tis is invital. Th onntion and prsistnof ths ontrovrsis WaS a
.iginationof thortial popula- rslt of a dpr, irrsolval idologial onflit. Advoats of th lassial
:h modrn synthti thory of and nutralist positions hld a onsvativ attitud toward hang and vri-
.t/right,s aility; thir fondnss for staility ld thm to s sltion as a mans of a-
ly Swall and J. B. S'
sptivly.Fisher's viws on th pting or rjting variations. Advots of th alan position' howvr,
:ly aptdtoday. mbradhang and variaility; thy favord th ida that variation ould
ativly maintaind in a population y alaning sltion.Ths dply
hld onvitions rgarding th valu of hang xplaind for Lwontin why
his ltrophorti data did not rsolv th lassial-balan ontrovrsy, but
f NturlSIctio.2ndd. Nw
' r u m d i t i o n y o f o r d U n i v r s i t y inspird its transformation.
Th Geti Bsis of uolutionr hng should not b undrstood as r-
London:Longmans. duing volutionary gntis to onfliting idologis. Lwontin prsnts a
n s .G t i s1 6 29 7 _ | , , - * . o . , .
arful analysis of th hllngs of produing a thory of gnti hang that
apturs how gnti prosss intrat and dvlop ovr tim. Aording to
Lwntin, information aout molular level hangs was finally providing a
hnge ody of data that would allow thoris of volutionary gntis to tstd
and rfind. In light of nw molular data, Lwontin laimd that th ntir
lrvard Univrsity' publishd an rlationship twn thory and fat would hav to ronsidrd. In addi-
)nry gntis.Lwontin's 1974 tion, Lwontin argud that thoris of individual gns would hav to r-
ge, olsiderd th trmndous plad with modls that onsidrd gnoms and thir ompl intrations.
Indd' h Gtic Bsis of uolutionr hge nds with th harg that
;ht y th intrdution of th-
qunt dvlopmnt of th nu- th ral prolms of volutionary glltis an only addrssd y undr-
in's ook is notworthy for its standing th intrativ ontt of allls.
s and th dynamis of sintifi
ILIoGRAP|.]Y
llogy. On n lvl, Getis How populations bam isoltd was a sujt of onsidral study in
s in th 1930s' oth zoologi- th 1930s.any disiplinsontriutd to that sr.rjt' and many ..isolating
: informatiorr and rtd an mhanisms''w proposd ('g., gographi, ologial, hvioral' m-
lg to nw spis.It also pr- hanial,physiologial). Anything tht prvntd gnti hang ountd
and orrfidnto iologists in that disussion.Getis survysprominnt ampls' strssingth impor-
rhr ontiution to th su- tan of gographi and physiologial mhnisms. Strility in hyrids -
'ion studisas a rsarh pro- twntwo spissvd as spial as study in this disussion. A whol
haptrjn Getics is dvotd to this topi, illr"rstratinghow isolting po-
t th gnti strutur of pop- ssslos off th t;rp of gn flow.
.!7right's
l d g i v n g n t i i s t ss n s i t i v Following shifting alan thry, Dozhansky plaind divr-
r gnfrquny'and hngs gnlargly in trms of sltion and drift, with irumstans influning
rtd onsidrabldata on th th partiulrrlan of fors. Impotantly, Dozhansky is not a kn.jrk
wild populations.H also d- adaptationist.vn whn it plyd a dtrminant ol in shaping organisms'
hangsto th gnti strutur h thought sltionwas many layrd.It frequntlyshiftd dirtion and irr-
nviroIlmntalonditions. In tnsity,and it frqrrntlyprssd populations in ontraditory dirtions.
-arndmpirial sid of pop- Dobzhansky prsntd xprirntal snarios in whih suh ontaditory
jtas rigrous,with th vo- rssLrrs hd n distinguishd.Dozlransky did not rqr.tirdivrgnf<lr
l of thir data. spiation.Famously, h namd Drosophil mird as a nW speis vn
.ms:a hang in th frquny though it Was xtrnally idntial to D. psedoobsur.Inthis as' individ-
ivly oprational. Getis of- uls faild to rd so th asnof frtil offspring was sr-rffiint for nm-
rsntsa dtailddisussion of ing a nw speis.
ssion,along with th latr dis- In th widr fram, Dozhansky largly rstritdhis fous tn Geetis to
r t h t w o t h o t i e sl p i n s i n volutionwirhin spisand to th form:rtion of spis,in his strit sns<lf
on drivs adaptlrtion and is a rpodutivlyisolatd omrunitis. To plain th oigin of highr t.
: of populations.Sltion uss goris (.g.,nw gnra' familis, and lasss),h saw no rson to invok
from ndom mutation as wll additional mlranisms!suggstingmrvolutiorrmrlv tndd th po-
mal prossst work within sssof mirovolutior.
Ky to Ceetics's impat was Dozhansky's <rnrnfor ojtivity,rigor'
nsky'snly volutionary mh- and mpirial studis. His ook ws old ssrtion <l[onfidtrin th
..shifting alan'' th- priod's approah to volutionary studis. Getis askd narrowly dfind
right's
n f adptiv landsaps (s qustionsand uilt arfully on thik vidnbas.Dobzhansky omind
rphasizdppr.rltionsiz and this with los tis to rnthmatilmdls nd rigorus primntal tsts.
:han.Muh of a whol hap- I n p a r t , D o z , h a n s k yw r o t a s a r p r s n t t i vof n w w v o f p r i m n -
''sapproah.In fat, many vo- tal iologists attaking volutiorrry topis. H also wrot with an y towad
nodls fronr rdirrg Getis' skptis from othr disiplins nious to avoid unsavory ompliations.
tiv IandsapsWre ssntial Sylpathtiollaguspointd to Getis as a sign of a nw r in volu.
tionary studis.Th produt of Dozhansky's 196 Jsup ltursat olum-
rng in lll frqunis. For bia Univrsit, Getis rvivd the olumia Biologial Sris,whih would
n.This is th sond thort- boman outlt for othr ky ooks in th making of modrn volutionary
th prosssin dtail lding iology' that thory usually rfrrd to as th volutionry synthsis (Cain
'latdand physiologially ina- 1993\.
isolatd,divrgnrsultd Dobzhansky pulishd a sond dition of Gtis d tb rigi of Spcis
Dozharrskyargud that in th in |94| and a third in 1951. Flis Geetis of the uolutior Procss (1970)
l . .u n i t s .T h i r u n i q u g n t i was ir-rtnddas Gtis's fouth dition' ah nw dition inludd sor
iar dstiny, was ntral to that signifiant hangs to Dobzhnsky's viws. In partiular, finding signifiant
systmatih:-rngs of hromosoml vrizrtionsin th wild, Dozhal-rskygrw
(t06 c offro Sit- ilir
'lrvolutionar,vpross. Using his rpatdlip srvi to transformist thoris mad him hro of radiirl
o u l d ( 1 9 8 ) ,n o g r e t l v r o i or priphal naturalists who ausd Cr'rvirf prostituting his sin to
r ..hardnit-lg'' of th synthsis' politial onsrvatism. Dply inflund y IJauy's rystallography and y
: it as nturl dvlornnt of Buffon's hints onrning th unity of animal typs, Goffroy initially on-
ivdvrtrats:ls ing omposd of th sam st of organs and prts, at
tims lmost invisil in som fmilis, fully dvlopd in thrs. His l-
ratd studis on th vstigil prsnin rptils of all th parts that om_
ivsolutiotls: Organizrtionl pos mammalian auditory organs liitd th admiration of Goth and
ls 84:1-2-5. Hgl and provd, in his ys' th thory of analogus, or, in th trminol-
Spcis.Nw York: llr-r.ri ogy stalishedy Rihard own, homology. livd that strutural or
: l i r i o 1n9 5 1 . )
mhanial Onstr.lints during dvlopmnt miglrt hv b rsponsii
. . \ w Y l , k :( o l t r l n h i . l I l l i r ' s i r y
for th diffrnt ..alaning'' of various sts of parts in diffrnt farrrilis
In . (irn, ed.,Dimsios of and vn lasss.Diffrnt ratios of prssion of an organ or parts indud
ity Prss. diffrnt animl forms, thus produing diffrnt funti<lns. During th
l Soil ostrtio|1? 1820s rrnd 1830s h toyd with tlre ida that hzrngingnvironmntal on-
ditions might miranially afft th mryo, thus prodrring modilid or-
'>ns.Gtis | 6: 97_1 59.
gan dispositions' modifid funtions, and modifid animirls. His growing
:rossrdingand sltion rn
nl ogrss of Gtis 1z thortialoldnss md Goffroy xtnd his thory of th unity of type to
-J.. all animal forms. His laim that zrrthropodswr uilt on th Sam plan as
vrtratsgav uvir th opportr-rnityto ngg his f<rrn-rr frind in a
'2-1844) hatd,famous disput on th uniry of typ that startd t th Aadmi ds
sinsin 1830. Although historians hv rpatdly ssrtdthat vitory
s orn in tamps n April 15, Wnt to uvir, many in urop sidd with Goffroy and with philosophial
rrtd mdi[studisin lrdr antOmy.
th patronagof th r1,strrllog. BIBLIOGRAPHY
almost rtain dath in Sptm.
Appl, Toy A. 1987. h uuir-GeoffroDbte:reh Biolog in the Dds
rtd profssr of zollogy at th
bflrDri. Oford: oxford UnivrsityPrss.
urll,not withtrr"rt som ltd osi'. 1'988.Th g of Lmrk:uolutiorhoriesi Fr,1790-180.
r o s h e r h i s t , r v t t p r t m n ri l l Brkly:Univsityof lifornirss.
sas.Unlik ltvir,who st;ryd GoffroySaint-Hilai,. 1818.Pbilosophietlmiq.Paris:lvlquignon-
]ptdth invitation to join th Marvis.
pol6onBonapart.His work on L Guydr,H.2004. ti GoffrlSit.ilir,1772-1844: Visilr
Nturlist.higtl:Univrsityof hiagoPrss. -P..
.t .ls Wrhis privtlyomnltl-
:rasinglsusPii()usin tlr rn-
Flmpir.Bak ttl Paris in 1802, Gillspi,John H. (. L944|
l' h was adiv lagging hind
orrlyin 1807 an]ws appointd John Gillspi has n th most prominnt hi.rmpionof th argumnt that
i r r a r i si n 1 8 0 9 . h u l k o f h i s pattrnsof molulr volution r plaind bttr y th ation of ntural
rringth l820s Goffroy ld th sltilnin vrial nvionmntsthan y tlr so-alld nutrl thory. His
llmnts in polvr, of whih u- work has inludd th dvloprntof omplx mathmatial modls that
i v .D u r i n gt h I 8 0 sh i s p r s i s . dsri natural sltion undr varying onditions and th omparison of
his mystial ()vrtons'nd his vrious pattfns of moleulr vriation within and among spisto th pr-
l l t i t t rl ' i t h r h P a r i s i l r l rs i n - ditilnsof thos, arrd otlrr, modls.
l l s i d r h lth r r l u g ' h o uru r t l p . A studnt of Knihi Kjima, Gillspi rivd his dotoral dgr in
\970 from th Univrsity of Tas at Austin. approahd th qustion of
), his ullgisof Lamark, and
608 Gillspie
frquntly thought to prss onisly in vrs that whih is diffiult tl ar- Baus of his wor
tiulat. So, too, th iologist in Goth smsto sk to xprss th elusiv pointd as on of th f
forms that undrwrit th priens of natur. fr Biology in Brlin in
frns twn gog
BIBLIoGRAPY
ras only rpsntd
Kant' I. 2000. ritiqu of th Poturof ldgmt.P. Guyr,d. amridg: lutionary transformati
ambridgUnivrsityPrss. mutations. His idas .
Rihards,R. J. 2002. h Rlmtiocptilof Lif. higo:Univrsityof
phnoopis and hom
hiago Prss.
Smon,D., nd . Zajon,ds.1998.Goth'sWof Sic:A hrlmolog phnotypi hangs.c
lf Ntur. Alany: StatUnivrsityof Nw York Prss. usd to dsrithsr
Stur,D.2002.In dfnsof xprin: Gth'snaturalinvstigations
and phors in th history o
sintifiultur.In [,. Sharp' ed.,mbridgompnion to Goth, 160-168. Mayr's Sstmtis
amridg: amridg Univrsity Prss. -1.K. part in ordr to rfut
ntly in th ontxt
tyi hangs ar oft
Goldshmidt,Rihard Bndit(1878_195s)
ntworks of gns, a r
Rihard Goldshmidt was on of th most ontrovrsial gntiistsof th shmidt's arlir idas
twntith ntury. Known as a skilld primntalistand a rativthorist' found rfug ftr his r
his idas about th natur of th gn and th rol of systmimutations (or
|BLloGRAPHY
larg rrrangemntsof hromosoms) in volution hav n onsidrd
hrtial, whil at th sam tim thy hlpd to rystalliz th so-alld modrn Ditrih , . R . 2 0 0 3 .R i
synthsis' whos partiipants wr' at th vry last, unitd in thir opposi- Nture Ruit,us Gt
G o l d s h m i d tR, . [ 1 9 4 0
tion to Goldshmidt. or rntly' StphnJay Gould (1982, in his intro-
S. J. Gould. Nw Hl
dution to th rprint of Goldshmtdt's Tb teril Bsis lf uolutio) and 1'958.Tbeorti
othrs (.g.,Ditrih 2003) hav rvaluatd Goldshmidt and argud that 1'96'1d u
his idas dsrv at last as muh onsidration as th prvailing dogma of Mayr, . 1942.Sstm
adaptationism. Univrsityrss.
Rihard Goldshmidt Was orn in 1878 in Frankfurt am Main into a pro-
minnt Grman Jwish family and grw up in a wll-to.do miliu. H gan
his studis of mdiin in Hidlrg' attnding lturs of oth Otto
Gould, StphnJa
Biitshli and Carl Ggnaur. owvr, his ral intrstwas zoology, and in Stphn Jay Gould w
1898 h movd t unih t study with Rihard Hrtwig. His first aadmi ultural stting that n
appointmnt Was as Hrtwig's assistant in Munih. As part of his tahing ionolasm, and his lI
dutis h organizd th laoratory in omparativ anatomy' whih in part Columia Univrsity h
xplains his vast knowldg of animal morphology' anatomy, nd ell iol- Norman D. Nwll. F
ogy. In his own rsarh h fousd on prolms of gntisand ytology, ollaorator, Nils ll
spially on th prolm of s dtrmination. Glldshmidt workd with on whih h would sr
th Gypsy mot Lmtri, whose gographial variants, whn matd to In his arly work, (
ah othr, produd an assortmntof so-alld intrmdiatintrss.To growth of individuals r
ount for ths intrmdiatforms, Goldshmidt prposd a dynami and Using quantitativ m
quantitativ modl of gntisthat was asd on th undrlying physiologi- portion of individual <
al ation of gne produts, hen th nam physiologial gntis.Th asi ats and timing f tht
ida of physiologial gntis Was that diffrnt variants of gn produts powrful tool for ana
should hav diffrnt physiologial proprtis' Suh as ration tims, whih xtint organisms.Mo
an aount for th diffrnt osrvalphnltvos. provid a potntial ml
Gould 611
that whih is diffiult to ar. Baus of his work in gntis and physiology, Goldshmidt was ap-
l s kt o x p r s st h l u s i v pointd as on of th founding dirtors of th nw Kaiser \Wilhlm Institute
for Biology in Brlin io \91' Soon his intrstslso inludd th gnti dif'
trnstwn gographi as of Lmtri. Ths diffrns twn
ras only rPrstd mirovolutionary vllts. To aount fo lrgr vo-
Juyr' d. amrilg: lutionary transformations, Goldshmidt postulated th istn of maro-
mutations. His ids on this sujt wr also motivatd y his study of
/e. hiago: Univrsity of phnoopisand homoti mutations, oth prsntinglarg oordinatd
phnotypi hangs. Goldshmidt's notion of ..hopful monstrs''' whih h
of Si: Phltltolog
usd to dsri ths rrovl phnotyps' is on of th most unfortunat mta-
Prss'
'tural invstigatilnsr-rd phors in th histoy of biology and was soon ridiuld y his ritis. rnst
,)mpdia|1 to G lth,',\:f. 1.r's Sstemtics d the rigi of Speis (|942) was writtn in major
prt in ordr to rfut Goldshmidt. Howvr, as w hav land mor r.
ntly in th ontt of volutionary dvlopmntal iology, larg phno-
Baus h hallngdprvailing viws, som rightly saw Gould as a riti Spkoski, D., and . I
of neo-Darwinian orthodoxy. Nonthlss, h was an ardnt admirr of Univrsity of hi
harls D:rrwin, singhis own work as an pansion of Darwin's ntrpris Stanly, s. . 1975. A
Ntil Adem,
and no l-nnsa rfutation of it. Gould's arlir work is most assilto
|979. rl
thos sking arr intodution to lris idas;his finl mgnum opus,T Stru- Stbins, G. L.' and F'
tr oi uolutior Thor1' Q002), is a rilr ut unwildy tom, mor rad- Since 21'3:967_
r I yq u a r i dt h a n r d i n i t s n t i r t y .
In addition to his original rsearh, Gould pulishd numrous ssays' most
notly in th magazine Nturl Histor. Ths wr olltd in sris of Grant, B. Ros
st-sllingooks, starting with uer sice Dr,in (1977a). Othr works ad- and Grant, Pt
drssd msllrsof human brirr siz and intllign(Gould 1981)' th izarr
Roslary and Ptr
tint irnimals of Camian sas (Gould 1'989|,and th rlation rwn si-
win's finhs in th (
n and ligion (Gould 1999|. H tstifid as an xprt witnss i th Arkan-
..rtion formd on of th
sas ourt as whr sin'' was shown to a rligious dotrin.
dsriing th fdin
Gould nvr hsitatdto join an intlltual fight. As a result, Gould amth
populations; doum(
prmir puli fa of volution in th Unitd Stats during th last qurtr of
trns of mt hoi
th rwntith ntrrry.
strutillg th phylog
th signal as study
BIBLIOGRAPHY Th Grants riv
.'J7. with honors from I
Allmn' D., P. . Klly, nd R. . Rlss,ds.208' StephJ Gluld:Rfl-
tilsltt His Vie of l-if. Nrv York: Oxford UnivrsityPrss. ridg. Thy mt an
ldrdg,N., nd S. J. Gould. 1972.Puntuatdquiliri:r: to phylti
An altrnativ lumia. P.R.G' tool
gradulisrrr. In T. J. . Shopf,d.'odIsi Plobillrlgy, 82-115. olumia; B'R.G. t
Srll-alliso: Frmirn'ooprnd o.
dotoral yar at Yal(
Gould, S. J' 1966.Allomtryand sizin ontognyirndphylogn'Bkllogil
Ruitl,sof the mbridgPhiloslphilSoiet41'z587-640. Univrsity of Mihig
1971. D'Ary Thompson and th sinof forrn.Nelu Litrr istor 2: positions at thsun
229-258. island id populati
1977,.l.uer si Druitt.Nw York: Nlrton. gos and morphlog
1977.togd Phllg1,. amridg'A: arvard Univsitrss.
diffrnt islands.B.R
1981.Tbistsr of n. Nw York: Nrrrton.
1988.on rplingth idaof progrsswitlr an oprationalnotionof among individuals i
dirtionality. In . H. Nitki, ed.,uolutilr Progress,319-338' hiago: aly pprs on sl
Univrsityof higoPrss. patri spiation. Th
1989. WderfulLif: h BurgssShld th Ntur of Historl' thm to th Galipag
Nw York: Norton. hallngsof logisti
|996, FilI Hous: Th Spredlf llfrtlm PItoto Drtui.
Th Grants hav r
Nw York: r.rgn.
|999. Rtlcksof gs:Scind Religio i tb Fullessof Lif. luding the Darwin,
Nw Yok:Ballanti. and tlr Darwin-.W
2002' Th Strturelf uoltiorThor.amridg,MA: Harvard Grants wr ltd
UnivrsityI)rss. thy wr oth lt
Gould, S. J., nd N. ldrdg.1977, Puntuatd quiliria:Th trpond rrrodof
2007, P.R.G. in 200
voltionronsidrd. Pllbilllg
3: 115_151.
1993.Pr-rntuatd quilibrir"rmotnsof ag.Ntur 66 Z2.3-Z27. S i n s( B . R . G . i n 2
Gould, S. J., and R. . L,lrrti.7979. h spandls of San ro and th
BILIoGRAPl
Panglossin pradigm: itiqtrof th adaptationist programme,Proedigs
of th Rl Soitof Llndrl'SrisB 205:581-598. Grant,B. R., and P. R.
Slzr,.J.,d. 1993. Udrstdig SitProse.alison:Univrsiryof wisonsin Popultion:h L
Prss. higoPrss.
Grt nd Grt 615
som rightly saw Gould as a riti Spkoski, D.' and . Rus, ds. 2008. Th Pleobiollgil Ruolutio. tago
ss, h was an ardnt admirr of Univrsity of Chiago Prss.
r pansion of Darwin's ntrpris Stanly,S. . 1975. A thory of volutin aov th spis |v|.Prodigs of th
Ntiol Acdm of Scis 72: 646-65.
s arlirwork is most assilto
1979. rouolutio, Pttr d Pross. San Frniso: W. . Frman.
his final magnum opls, Tb Struc- Stbbins,G. L.' and F. J. Ayala. 1981. Is nw volutionary synthsisnssary?
ih ut unwildy tom, mor rad- Sic2I3: 967_971'. -R.D...
Grant, P. R. 1986. olog d uolutkl lf Drtui's Fis. Prtnton, NJ: Tbulr Vie of the P
Printon Univrsity Prss.
praisd Darwin's on t|
Grant' P. R., and B. R. Grnt. 27. I]o d Wh Spies ultipl: Tb Rdition
-].. thir dinurgh days t
of Drtui's Fihs, Printon, NJ: Printon Univrsity Prss.
trs srouting from sp
two mn diffrd. A s
Grant, Robrt dmond (793_1874)
and fw mournd his p
Robrt dmond Grant' an dinurgh-orn spong xprt, ir-rtrodudom-
|BLloGRAPHY
parativ anatomy to Britain in th 1820s and 180s. H inrporatd th
studis of Jan.BaptistLamak and tinn Goffroy Saint-Hilair' apt- Dsmond,A. 1984a.Ro
ing th ..transformism,''or volution, aompanying thir works. Grant also transmutationist. /O
indutd th young harls Darwin into Lamark's viws. Darwin, ord y 1984.Rort.
l,turson ..palo
mdiin at dinurgh Univrsity ln 1825-1827, studid Noth Sa invrt-
95_413.
rats with Grant, who had om an MD in 1814. As an old man writing |989' Tha Pliti
ls Autobiogrph, Darwin ralld Grant's admiration for Lamark. Rdil Lodl.ht
Grant hampiond Hnri d Blinvill's ontinuous hin of lif (rathr S l o e n , . R . l 9 8 5 .D r w i
than Gorgs uvier's disrt embrnchemets) and Goffroy's lif that th transformism. In D. I
N J : P r i n U
nrliv
sam organs orrd throughut th hain-hn Grant's laim in 1825 to
hav found th homology of th vrtrat's panras in mollusks. Studying
spongs in 1825-1828, Grant oind th group's nam, Porifra, and h
mootd a simpl, dfnslss,frshwatr form as th ..parnt'' of th armord Gray, Asa (1810-1
marin spis. Darwin xamind th larva of th llal ryozoan Flstr un- Asa Gray was on of tI
dr Grant. Grant livd ths lrva wr rlatd to th ,.ova'' of alga and ia. H ame on of
pointd ak to the spontanouslygnratd..monads'' lying at th juntion mnts drivd from th
of th plant and animal kingdoms. At dinurgh Darwin was alrady roah- of volution. A dply
ing issus surrounding th birth of individuals and spis' vn if h rjtd logial impliations of l
Grant's monadism upon bginninghis own vlutionary spulations. ural sltionould
As th first profssor of omparativ anatomy at London Univrsity in y a wis and nvol
1827, Grant alrtd th mtropolitan lite to th Lamarkian thrat. H Gray was orn in Sr
joind th Gologial Soity ounil in 1832, just as harls Lyll was pu- i n m d i i n n d p r a t i
lishing his antievolutionry volum Priniples of Geolog. Grant's fossil l- tahing so that h w(
tures promotd th ..dirtgnration'' of sussivspis,on giving irth Fishr Profssor of Nat
to anothr. And his studntsw amind on th onditins rspnsilfor stood that h would o
..originating
and ffaing th tmporary organi film on our plant'' (Zoo|- onsult with othr bot
ogy amination Paprs, 1857_58, p. 6, in ..Grant on Zoologial Sujts,'' Amrian plants. H w
ollg olltion DG 76, Univrsity ollg London). ( 1 8 3 8 _ 1 8 4 3 )a n d v
Grant took th Fullrian hair of physillogy at th Royal Institution in ot of th Northert
187, ut that markd his apog.H had lost his ounil sat at th Zoo1og- January 30, 1888.
ial Soity in 1835, whn th zoo's aristoratipatrons gav his onsrvativ Gray,s work on th
rival, Rihard own, pfrntial tratmnt. At th Gologial Soity, Own Amrian plants mad
ountrd th fossil impliations of Grant's transmutationism. Grant lost r- voatd y Louis Agas
sours and his finans ttrd.Th 1840s found him living in a slum, his liation of th ri
shay swallow-tail oat as unfashionl irs his lturs. A rising youllg lttr to Gray, datd S
Thomas Hnry Huly in 1852 mrly laughd t Gfant's ..ntriity.'' Darwin and A. R..sa
At th nd, Grant nvisgdplantary ooling as th motor to powr th vo. 264-267|. Gray sus
lution of warm-loodd mammals and irds from dinosaurs and otrosaurs.In taks from Agassiz at t
Gr 617
,'ri'sFihs. Printon' NJ: Tblr Vieu., lf the Primr Diuisios of tb iml Kigdom (1861) h
praisd Darwin's the origin of Speis whil rminding his old pupil of
Vh Spisltltipl: h Rdition
-J.T. thir dinurgh days togthr. But th bulr Vi,s ountlss volutionary
ln UnivrsityPrss.
trssprouting from spontanously gnratdmonads showd how dply th
fwo mn diffrd. A sminal figur in th radial 1830s, Grant outlivd his ag,
4)
and fw mournd his passingin |874'
spong prt' introdud om.
lBLIoGRAPHY
nd 180s. H inorporatd th
r Goffroy Saint-Hilair' apt- Dsmond,A. 1984a.Rlrt. Grant:Th soialprdimnt of a pr-Darwinian
npanyingthir woks. Grant also transmutationist.lournl of th Histor of Biolog \7: \89_223'
1984.Rort. Grnt'sltrviwson organidvlopmnt: Th Swiny
mark'sviws. Darwin, ord y
Lturson ..palaozlogy,''1853_1857.rhiusof Nturlistor 11
| 8 2 7 , s t u d i dN o r h S a i n v r r - 95-4\.
) in l8l4. As an old man witing 1989. Th Politis of uolutio:orpholog,ediie' d Rform i
; dmirationfor Lamark. Rdil Lodo, hiago:Univrsityof hiago Prss.
ontinuous hain of lif (rathr Sloan,P. R. 1985.Darwin,sinvrtbrat rogram'1826-1.86: Pronditionsfor
zts|and Goffry's lif that th trnsfomism.In D. Khn, d.,Th Drii Hritg,71-120,Prinrl,
NT: Printon Univrsitv Prss. -4.D.
-hn Grant's lim in 1825 to
's panrasin mollusks. Studying
group's nam, Porifra' and h
m as th
..parnt''of th armord Gray,Asa (1810-1888)
l of th loal ryozoar' Flustr un- Asa Gray Was on of th lding otanists of mid-nintnth-ntury Amr-
..ova'' of alg and
rlatdto th i . H m o n o f h r l s D a r w i n ' s m o s r a t i v s u p p o r r s u
. sing gu-
. . m t l n a d s .I.y i n ga t t h j u n t i o n
d mnts drivd from th gographi distriution of plants to dfnd th thory
urgh Darwin was alrady roah- of volution. A dply rligious man, h also wrot xtnsivly on th tho.
als and spis,vn if h rjtd logial impliations of Darwinism' attmpting to show how volution y nat-
volutionaryspulations. ural sltionould ronild with th lif that th world was dsignd
n t o m y r L o n d o n U n i v r s i y i n y a wis and bnvolntGod.
to th Lamarkian thrat. Gray was orn in Sauquoit, Nw York, on Nvmr 18, 1810. H traind
32, just as harls Lyll was pub- in mdiin and pratid for a fw yars, ut aandond this for part-tim
ls of Geology. Grant's fossil l- tahing so that h would hav mor tim for otany. H was appointd
ussivspis'on giving irth Fishr Profssor of Natural History t Harvard in 1842, whr it was undr-
l on th onditions rsponsil for stood that h would onntratlusivly on otany. H visitd urop to
:ganifilm on our plant'' (Zoo|- onslt with othr otnists, ut his own work fousd mainly on North
..Grant on Zoologial Sujts,''
Amrian plants. workd oo a Flor lf Nortb Ameri with John Torry
lgLondon). (1838_1843) and vntually produd a highly sussful ul of the
o l o g y t t h R o y l I n s t i t u t i o ni n Botl of tbe Northr Uitd Stts (7848). H rtird tn 187 and died on
ost his ounil sat at th Zoolog- J a n u a r y3 0 , 1 8 8 8 .
:ati patrns gav his onsrvativ Gray's wlrk on th lassifiation and gographi distriution of North
. At th Gologial Soity, Own Amrian plants mad him suspiious of th ..rationist''position bing ad-
; transmutationism.Grant lost r- voatd by Louis Agssiz. harls Darwin onfidd in Gray for th pu-
)s found him living in a slum, his liation of o tbe rigin of Speis in 1859, and an astrat of Darwin's
. rising young
a s h i s | t u r sA lttr to Gry' datd Sptemr5, 1857, formd part of th joint papr y
..ntriity.''
;hd at Grant's Darwin and A. R. rJallain 1858 (rprintd in Darwin and !7al1a1958,
ling as th motor to powf th vo- 264_267). Gray susquntlydfndd th thory of volution against at-
l from dinosaurs and ptrosaurs. In taks from Agassiz at th Amrian Aadmy of Arts nd Sins.
618 Group Sltio
)nismon th asis of otanial g- othr volutionary fos. This dsristh first prt of Darwin's passag
l876). Gra saw how th disprsal quotdaov. Fortunatly, groups of individuals who nfit ah othr will
d with adaptation of populations ontriutmor to th total gn pool than more slfishgroups' as dsrid
w homs,ould aount for th ..for th good of th
in th seond part of Darwin's passag.Traits that ar
Lritytwnth plants of astrn group'' an volv y naturl sltion, ut only y a pross of sltion
nd y noting that in arlir go- among groups in a largr population and oftn in opposition to sltion
:adarossAsia and North Amr- among individuls within singl groups.
g th i ags, with only th two It is impossil to dsri th ()npt of group sltion without also
r nw rmaining. dsriingits turulnt history. any iologists during th first half of th
ious to show that Darwin's thoy twntithntury did not shar Darwin's insight; thy assumd that adapta-
In a srisof rtils (rprintd in tions straightforwardly volv ell lvls of th iologial hirarhy, fom
y prossy whih spis ould gnsto osystms.This position am known s navgroup sltion
npatil with th lif tht th and was widly ritiizd in th 1960s, spially y Gorg . Williams in
y a wis nd nvolntCrator. his ook dpttio d Nturl Seltil(I966), whih am a modrn
.astfulnss and rulty of natural lssi.Williams affirmd th importan of multilvl sltions a thort-
oah, and h suggstdthat varia- ial framwork, agring with Drwin that group.lvl adaptations an
tto.Darwin ojtdto this viw volvonly y a pross of twn-groupsltion.Williams thn mad th
of thistivolutionism, in whih mpiriallaim that twn-groupsltionis almost invarialy wak om-
y suprnaturalrathr thn natu- pard to within-group sltion. It was this mpiril laim that turnd
..th thory of individual sltion,'' whih b-
multilvl sltion thory intr
am th onsnsusfor th rst of the twntith ntury. Susquntvolu-
tionary thoris of soial havio, suh as kin sltion, riproal
tio b Nturl Sltil,l. altruism, gam thory' nd slfish gn thory, wr dvlopd pliitly as
altrnativsto grOup sltion.
\: Hrvard UnivsityPrss.
tthr Uitd S,a,es. Bosron: Th 1960s onsnsuswas asd upon thr argumnts:(1) twn-group
sltionis thortiallyimplausil, rquiring a dliat ombination of pa-
; Prtiig to Driism. New ramtrvalus to prvail against within-group sltion;(2) thr is no on-
vining mpirial vidnfor group sltion;and (3) th thoris proposd
. North Amri. 2 o|s. Nw York:
-P.J.B. as altrnativs do not invok multilvl sltion in thir own right. All thr
argumntsganto qustiond,vn as arly as th 1970s, although it was
diffiult to argu in favor of a topi that for many had om taoo. Modrn
thortial modls of group sltion ar fr mor plausil than thir prd-
ssors.Williams himslf aknowldgdth importan of group sltionfor
,rolmwith soial lif and its po- traits suh s sx ratio and disas viruln (.!illiams 1'992: Wll'Liams and
assagfrom Th Dscnt of Nss 1991). All volutionarythorisof soial havior must assumth -
ttn that although a high stndard istnlf multipl groups to iologially ralisti.!hn ths groups ar
tgto ah individual man and his idntifid, olprativ nd altruisti traits ar sltivly disadvantagous
yt that an inrasin th numr within th groups and rquir twn-grlupsltion to volv, as always
th standardof morlity will r- nvisiondy multilvl sltionthory (Sorand.Wilson 1998).
:iovr anothr.'' A major vnt in volutionary thory ourd in th 1970s with th dis-
amth asis of a thortial ovry that th singl organisms of tody wr th groups of past ags (Mar-
:rultilvlsltion (Sor and \WiI- gulis 1970; aynard Smith and Szathmary 1995)' volution prods not
sd on rlativfitnss' a trait that only y small mutational hang,ut also y singl.spisgroups and multi.
ngin frquny within th group. spissymioti assoiations oming so intgratd that thy om highr-
lf th slf atually drassin fr- lvl organisms in thir own right. Ths major transitilns of lif, as thy ar
v omtint in th asn of alld,hav ourrd ptdlyand possily inlud th origin of lif itslf as
620 Grrlt,tpSeletiolt
groups of ooprativ molular rations. Dspit multilvl sltion th- Rihson,P. J.' and R. B
ory's turulnt past' it is th aptd framwork for studying major transi- rsformed um
tions. Th alan btwnlvels of sltionis not stti ut an itslf volv. Sor,., and D. S. Wilso
UselfishBhuior.
A mjor transition ours whn sltion within groups is supprssd,n- .!illiams,
G. . 1966.Ad|
ling etwn-group sltion to om th primirry volutionay for. urrt uolutior
Tlr paradigm of rnajor transitions did not appar until th 1970s and was 1992.Nturl Sl
not gnralizd until th 1990s, ut i.rlradyit sms likly to plain th vo- UnivrsityPrss.
lution of soial inst olonis (.Wilsorrand Hlldoblr 2005) and humans as Williams,G. C., and R.
th first ultrasoial prirnt spis (Boehm 1999). Th traits assoiatd with QurtrlRuie'o
Wilson, . o., rrdB. Hol
human morality Ppar dsignd to suppress sltion within groups' similar Proedigsof th N.
to gnti mhanisms suh as hro-losomsand th ruls of miosis, nabling
sltion among grollps to om th primary volutionary for. ultural
prosssan aus groups to om vry diffrnt in thir phnotypi prop-
rtis,vn whn thv ar gntillysirnilar.Multilvl sltionhirs om
th aptd framwork for th study of human ultual volution, in xatly
th sam way as th study of major transitions (Rihsn and Boyd 2004).
Th rvival of grop sltion might sem traordinary until w rmm-
r th:rtWilliams (|966) aptdmr.rltilvl as a thortial fram-
slti<lrr
work, riting group sltion on th sis lf an mpirial lim that it is
lwas wak oIrp:rrdto within-group sltion.Whn Williams hangd
his mind aout th thortial and mpiial vidn' h himslf rvrtd
ak to multilvl sltion. For fml.iasd s ratios (\Williams 7992'
49), he statd, ..I think it dsiral . . . to raliz th:rt sltion in fmal-
isd ndlin popul:rtionsfavos nrirls,nd that it is only th sltion
among suh groups tht an favor th fmal ias''' For th volution of vir-
..Th
uln in disas ol.]nisms(Williams and Nss 199|, 8), h statd,
volutionary outom will dpnd rn rlativ strngths of within-host and
etwn-hostomprition in pathogn volution.''
Unforrunatly, th fild as a whol hzrsnt yt ahivd a nW onsnsus.
Books nd artils Writtn from multilvl prsptiv appr alongsid
othr ooks and artils that ontinu to trat group sltion as ir rjtd
onpt. Ttooks hav en partiularly slow to rflt th many dvlop-
mnts that hav takn pla sin th 1960s. Th trr-rlnt histry of this im-
ortant sujt is rviwd y Borrllo (2005) in an rtil appropriatly
titld ..Th Ris. FalI. and Rsutionlf Grourr Sltion.''
BIBLIOGRAPHY
Bohm, '. \999' Hirrh i tb Forest:glitriismd th uolutio of
ttl tt ltrui sm. arrrridg,MA : Hzrrr,i-r
rd Univsit,vPrss.
"t
Brllo,. .200.5.Th ris,fall' nd rsurrtiln of goupsltiorr.
deuor
29:4347.
Darwin, . 1871. Th l)sctof n, tdSltilli Rltio,o Sex. Nw York:
D. Appltonar-rd .
arguIis,L. \970. rigi of ukrotil/s.Nw Havn,T: Yal Univrsity
Prss.
MaynirrdSmith,.|.,nd . Szthmarv.l995. Th jorrsitiosrlf Lil?. Nw
l,tlrk: !7. . Frman.
Group Sltio 62
)ns. Dspit multilvl sltion th- Rihrson,P. J.' and R. Boyd. 24.Not b Gsll:olt, Itur
rmwork for studying maior transi- TrsformdHum uolutio.Chiago:Univrsityof hiago Prss.
.!ilson.
Sobr,., and D. S. 1998. I.JtoOthrs:Th uolutio d Pshologof
rtion is not stati ut an itslf volv.
UslfishBhuior.amridg,A: Harvard UnivrsityPrss.
on within groups is supprssd, n- Williams,G. . 1966.dpttiod Ntrl Seletio: ritique of Som
n th primary volutionary for. rrtuolutiorThought.Printon,NJ: PrintonUnivrsityPrss.
J not appar until th 1970s and was 1992.Nturl Sltil:Domis, Leuelsnd hllgs.oxford: Oxford
ady it smslikly to xplain th vo- UnivrsityPrss.
\fillims. G. .. and R. N4.Nss. 1991,.Thdwn f Darwinian mdiin'
lnd Holldolr 2005) and humans as
'm 1999\. Th traits assoiatd with QurterlRuituof Biolog66:1_22.
Wilson,. o., nd B. lldolr.2005.usoiality:Origin and onsquns.
'prsssltionwithin groups' similar Prodigsof the Ntiol dem of Sics\2: 167_1371.
lmsand th ruls of miosis, naling -D.S.\{1
primary volutiony for. ultural
ry diffrnin thir phnotypi prop-
nilar. Multilvl sltion has om
f human ultural volution, in xatly
sitions(Rihrsonand Boyd 2004).
smtraordinary until w rmm-
ilvlsltionas a thortial fram-
asis of an mpirial laim that it is
tp sltion. \Whn \/illiams hangd
rpirial vidn, h himslf rvrtd
,l-iasdsx ratios (.Williams 1992,
. to raliz that sltion in fmal-
nals,and that it is only th sltion
[malias.''For th volution of vir.
..Th
and Nss 1991' 8), h statd,
rlativ strngths of within-host and
volution.''
las not yt ahivd a nW onsnsus.
rltilvl prsptivappar alongsid
to trt group sltion as a rjtd
arly slow to rflt th many dvlop-
960s.Th turulnt history of this im-
lo (2005) in an artil appropriatly
r of Group Sltion.''
d th uolutio of
Zglitriism
arvardUnivrsityPrss.
of groupsltion.
iurrtion deuor
L
rviw, Ludwig Rittimyr (1825_1895), an mryologist at Basl, hargd
Hakl with fraud. H osrvd that in illustrting th prinipl of
rapitulation-or th iognti lw' as it am known-akel had
rprsntdvry young mrvos of a dog, hikn' and turtl as morpholog-
ially idntial. Riitimyr maintaind' howvr, tht Hakl had mad th
asy using th sam woodut thr tims. In th nt dition of th ook
(1870),Hakl usd only on illustration of a vrtratmryo at a vry
arly stag and said it might as wll b th mryo of a dog, hikn, or tur-
tl aus you annot tll th diffrn. Th damag, howvr, was don,
and th harg of fraud would haunt Hakl for th rst of his days.
Dspit th ontrovrs, |trtiirlihe Shopfugsgeshicbte mad a powf-
{ul impat on its rdrs, spially on th topi of human volution. H
rprsntdnin spis of humn ings along a tr of volutionary d-
vlopmnt,with th Papuans and ottntots at th lowst rnhs, losr
to roots i te Urmsh, or ap-man' and with th auasian branh at
th highst lvl, arrying at th top rahs th ditrranans' Grmans,
Jws, and Aras. Although Hakel shard many of th raial viws om-
mon to nintnth-ntuyuropans, h was diddly not anti-Smiti'
an attitud whih on of his disfftd studnts hld against him. H ar-
gud' fllowing his frind, th linguist August Shlihr (1821_1868), that
grads of human mntal aility prssd grads of languag omplxity
and that th uropan and Smiti languages hlpd rat a orrspond-
ingly ompl mind-a gnral thsis that Darwin adoptd in The Desnt
of .
| 1867, aftr visiting Darwin and othr British sintists in ngland,
akl travld to th nary Islnds with thr rsarh assoiats.H
prformd th kinds of primnts on dvloping siphonophor mryos
that would garnr fam for Wilhlm Rou and Hans Drish som 20 yars
latr. H also gan work on a systmati analysis of alrous spongs that
would yild a thr-voium study, Die lks,ijmm (Th alarous
spongs' 1872). In this work, Hakl attmptd to provid what Bronn
maintaind was nssaryto show th viaility of Darwin's thory, namly,
mpirial proof that spisdsntWas mor than a thortial possiility.
Hakl also argud, mploying th ignti law, that in anint tims an
organism, having th strutur of a primitiv spong (and th form takn y
mtazoansin gastrulation),plid th anint sas. This am his gastraa
lld his ooks with his owrr
thory.
rthosisof volutiona11-
mdrbildr' 1905), is an
Bausof his various invstigationsof marin invrtrats,Hakl r-
ws griving th dath of a ivd th ommission in th lat ]870s to dsri systmatially svral
with whlm h had just had lasssof organisms drdgd up y HS hllegr. ovr a 10-yar priod,
l wll as th hnging tims. h omposd svral larg volums on mdusa, alarous spongs'
Sulit (grtly inflund y siphonophors, and radiolaria-with mor pags produd than y any
hd in th sm yr this othr author in th sris of bllgr rports. Th ommission indiatd
th high rgard of th sintifi ommunity fr his work in marin iology.
That rgadwas lso prssdy th many honorary dgrsand wards h
r i v dd u r i n .h i s I r t i m .
626 Ifckel
lian hrdity.Indd, Haldan al Institut.In his famous pap, ..Th ost of Natural Sltion'' (1957),
ortialpopulation gntis. In Haldan argud that thr is a ap on th rat of adaptiv volutiorri.tus
of 10 mathnratialpaprs - of sustitutionl<lad,that is, usa sustitution (y mr.rtationor hang
Mndlism; th last in th sris in nvironmnt)ould not start mor frquntly than vry 300 gnrations.
r puliation of his ook e Haldan'slaim was latr alld Haldan's dilmma.
Haldan was awrdd th Drwin-.!llaMdal of th Royal Soit in
aldn'ssrisof paprs on vo_ 1 9 5 8 , a n d i n 1 9 6 | r i v dt h K i m r d a l o f t h L I . S .N t i o n a l A d -
o f t h o r t i apI o p u l t i o n g n - my of Sins.Aftr a prodigious arr)ldn did of olon an in
his ontmporaris;h wntd Bhubanswar' Orissa, India, on Dmr 1,,|964' Srious illnss, howvr'
natural sltionin N{ndlian did not dampn l-lldan's spirit. In his por, ..anr Is a Funny Thing'''
ry pimartly fo its ritial om- h wrot' ,.I know that anr oftn kills / But so do ars and slping pills /
own mathmatialxplorations And it an hurt on till on swats / So an ad tth and unpaid dts / A
n that. Haldan aomplishs a spot of laughtr, I am sur / oftn lratson's ur / So lt us patints
dlian hrdity,ut also, undr do our it / To hlp th surgons mak us fit'' (lark 1968). Haldan has n
sonrlmhanis,ytology, and dsridas larg than lif du to his srvi i th Blak Wath, his famous
wit, nd his politial viws. Haldan Was' tO sur, iologist anl thn
's work was by no mans r- som.
lishd his famous Dedls, or
BIBLIOGRAPHY
: on sinin its soial ontxt.
's lark, R. \).1968..|BS:h Lif d Worh rf .|.B. S, Hldna.Lodon: oward-
frind Aldous tlu|e,s Bru
ibutd Haldan's prinipl, th Mann.
Hldan,J. B. S. 192-3.Ddlus,rlr Sid the l-utur.Lonlon:KgarrPul,
y quipmnt animals rquir, in
Trnh,and Trunr.
a th olltion Possibl Worlds 1924'^ mtlrmatial thovof naturaland artifiials|tion, I.
ationson th origins of lif wr rstionsof th mbridgcPhilosopiISlict23: 19_41'
r Oparin (1894-1980). In 1930, 1927.PossihlWorldst| othr ss1,5. London: h:ltto and Win]us.
ns of nzym kintis. Dspit \929.The <lriginof IiI.Rtiolistttl:.
190.zms.London:Longmans.
l harlott Frankn Burghs in
192.Th ssof uolution.London:Longmns.
l harlott was marrid at th 1934. matlrmatial thoof naturalad rtifiials|tion' X' Geetis
formarring Haldan. J. B. s. 1.9:,41.2429.
st ln Spurwy in 1945. |97. Te fftsof variationon fitnss.AmrinNtrlist71 37_49.
ol1g,London, and am 1957. Te ostof naturalsltion. Jourlof Getis.5.5: 511-.524.
,.Th H a l d a n J,. B . s . , A . D . S p r u n t, n d N . . a l d a n 1
. 9 1 5 .R d u p l i a t i oinl l m i .
ldan publishdhis papr,
.|rlrl of Gntis5: 133-135. -R.'s.
th dlinof th avragfitnss
LalalllWas proportional to th
: in fitnss.Haldan's onlusion
Hamilton,W. D. (1936-2000I
utation load.'' At this tim, Hal-
Ltyand, in fat' wrot a numr !7illiam Donald Hamilton was on of th major volutiorraryiologists of
ill Workr. In 1950, aldan th twntith ntuy. any of his ontmpor:risompard lrim to harls
lism and to th rims of Sovit Darwin' partly ausoth rrrnshard dp lov of natural history and
: Sovit Union during whih si- partly aus both attemptd to solv som of iology's iggst mystris.
)arwin Wrsnt to th gulags. Prhaps th largst of ths mystrisis th qustion of altruism, prforn-ring
l of th Royal Soity in 1953, ats that nfitoths at a ost to th altruist. Strprfiilly,volutionary th-
th Univrsity of Paris in 1956. ory suggeststh21tmutants hilliting mor ..slfish''hzrviorshould zrlto
rtd to India nd ultimatly - wip out altruisrn, aus y dfinition th altruists hv rdud fitnss
)tstof British poliy during th whil th slfishindividuals prospr at thir pns.Yt altruism aounds in
t DofSsort th Indian Statisti- natu. Afrian kingfishrs fd and prott young that ar not thir own;
60 Htilto
lronyesommit suiid whn thy sting intrudrs, and vn mor notaly sltir.prssursmust fl
from an volutionary prsptiv' workr s ar stril, sarifiing thir prodution still wins in th
own rprodution to hlp th hiv flurish. i n d e i l L r s i n gp p u l a t i o n
Harnilton's ir-rsightfor solvir.rgtlr puzzl rvirs dptivl1,simpl: slf- vid suh flututing sl
sarifiial vior an volv and prsist if it nfitsrlativsof th altruist attak as thir hosts vol
austhy shar th altruist'sgns.Thus aiding a siling, ousin, or othr th ontinual arrag of i
rlation an still favord y sltior-r' vn if it nrans forgoirrg th pro- ual rprodution advanta
dution of th altruist's own offspring. Hanriltn xprssdthis in a qua- tlrn turnd his :rtt
tion that lras om to alld Hrilton's rul: ltruism is favord whn ing tlrat thr, too, prsst
lc>1'lr, whr is th nfit, is th ost of th at, and r is th dgrof tion of showy sxual orn
rlatdnsstwn th dnor and th ripint of tlr altruisr. on should malsrsistant rl plrrlsit
thrfor b ror likly to s th most s.lrififor thos most losly r- flashy tr:ritsas indiirtors
latd, sulr as silings, nd lss sarififlr thos lss losly rlatd, suh as as w,ll.Parasits lrr of
third ousins. Th pross is alld kirr sltion,arrd th ida has n usd u d p r s s u r t r l m a i n i
lvith grat sussto plain a varity of prosssin animals, inluding th hang thir dfnss,tht
ooprativ irds nd suiidl smntiond rlir, although it dos not al- ass. Ftnalsthus s
ways provid a omplt answr. T s,it turns out' ar though a gnti th spifi gns onfr
puliarity of thir inst group vry lt>slyrlatd to thir hiv mts, mor n tstdin nurrlrous
so than thy rvould to thir own offspring, and h.nhlping thir mothr ata1,parasitsdl s
(th qun) to ris thir sistrsand rotlrrs,who rvill om fr.rtr-tr rpro- vlopnrnt of som su
dutivs,will favord undr manv irumstns. Ntul histor-v,part
In addition to his work on altruisr, amilton wrot aout th volution arilton, and his thor
of animal groups in
..Gomtry for
th Slfish Hrd'' (|971), aout sx ra- organisms havdin th
tios, aout aging and snsn'and :rout th origins and funtion of o- of 6, following orpli
opration and fightir-rg.uh of lris work is mllrkd y a r,villingnssto ongo to follow up tln
ntrtain nw idas and viw old inforrtion witlr a frsh y' and h was a polio vain. H won
rathr solitary sintistthroughout most of his lif. Th ida of kin sltion Kyoto Priz, th rafo
wnt rlativly unappritd for svrirlyars ftr its initial pr"rlitionin mdial iology), tlr Di
7964,ut vntully it ron f th ornstonsof mdrn volution- ml Bhvior Slit1,
ary ilogy' His ids almost always am ky omponnts of thir fild, Soity of Natuarlists.
whthr immdiatly or aftr a lag whil othr sintistsaught up to him. iplirrs,from ntomol
Hamilton vntually .rmintrigu1with on oI th gratstprolms of furthr tstimon to thr
vollttion' th xistnof sual rprllr.trion. This ld to an intrstin p- B|LIoGRAPl{Y
ilsits and thir ffts or-rhost olog,v,havior, nd volutiln that lastd
for th rst of his arr. S is a onundrum aus,just as th altruist H a m i l t o n ,W . D . 1 9 7 1 G'
1:29.5-311'
slrould los in a ra with a slfish onlptitor' an :rsualspis (on that 197.5.Gmblrss
simply rpodr'rsitslf rnaking gtltiallyidntil opis) shltrldout- rf BIogl'50:l7.5-1
ompt a sual spis tht rquirs two individuals to produ a singl 1 9 8 0 .S v r s u
offspring. rtainly som organisms' ir-rludingsorrustaans!lizards, in- 1995a.NrroF
sts, and plants' do rprlduasully. ost, howvr' pay th pri of Nw York: \(. H. Fr
1 9 9 5 .N r r o I
s, with all of its onsquns.As Harrrilton askd in a 1975 ook rviw, P
..Why all .Wy Oird Llnivrsitr'
tlris silly rigmtrrolof s? Why this gavottof hrot-l.losoms? 2005. N.lrr Rr
:rll ths uslssmls, this striving nl wastful loodshd, ths grotsqu Hmilto.Vol.3, La
horns, olors, . . . and why' in th nd, novls . . . ahout lov?'' (p. 175).
Hamilton, long with a trv othr sintists,suggstdthat sltionmust
favor th shuff1ingof gnotypstrvt-t gnrirtiol.ts
and hrrsx. But th
Hmilton 61
phylognti systmatis was long ft, a short ig to, a tiny rainas, and an ap-sizd rain-to-ody
3 8 ) .S o m o f H n n i g . s n m i s ratlo.
s on ognizingmonophylti To dat, th rmains of twlv Homo florsiensisindividuals hav n
follows quikly adoptd th found in assoiationwith faunal rmins of dwarfd lphant-likeStegodo,
nt ladists argud that th a- giant Komodo dragons, rts. and ats. harrd ons nd lustrs of rd-
ary rlationship ut th most dnd, fir.akd roks wr also found in Liang Bua. High dnsitis of
: data (Ridle 1986). othrs ston tools wr avatd from th av' inluding ors, anvils, rtouhd
,arsimonywas only on of sv- tools, and flaking dris. Baus of ths assoiatd finds, th disovrrs of
vlutionry rlationships on Ilomo florsiesissuggstthat th spismad tools, hul-rtd,usd fir, and
rtesinspird y nnig's rvo- uthrd and ookd mat. Although th suggstion of highr ognitiv
ran 50 yars aftr its original ailitisin suh a small-raindspisrmins ontrovrsial,it is sr'rpportd
y an analysis of th impssions of tlr rain that wr imprintd insid
LB1's rainas.A ast of th rainas (or ndoast;s figur) rvals that
th rain was highly onvolutd and uniquly organized (wird),with a num-
<i,http://www.ilogi.d/ br of advand faturs.For ampl, th tips of th frontal los wr -
pandd in a gion that is known to impotant for ronitoing intrrral
mi thought. ul Ruielu of
thought and for planning ahad. Ths and othr dtails of LB1's ndoast
rilosophy. In Iferrig ontradit th suggstion y a fw rsrhrs that it may from a modrn
rr Assoits. human afflitd with a pathology known as mirophaly.
logtish Sstemtik. Blin:
Th disovr of Homo fllresisis Was tmly surprising aus it twn dissimilar hal
showd tat Homo spis Was not th only hlminin arund 18,000 yars of intrmdiat forr
go (and for),as had n widly livd.Sintistsar mixd on thir trirrtionof sinrilr
intrptationsaout whr rhis nw spism frm. Som tlrink that it is ogy ritria aptur t
a dsndant from iggr.odid Homo rtusallstors that lramdwarfd asd on pattrns of '
aftr migrating to th island of Flors. This hypthsis is in kping with th lotid sustitutiorri
wll-known island rul, whry larg nimals onrsmallr (lik th aov- squns.Belttso
mntiond Stegodo\ and small animals lmlargr (.g.'th giilnt Kornodo volurion, fr.rrtlr
dragons) on islands whr rsours ar limitd. othr workrs liv that whih divrg as a r
Homo florsisismay lrav n dsnddfrom a small anstor' suh as paralogous gns,wh
thos in Luy's genus (tlstrlopithus), nd that its nstors simply r- Th onpt of hor
rnaind small aftr arriving On th islirrrdof Flors. Tim will tll. tirns. Aristotl fotrn
irl' and vn ftlr
|BLloGRAPl{Y
analogy, th idsa
Brown,P.' T. Sutikna,tr4.J. orwod, R. P. Sojono,.|tikmo, FJ.Y/. Saptlmo, and dat tw G
R. . Du..2004.^ nrvsmall-did hominirrfrllth Lt Pliston lf a r l v 1 8 3 0 s ( s A p t
F l o r s I. n d o n s i aN. t ' 4 . 3 1 1
: 0 5 5 - t 0 6l .
st undrstoldin tL
Flk, D., . ildolt,K. Srnith'..|. tlrwood,. Srrtikn, P. Bror,vn,Jatmiko,
. W. Saptml, B. Brutlsdn,and F. Piro.200.5.Th rainof LB|, Homl ments) nd within th
floresisis'Si8:2"42_24 5. Goffroy argud fr
lv{orwood,.J.. P. Brown'Jatikmo,T. Strtikn'. W. Saptomo,. . wstaway' plarr of all ninrals,
R. A. Dr"r, R. G. Rlbts, T. da' S. Wsisttl'nd . Diubintonl. 200.5. rognizd thirt idn
Furthrvidnfor small-bodid homininsfrm tll l,tlistln f Flors.
-D.F. arranging diftrnt ft
I n d o n s iN . I r 4 3 7 : | | 2 _ | 0 | 7 .
vids a rational syst
th rrotion of th ar
similritis 1m()ng al
Homology
In th ontt lf t]
Homology is on of th ntral onpts in iology. It rfrs to th similarity tnth ntury' th h
twnmorphologial haratrs'gtls'and othr iologial rraits, suh as th osrvd pttrn
haviors,al-nongdiffrrrtspis.Srn of th st-krrownnrplsof ho- in thir ommolt vr>
mology ar (1)th wings of irds, th arms lf htlnlans, and tlr forlims of for th ronstrutio
othr ttrapods and (2) th ony lmntsin th gill rhs of sharks and th phologial ritia,
thr middl ar ons (mallus,inus, nd staps)in humans. Ir-roth ass, asrtain homology r
th similaity btwn haratrs is stlishd indpndntly of thir siz, ws sd on th ssl
shap, and funtion. This fat is rfltd in Rihirrd own's dfinition of a si.rilarmryolrian
homologu as ..th sam orgn in diffrnt animls und vry varity of ogy would soon run i
form and funtion''' to distinguishdfrom an analogu as ..a part Or Organ gl ll, it ws not lt
in on animal whih has tlr sam funti<lnas anothr part or lrganin a dif- samnsstwntrv
frnt amphibian lss,lf
What is alady lar trom Own's dfinition is that in ordr to stalish th sion that th original l
istn of homology, w nd to hav a st of ritria tht allow us to dtr- prolemati.Anothr
nrin th samnss btwn diffrnt lraratrs. Irr th past' omparativ Th rognition that l
anatomy arrd mryology providd th tools for lromology assssmnts;tody ph ologial strtlturs
w also us molular and sequndata. Th morphologial homology ritriir tu.n haviors ws
inlud (1) position-homology is mor likly if two haratrs ar found in Today, th homol
sirnilar positiorr within two organisms; (2) strutur-lromology is mor likly if disr.rssions withirr i<
two haratrs shar many individital faturs; (3) trar-rsition-homology - irls nd dfinitionsof ]
oologv 6.7
Josph Daltorr L{ookr was a lading otanist of Vitorian Britain and on of Alan, NI. 1967.Th Hl
harls Drwin's staunhst supportrs. H was on of th first sintists to I{ookr,J. D. |845, he
Disot,rShipsre
informd of Darwin's thory and playd a lading rol in hlping Darwin Bros.
to formuiat his idas, spially in th appliation of volution thory to | 8 5 5 .F l o r l d
ioggraphy. Idi. I-ondon:\/ . 1
Hookr ws orn on Jun 30, 1817, th son of Sir \WilliamJakson F{ookr, 1860. FIor ,ls
who was dirto of th Royal Botnial Gardns at Kw. otaind an
MD at Glasgow in 189 and latr that yar joind }i.S rbus on a four-
Hrloker 6L)
I
mology onptsand (2) iolog- yar ploratory voyag to Nw Zea|and, Tasmania, and th Antrti. H
O g y o n p t s r p r o m i n n i n susquntlyprodud a six-volum survy of th otany of ths rgions. In
[y usd approah, ladistis or 1847,e travldto Sikkim, Npal, and th astrn Himalayas, snding ak
ls of spis(lads)y th prs_ xoti plants suh as rhododndrons' whih soon am favorits in uro-
orphis).Ths synapomorphis pan gardns. H pulisd Flor Idi in 1855, th yar in whih h -
i onsqunof thir phylog- am assistant to his fathr t Kw. In 1865, h sudd his fathr as
l r s u l t i n gp h y l o g n t i s y s t m diretor, ontinuing Kw's rol as a lading entr for th onomi otany
haratrizdy a uniqu st of of th British mpir. Hook did in 1911.
|n 1'844, Darwin informd Hookr of his idas on volution and in 1'847
o t h r h a n d , I s o m p h a s i z s ,i n ..ssay'' in whih his thory was first dsried in dtail. Th
snt him th
ry' th dvlopmntal origin of two argud for many yars ovr th thory's impliations for th gographi
rnotypistrutursar sn as a distribution of plants. ookr livd that disprsal had takn pla ovr
1ram.As Girntr.!7agnr dfind anint land onntions twn th isting ontinnts' whih had sus-
th sam individual ar homolo- quntly n sumrgdy gologial ativity. He was partiularly ativ in
straints,ausd y loally ating dfnding th viw that thr had on n a grat ontinnt in th south-
ation'' (Wag'er 1989, 62). Te rn hmisphr, of whih Antartia, Australia, and Nw Zea|and ar ut
aptursth distriution of h_ sattrdrmnants. Darwin argud instd for th disprsal of sds aross
intndsto plain th mhanis- th oans y flotation, winds, and irds. Hookr was at first skptial of
l ar to found in th dtails of volutionism, ut vntually h am on of th arlist onvrts to th
nology onpt is thrfor at th thory.
th rnt synthsis of volution. Alng with harls Lyll, Hookr advisd Darwin on th ours of ation
h should tak aftr reiving Alfrd Russl Iil7alla'spapr in 1858. Hookr,s
introdutory ssay to is Flor Tsmnie of 1860 was on of th first major
puli ndorsmnts of th thory. H spok aftr Thomas Hnry Huly in
ch Biolog i tb Deds befor th famous dat with Bishop Samul Wilrfor at th oxford mting of
th British Assoiation in 1860, and som obsrvrsthought his ontriutin
gy. Novartis F-oundation
was mor prsuaslv.
l Bsis of omprtiu Biolog. Darwin and Hookr ontinud to dat iogography in th following
dads.In 1878, Hookr usd his presidntial addss to th Royal Soity
:nt and th dvlopmnt of th of London to dfnd his thory of an anint ontinnt in th southrn hmi-
777-788.
sphr. At th sam tim' h ndorsd th viw favord y Darwin and many
,tom d Phsbklg of the
othrs that th main sour of mor highly volvd spis Was th northrn
| ollg of Srgeos. London:
rgions of urasia and North Amria. Ltr in his lif' he gan to ak
nept.
nlReuietu away from th thory of sunkn land ridgs' undr prssrrrfrom Darwin,
"'\i}'.il',. !alla, and othr supportrs spousing th prmann of th ontinnts
and oans.
r.) BlLIoGRAPHY
lisess
i World ut of ontrovrsil, inspirirrg th first intrnational onfrn on th topi, fol-
lowd y lfticid: omprtiu d uolutior\' Perspctius (1984),
A. Nw York: Random
oditd with Glnn ausfatr. By thn, Hrdy had om inrasingly in-
of Ntur'sost trstdin dilmmas onfronting working mothrs thrrlughout history and
-.L. dp into prhistory. Sh st out to amin aross spis and ulturs th
options that mothrs hav -rployd,from sking th assistnof ..allo-
mothrs'' to nglt and ;rndonmnt. (Allomothrs ar mmrs of a
goup who assist a mothr in ring hr infant') rJ?hrrorr Ntur ap-
:lt nthr()pologistsnd a pard in1999, I-illrr Jlurl namd it on of tlr st ooks of th yar.
rtlrropolgistand prima- lnrasingly drawrr to th qustion of infant nds, irr 2005 sh oditd
m w o r ko f s o i o i o l o g i - Dhlm !orkshop rpot on attahmnt and ondirrg.
:riutions.Hrdy is most Hrdy's ooks hav n translatd into svral lnguags.Although hr
lutionary stratgisusd idas of irrfantiidwr lng rgardd with suspiion within th anthropol-
ift away from primatolo- ogy ommunity, in 202, at th annual mting of the Amrian Anthro-
ors.Mo rntly' Flrdy pologial Assoiation, other Nture was awardd th Howll Priz for
ship to thir hildrn. Al- outstanding ontriution ir-rBiologial Anthropology.
od'' ntrirl to soioiol. Hrdy hld th positiorr of Profssor of Anthropolog ilt th Univrsity of
r q u l - r l p p o t u n i tyr i t i aliforia at Davis from l984 to lrr arly rtirmntln I996' As Profssor
nr1,pshologistsfo ig_ mrit, sir ontinus loirrgrsrh,writing ooks, ad svil-rgorr th d-
itorial boards of t,llutilr Athropolog anl ut Ntur. Hrdy and
nily. Sh graduatd from hr husand liv in lionia, whr thy ar involr,din hitat rstoration
'nthropology ours that and sustainalagritrltur.
n a m ) ,w h o m s h m a r -
BIBLloGRAPY
Vhil raring thir thr
d to updt Attahmnt artr'. s.' L. Ahnrt,K. . Grossrann,S. B. Hrdy, ' . Lam,S..W.Porgs,
ids as ooprativ rd- and N. Sahsr,ds.2006.ttbmtd Bodig.ambridg,MA: IT
Prss.
iowly, arly ttahmnt
Hausfatr,G., and S. Hrd' ds.[1984]208.Iftiide: tplt'tttiue
d
and motionir1dvlop_ ulutior Prspti us, Nrv Brunswik'N.|: Aldin Transatiol-l.
dy, SaralrBlff.1977 L1980|.b Lltgttrstlf btt: Fnlld NIlStrtgies
1'970,about th sam lf Rprldtttilll. rrrridg, Prss.(1980,aprak
A: Hrvrd Univsit1.
Soiobiolog: Th Ntu d i t i o nw i t h r l r rp' r i ; r . )
r it was pulishd. Irvn 1981 [1999].T Womht Nueruolud.arnidg, N{A:Harvard
.Wilson UnivrsityPrss.(l999, papraklitiln with nrvpr[')
nd y and by
1'999.l>tbrNdtut : Histlrof otrs,Iftsd Nturl SItio.
rn th anuman langurs Nw York: Pantlrrr.
d le Strtegies of \h ag. h ttp://www'itrna.m/srhhrdy.htm. -u.s.
t h r r s ho n i n f a n t i -
pathologial rSpons t)
sardthat hypothsis. ln-
Hutton, Jams(|726-1797)
d infnts bon to fmals JrrrsHrrttln, a Sottish-orrr]otr and gntlmn firrtnr' is onsidrd
s wr f from pssiv. th fathr of gology irr Britain. H livd ruh of his latr lif in dinurgh
xampl,y mating with and was a major figur in th so-alldSottislrrrlightnmnt.A ontmpo-
trop, fmals strov to rar of David Hum nd Adam Smith, h hampiond h th Plutonist th-
ory of gologyas wll s th mthodologyof uniformirrianism.As a Plutonist,
i (198\), Hrdy provoa- h argud-against th so-alld Nptunists, who thor"rghtthat th world had
h prinrat origins of pa- n formd y pripittiln from Wat-that th ntr of th arth is vry
on infantiid rmaind hot and that roks ar formd y moltn lava pushirrg up to th surfa,
644 Hutto
mitarian.he argud that th half of th nintnth ntury. It took harles Darwin in tbe origi of
ausstoday ar th san-las Spiesto turn suh idas into full tlrory of volution.
qologialformations an
|BLIoGRAPHY
l-volunr work titld Thor uvir,G. 1813. ss\'o th Theoro|.th rth.R. Krr' trans.dinurgh:
.W.
; ddonly y its osu- Blakwood.
Huttoll'J. 1788.Thoryof th arth;Or an invstigation of th laws osrval
in
i s h d m o r r a d i l y a s s i -
th omposition'dissolution,and rstorationof lnd upon th glo.
ition arl rognizd and T'rstios of th Rol Soitof dibrgb 1, no. 2: 209_304.
mor ppular thorizir-rg of |794. luestigtiooi th Priiplsof Kou,ldgd of t|lProgrss
:rar lnny onvuIsionsin of Reso,from Sesto Sid Philosophy.dinurgh:A. Strlranand
H u t r o n ' s u n if o m i t r i n i s m T. Cadll.
r's atastrophism.This am 1795. hor of th rth, ith Proofs d ]llstrtiozs.dinurgh:
willim rh.
harls Lyll, proly th
Lyll, . 1830_1833.Priiplsof Golog:Big n Atte?ttrl plin tb
rg h:rrlsDarwin. Formr hgesi th rth's Surf,lb Rfrto ussNott''ilt
Hutton's importan lis in ()prtion.3vols.London:John urray.
famously spok of th world Playfair'J. \802. Illustrtionsof th Huttoi Thor of th rth. dinurgh:
) to f a n d , ' ( 1 7 8 8 , 3 0 4 ) , \filliam rh.
1805.Biographialaountof th latDr. JamsLluttonF. R. S. din.
l, ..thmind smdto grow .]-rstios
, (Plyfir of th RrlISoitof diburgh 5, no. 3: 39_99.
1805) oviusly, Rphk,J. 2003. Th Vlho Ftlud Time:Jnls uttld th Disourof
al history rad litrally. Al- the rth,sAtiquit'London:Simonnd Shustr.
proaly h was (like many Rudwik,M.J.S. 25.Brstigthe Limits of Time.Chiago:Univrsityof
t, on who ss th glory of hiagPrss. -.R.
tht has n muh ndorsd y suh thinkrs as Rihard Dawkins (1986).In ian intrvntion in url1
1910, Huxly rivd his first taulty ppointmnr, irr zoology rtoxford. mmrs of th Zoolgia
Whii in Oford h undrtook his first majrr rsarh suss, study of soity. Thy ford his
ourtship rituals and sxual sltion in th grat rstd gr. H also d- Aftr sveral ontriuti
vlopd skill in primntal mbryology. onstrution, in 1'946 l
Huxly's arr irrludd many mployrs. In 1,9|2, aptd an offr Nations duational, Si
from th Ri Institut in Houston to dvlop a nw dpartmnt of iology, rol suitd uxly prf
.sfhn .W.ar
whih provd diffiult. World I rupted, h rturnd to ngland. l i t i a l a s p t so f t h d i r
During th war, Huxly srvd mainly in rmy intllign.In 1919, h r- gav UNSCO an agndz
turnd to Oxford's zoology dpartmntnd onntrtedhis attntion on d- vision h had advotdf
vlopmntal iology and physiologial gntis.In 1920, h rportd on th s l f f o r i t s f u t u r v o l u t l
aility of hormons to provok mtamorphosis in axolotls. This rivd W hav to tak rsponsih
onsidrabl prss attntion and srvd as zratlyst for his liflong involv- family planning Programs
mnt in th nrdi. uly also invstigatdth rgulation of rats of gn t i s i n d v l o p i n g r r n t r
xpression and prolms of rlativ growth' suh as allomtry. In |925, sions, spiallyrgarding
movd to King's ollg London' apting th position of profssor in zool- md a vision of progrs
ogy and laoratory dirtor. Two yars lar,uly rdud his position to Aftr laving UNS
an honoary on and lft King's to pursu a arr as a frlanwritr. His sintist-lrity.H wr
ontriutions to his sin ar summarizd in Problms of Reltiu Groruth Lysnkoism), travld, at
(Huly \932) and The lemets of perimtI mbrology (Huly and Progrss and ultual d
d Br 194). Ths arnd Huxly ltiorrto th Royal Soity in 1938. ampaigns (Huxly 1954
Huly's most important frlanprojt was a ollaoration with . G. vious work and noua
.slls
and G. . to survy iology and th history of volution, titld Th Si- rsarh again. uly di
e of I'if (1929). During th 1930s, Huxly ontriutd to nwspaprs' As a sintist, Huxly l-
magazins,radio, and film. took on book and ditorial proits,as wll as bhavior,dvlopmnt,
spakirrgtours (sfor mpl uly |94). Smir-rgly vrythinginvolv- howvr, Huxly norml
ing puli sinin Britain in th 190s inluded Huly in som way. H is fousd attntion on iol,
st known for his rol on th BB radio program ..Th Brains Trust'' during thory and xplanation.
\forld \/arIl. Huxly's aility to ass th mdia provd ssntillatr in and his intrsts shiftd il
his lif. w o r k k t o t h s p i n r
In 1935, Huxly rturnd to institutional mploymnt, oming srtry topis am lss impl
of th Zoologial Soity of London. This ntaild marragingth soity and gav ohrn and ml
.Whil
its zoologil gardns in Rgrrt'sPark and \il/hipsnad. srtary,u- ply fat gathring.
ly kpt up his puli ontriutions. H turnd his rsarh intrsrsto sys- Huly's prfrn fo
tmatis and volution. Huly was partiulrly intrstdin volution lading tndd for nonspialist
to adptation and to th formation of nw spis.Polytypi spisnd go. ( 1 9 6 ) ,a n d i n h i s f i l m P l
graphial varitis (suspis)wr of intrst as thy suggstd..volution a m p l st o i l l u s r r a t g
in ation,'' a phras Huly oftn rpatd. He oind th trm line for car- providd a as for th
atrs that show gradual hang ovr a gogaphigradint.FI lso nour- mating havior' flight,
agd sarh into natllral nd sxual sltiorr'Dr"ringthis piod, uxly sud admiraly in pr<
organizd he N Sstemtis (1940) irnd wrot uolutio: The odr mation and th trati
Sthsis (1942). a supr xampl. This r
.!Var
Th outrak of world II in urop radially hngd Huly's om- stad, th ook's aom
mitmnts. H wrot xtnsivlyagainst fasism and against Nazi raist iol- ordination ot informt
ogy, nd h strssd th importan of intrnationalism and dmoray mriad fornls of slti
(Huly 1941,194). Aftr 1939, Huly ampaignd vigorously for Amr- Britain for a gnration
Hle' Juli S. 647
sid. In rfrring an ap anstor ovr a ishop who prostitutd his talnts' fid to lad industrial so
Huxly, th Puritn, oozd moral sority to sham th worldly Nilrfor. him. As Dissntrs (out
Suh onfrontations only xaratd th Vitorians' ..risis of faith.'' In taking Oford and ar
1861, uxly denid own's laim that aps lak th human's third rral tion'' lgitimizing thir g
lo' whos avity has a postrior horn with a projting ..hippoampus Huly mad th wor.
minor.'' hrls Kingsly tnTh'Wter Bbis laughd aout ..hippopotamus a naturalisti maning, .
majors,'' and th affray rought human volution squarly to th puli's at- spiritulist A1frd Russl
tntion. Huxly mad his nwly aquired Nturl Histor Ruieta an anti. Huly oind th word
Own organ and st th trnd with his artil ..On th Zoo\ogtca| Rlations to ..agnosti'' aout
of an with the Lowr Animals'' (1861). Huly's wrking-lass talks, ou- with lean hands to tr
pld with a ltur on fossil humans (fousing on th rntly unarthd N- aS th nutral profssion
andrthal Man). formd th ornrston of his uidece s to 's Plc i his lst yars rguing h
Nture (1863). shaping a ntury of vo
uxly studid Dvonian fishswith lo fins and similar-looking amphi. Although volution w
ians, alld lyrinthodonts, from th strata surrounding oal sams.Rathr form, Darwinism's wal
than draw volutionary onntions' h mphasizd that layrinthodonts ltion had raisd manki
had prsistd unaltrd for immns priods. Initially, Huly livd that ..volution and this'' (
..moral indiffrn.''
th major groups had volvd for Silurian tims and had hangd littl t
sin. Only whn Darwin's Grman supportr' rnst Hakl, pulishd on tur within natur'' som
phylogny' or raial anstry' did Huly ronsidr. In 1'867,Huly rlas- Ultimatly, th supr
sifid birds gnealogially,and in 1868' h famously suggstdthat th small wht onrnd uly z
dinosaur ompsogtbus oti.d hav n anstral to irds (and may hav rd. Whras his Siet
orn fathrs). H dramatizd ths rlationships and promotd o. . tiv' his nin volums o]
Marsh's hors fossils from Nrask on his Amrian tour of 1876. on il- showing the Vitorian a1
lustration in Amri ddresss (1877), showing th inrasing lg and H. L. nkn alld H
tth siz from th small thr-tod on rohipps to th tiptoing who has vr livd.''
quus, was itd as th dfinitiv fossil vidn of volution for gnrations.
BILIoGRAPHY
Latr h dsrid th volution of th sondary palat in roodils.
.!hr Barr, A. ., ed. 1997.ho
Huxly's attitud toward Darwin's mhanism rmaind quivoal.
Darwin nvisagd variation ourring in minusul stps, Huly in 1859 eterssays.Ath
Dsmond,^. 1997.ul
piturd an innr for produing nw spis at on. His ntry on ..volu-
Rading,A: Addiso
tion'' in the cclopedi Britic (1878) mootd natural sltion only Di Grgorio,' 1984.. I
on' and Huxly's admi work ypassd it ompltly. uxly rsrvd Y | U n i v r s i tPy r s s
his disussion for popular talks: hr natural sltion was liknd to Lyons, S. L. 1999. homs
Napolon's troops rtreating from Mosow and rossing the Brsina Rivr, NY: PromthusBk
..vryon Paradis,J. G., and G. ' !(
hding only himslf,'' th fittst alon rahing th far ank.
PrintonUnivrsityP
ostly, Darwin's althusian mhanis srvd uxly in omating soial- Y/hit,P. 2003. ThlmsHl
ism or dunking rationism.Also, h avoidd volution in lass aush ambridgUnivrsity
Was training sinshoolmastrsaftr th duation At (1870). stalish-
ing his first propr laoratory in South Knsington in1'871', h fashiond a
pratial ..biology'' asd on th ommon plant and animal ..Typs.''This in.
flund shool urriula until th mid-twntith ntury. But inulating a
disrt ..Typ'' systm lft littl room for volutionary disussions. opning
Birmingham's nw Sin ollg, Huxly ontrstd th ryling of th
lassis at th Anglian univrsitis with sin's primntal approah.
Th undrvalud sintifi rsarhr' h ontndd, would ttr quali-
Htll, Thoms Hr )/
p who prostitutd his talents, fid to ld industrial soity.This dlightdth stl magnatswho finnd
.!ilrfor.
ham th worldly him. As Dissntrs (outsid th Anglian huh), thy wr ludd from
/itorins'
..risis of faith.'' In taking Oxford nd amidg dges and saw Huxley's ..nw Rforma-
ak th human's third rral tion'' lgitimizing thir growing ivi authority.
..hippoampus Huxly mad th word euoltloz fashionalaround 1870. Fo him it had
h a pojting
..lrippopotamus a natllrlisti mning, wlrras for th atholi St. Gorg ivart nd th
laughdaout
.slla
ion squarlyto th puli's at- spiritualist Alfrd Russl it onvyd th fiat of a highr powr. n,
Iurl Histor Ruitu an nri- Huly oind th word gosis in 1869. Biologists, as ultural ladrs, had
..on the Zoologial Rlations to ..agnosti'' aout suprnatual issus. They had to appoah sin
<ly'sworking-lasstalks, ou- with lan hands to trustd.Huxly hlpd mold th imag of th sintist
on th rntly unarthd N- as th nutral profssional,nd of volution as dmystifidsin.H spnt
; uide s to is Plce itl his last yars arguing his antimiraulous as in thologial ontrovsis,
shaping a ntury of volution-rationdat.
rs and similar-looking amphib- Althlugh volution was fin for dmolishing Anglian doors losed to r-
urroundirrgoal sams. Rathr form, Drwinism's wak-to.tlr-wall r"rpshotprturbd Huxly. A rural s.
phasizd that lyrinthodonrs ltion had raisd mankind, ut it ould not plain our ar of th wk. In
..volution
Initially, uly livd that and this', (189) h portrayd thial man rnouning natur's
..moral indiffrn.''
1 tims and had hangd littl this wr fort-ndy volution' yt mankinl's ..na-
r, nst Hkl, pulishd on tur within natu'' somhorvworkd gainst th primitiv sltivfors.
nsidr.\n 1867, Huxly rlas- Ultin-latly, th suprmay of sin' naturlism' and agnostiism wr
nously suggstdthat th small what onrnd Huly as a populist. This ws vidnt as his works wr gath.
strlro irds (nd may hav rd. Whras his Siet moirs (fiv volums, 7898_|902) wr dsrip-
onships and promotd o. C. tiv, his nin volums of ssays (789-7894) wr sintillating pros works
tmriantour of 1876. on il. showing th Vitorian ag aommodting itslf to volution. Rading thm,
rowing th inrsing lg and H. L. nkn lld Huxly ..prhaps th gratst virtuoso of plain nglish
e hippus o th tipoing who has vr livd.''
of volution fo gnrations. BIBLIOGRAPHY
|arypalat in roodils.
.!hr Barr, A. P., d' 1997. hr.lmsI7rull',sPl irt Scid Lttrs:
sm rmaindquivoal.
tenrssys.Athns:Univrsit,vof Gorgia Prss.
nusulstps, Huxly in 1859
..volu- Dsmond,^. 1997. Hule: From Deuil's Disipl tl uolutio'sHig Prist'
s at on. is ntry on Rading,A: Addison.IWsly.
mootd natural sltion only Di Grgorio'N4. 1984. T. H, Hul's Pl i Nturl Sie.N'"vvn' T:
it ompltly.uxl rsved Y l U n i v r s i tPr 'r s s .
rral sltion Was liknd to Lyons' S. L. |999. I'homsHr Hl: he uolutilof Sitisf.Amhst,
NY: PromthusBooks.
nd rossing th Brsina River,
Paradis,J. G.' and G. . Willirs,ds.19ti9.uolutild thi<:s.Printrr,NJ:
alon rahing th far bank. PrintonUnivrsityPrss.
d uxly in onrbating soial- Whit,P. 20.Tboms Hul: kig the ,, of Sin'''amridg:
:d volution in lass us h CambridgUnivrsityPrss. -A.D.
d u a t i o nA t ( 1 8 7 0 ) . s t a l i s h -
ington in \871,e fashiond a
..Typs.'' This in-
nt and animal
'ith ntuy. But inulating a
'lutionaydisussions.opning
ontrastd th ryling of th
n's primntal approh.
.tndd,would tt quali-
Industrial mlanism
Industrial mlanism rfrs t th volution of dark ody olors in animal
spis that liv in haitats blaknd y industrial soot. Of pproximatly
1 0 0 r p o r t d a m p l s 'm o s t a r i n s s p i s .a n d o f t h s t h e v a s t m j o r -
ity ar moths' Th most thoroughly doumntd ampl is th ppprd
mot|t, B i ston b etlr i.
Th ommon name dsristh apparan(or phnotyp) of th typial
adult, a pal moth ..ppprd'' with lak-and-whit sals (s figur).
Pal and dark (mlni)f
Mlani phnotyps' namd aronaria, ar ssntiallysolid blak. Intrm-
diats, or insularia, also our ut riv rlativly littl attntion in gnral
summaris. xtnsiv gneti analysis onfirms that th phenotyps ar d-
trmind y multipl allls at a singl gn lous. akgrounds; (2) dirt
Th mlani phnotyp Was first disovrd nar anhstr, ngland, in onto tr trunks; and (3
1848. By 1895' about 98' of th spimnsnar anhstr wr mlani; in pollutd and unpollr
this on rar phnotyp had sprad aross th industrial rgions of Grat tlwll's pionring pt
Britain. With only on gnration pr ar' th narly omplt rvrsal in th yrs. ritiisms a
.!7ll
phnotyp frquny was astonishingly rapid. away from industrial r- usd' th positions on tr
gions, mlani spimns rmind rar. moths wr rlasd,an<
British lpidoptrist J. \/.Tutt advand th following Darwinian xplana- inspird nin additiona
tion in 1896: Ppprd moths ar ativ at night and rst during daylight was a svn-yar study I
hours' hiding on th surfas of trs. ost instivorous irds hunt y day N4ajrusrlasdliving
and loat pry primarily y vision. oths that ar wll onald against thir own 'hiding plas
akgrounds upon whih thy stay motionlss ar mor likly to sap d. propotions to math l
ttion y irds than ar onspiuous moths. Tutt proposd that typial pp- inoulars h mad dir
prd moths in undisturd environmnts gain prottion from visual ppprd moths. By ol
prdators y thir rsmlanto lihns on tr ark. In manufaturing r- usd in ths prdatin
gions whr lihns hav n dstroyd y pollution and th surfas of virtually mathd th s
trs hav n laknd y soot, th typil phnotyps los thir amou- t h m l a n i p h n o t p i
flag and fall vitim to ird prdators' whras th mlani phnotyps in Th wight of vidn l
suh pollutd habitats sapdttion long nough to pass thir gnson to prd moth phnotyps
progny. y xprimnt.
Tutt's idas wr not tstd until th 1950s whn Brnard Kttlwll Baus th rfltar
(197) initiatd a srisof xprimntsto dtrminwhthr or not irds a- ngativly orrlatd w
tually at mlani and typial ppprd moths sltivly. His xprimnts tstal prdition is th
inludd thr main omponnts: (1) quantitativ rankings of onspiuous- s o o t y ' p o I I u t dr g i o n s
nss (to th human y) of typial and mlani phnotypsplad on various ws lrly mt y th
6.t2
]ndustril elnism 65
trv)rdat 83 loiltions aross dlin in mlanism in Amri:rn ppprd moths oinidnt with rdutions
in atmosphri pollution is omplmntd y omparisons of So. onntra-
in ppprd moths is mor tions from southwestrn Virginia, whr mlnism nvr dd 57" ovr
;ulfurdilide (So:) tlran with th sam tim intrvl. Th Mihigan lotion rodd highr onntra-
from point sours than par- tions of So, than th Virginia loation in 23 of th 25 yars that masur-
lt. Although sltionmay op- mntsWr takn at oth plas.
ldividuals mong populations Amian and British ppprd moth populations ar now approhing
rism in pprdmoths' a rl- monororphism for thir rsptivtypial plrnotyps,orrlating with r-
moth spis(.g'' Goodo- dud lvls of atmosphri pollution on oth sids of th Atlanti. Vhil
ny distriutions of mlani orrlationsalon d<lnot stalishausal rlationshios.ommon orrlations
rir rngs.Th porvr of gn suggstommon auss.
nditions rsulting fronr sl- volution is dfind at th oprational lvl as a hang in gn (alll) fr-
- ' r s ,w h o n t r t i n dv a i o u s qutlyin a population ovr tim. Th hangs w hav osvd (and ontinu
,unttr apparnt nomalis in to obsrv evn now) in ppprd moth populations mt that dfinition pr.
4 DNA squns hav now isly. of th four primary volutionary fors-mutation, gnti drift, gn
hri and his ssoiats(Daly flow, and sltion-only strong sltion an ount for suh rapid, dir.
, loi fron-rpopulations aross tional lrangs.Whtlrr o not diffrntial ird prdation of moth Phnotypes
-ds'ngland, indiats high is th sol, proximl agnt of sltion' th ruial vidne for ntur:rl sl-
Lrwork shuld hlp quantify tion in th volution of mlanism is sd on th gnti hangs that hav o-
sm in this spis. urrd in populations. This vidn is indisputabl and wholly indpndnt of
l stablishso_lldsroklss primnts aimd at idntifying partiular olnponnts of fitnss (e'g', am-
4nifiantrdutions in atmos. ouflg from prdtors and physiologial diffrnsamong gnotyps).
: dlind in frquny as th In all aras of ativ rsarh' partiipants routinly hallng assump.
.lfhil
mlanism in th lattr half of tions and intprtations. w ontinu to invstigtth rlativ im-
than th irrrsirr rlarrism portan of gn flow, lihns, prdators, potntil nonvisual sletion' and
unintrruptdnnual rord th havior and ology of this moth spis,industrial mlanism rmains
y yril lark (lark t al. an outstanding ampl of natural sltion asd on th most massiv data
ng a drop in th frqueny of st on reord doumnting long-trm alll frquny hangs in natural
2003 (Grant2005). Tlr most populations.
t a|, 1998) shows markd d-
in Britain. h prditd or- BIBLIOGRAPHY
ion and in th fr1unisof
lark,. A., G. S. irni, and G. Wynn.1985.vlutionin rvrs: lan ir nd
ttions lrs n firnrly sta- th pprdmorh.Billogil.|<trl of th LitlttSoit1'26:1'89_199.
ook, L. . 200. Tlr ris and iIl of the rbrtriform of th ppprdoth.
996) hav doumntd paral- Qurtrl Ruituof Biolog 78 99417.
.lfaltham, .!atts,
h North Amrian pppred Dly, D., K. J. ully, P. . A. Rosin, S. J. Kmp' nd I. J.
Sahri.2004. Trinulotidmirosatllitloi fo th ppprdmoth (Bistrl
. ' n O t y p isn t h s u s p i s -
htulri) . llulrcologNo/,s[doi:10, | | | |li.147|-8286.2004.00607l.x].
1004),ut th ris and sprad Grant,B. S. 1999.Fin runingthppprdmoth paradigm.uolutio53: 980-984.
abotrt 50 yars latef than in 2004. Allli mlanismin Amrin nd British ppprdmoths.lournl lf
lud irny mlani spimns Hrdit95:97_|02.
Ln,ut 1959 th frquny 2005. Industrilmlanism.Ilpedi rlf Lif Sies,www.ls.nt
tio was thn inauguratd in I d o i l:0 . 1 0 3 8 / n p g . l s . 0 05004l1.
Grar-rt,l}.S.,A. D. ook, D. F. Own, and . A' lark.1998.Gogrphid
strn ihigan show that at- tmporalvriationin th inidnof mlanismin ppprdmoth populations
luntly dlind signifiantly. in Amria nd Britain.Jourl of Hrdit 89: 465-471.
'ths had flln tl 20", drop- Grant,IJ.S.' D. F. Own' and ' A. lark. 1996.Para||e| risrrdfall of mlni
:).Th sigrrifianof a shap ppprdmothsin Amrind Britain.llurlof Hrditl 87: 351_357.
656 Isets
Grant' B. S., and L. L. \/isman' 2002. Rnt history of mlanism in Amrian Thr ar striking pat
ppprd moths. Jourl of redit 93: 86_90. sity. To appriat ths
Kttlwll, H. B. D. \973' The uolutio of lnism: he Stud of Rurrig
txt. Insts,along witl
Ncessit. Oford: larndon Prss.
th four major groups t
ajrus, . . N. 1998. lism: uolution i tio. Oford: oxford Univrsity
Prss. a r t h s g m n t d n i n
27. The ppprd moth: Th proof of Darwinian volution. http://www alld suphyla, ar th.
.gn.am.a.uk/Rsarh/ajrus/SwdnPpprdmoth207.ppt;http://www tivs), and th rusta
.gn.am.a.uk/Rsarh/M jrus/Swdnta|k220 87.pdf . many othrs). Th inst
Tutt, J. W. 1896. British oths. London: Gorg Routldg and Sons.
th suphylum thy ol
-B.S.G.
suphyla has n quit
varlet of hpods ar
Th haratristi fa
Insts
organizd into thr ma
It is diffiult to argu that volution has produd any singl group mor lgs mrging from th r
sptaular than insts. Thir numrs alon ar staggring:ovr 820,000 nd outr shll that is r
spis hav en dsrid, and stimats for th atual numr of spis od; and a varity of d
xd 2 million. jor groups of insts ar still ing disov ered.|n 202, sts hv no lungs, r
for xampl' an ntirly nw ordr of mantidlik insts was dsrid, and th body that opn at tt
hundrds of nw spis ar dsrid from th tropis vry yar. molting' shdding thrr
Th divrsity of insts is no lss staggring.Insts oupy vry haital panding into a nw skin
rgion on arth. Thy hav invadd vry haitat pt th open sa, and thy On of th striking pl
hav radiatd into narly vry onival fding nih. any spis uild th 31 tant ordrs ar(
shltrs nd nsts, ranging from th asings of sand and dbris onstrutd by an found in vry ol
addis fly larva to the intrit mound and tunnl systms of tropial trmit vry othr major anim;
olonis. Whil th majority of insts lad solitary livs, thr is a sptrum of fossils or in whih ord
soil organization that inluds family aggrgationsin atrpillars, small fam- springtail that is stima
ily groups of wood roahs, olonial nsting in wasps, and highly rfind, om- mian, aout 250 millior
pliatd soitisof silings that form ,wasp' ant, and trmit olonis. Th and fossils of thm ar '
ants alon inlud spis that ar savngrs,prdators, and gardnrs;som ordrs that had not ap[
ant spis tnd aphids for thir sugary srtions and othrs ptur ants of and mammals, whih at
othr spis and nslav thm as workrs in thir own olonis. mals in th fossil rord
Insts provid som of natur's most sptaular phnomna and display olaanths, Ltimeri s
som of natur'S most sophistiatd faturs.Th ilial plagu of lousts r- ognizabl mmbrs of th
flts a sptaular natural ourrn' th ruption and organizd movmnt this standrd' w might
of millions of individuals of a speis namd Shistocr grgri. Priodial Th only inst ordr
idas mrg in th forsts of astrn North Amria y th millions on 13- dityoptra, whos m
and 17-ear yls, only to mat' rprodu, and di within wks. Monarh Unlik dragonflis' th
uttrflis mak sasonal migrations aross hundrds of mils twn North C r o n i f r o u s ( a l u rt
Amria and Mio. Many insts appar to dfy iologial onvntions: othr insts,and disap1
thr ar spis that onsist only of fmls, and many spis an produ of th Plaodityoptr
many individuals, from dozns to hundrds, from a singl gg. Instshav so- normously sussful,
phistiatd snsory systms' sing in wavelngths that humns annot' suh sts from thos priods
as th ultraviolt, nd haring at frqunis tht humans annot. Insts ar disappardso suddnl
not asy to poison; thy hav th apaity to assimilat a varity f toxi mol- !hil narly all of th
uls, somtims raking thm into harmlss omponnts and somtims hav n four prids l
squstringthm as dfnsiv ompounds that dtr prdators. dramatially. In th first
lnsets 657
.y of mlarrismin Amrian Thr ar striking pttrns in th organization and history of inset divr.
s i 1 'T. o p p r i a t h s p a t t n s i. t i s i m p o r t a n t f i r s t o p I i n s t si n l n .
m: Tb Stud of Rurrig txt' Insts' along with rnyriapods (ntipedsnd nrillipeds),ar on of
t four major groups that ompris th arthropod phylum. Th arthropods
fio. oxford: Oford Univrsity ar th sgmntd aninrals, and th othr major arthropod groups, oftn
.wirrianvolutio.http://www alld suphyla, ar th triloits, th hlirats (spidrs and thir rl-
rdmoth2007.ppt; http://www tivs),and th rustaans(a divrs group that iniuds ras, llstrs,and
:0807.df. many othrs).Th insts ar a distint lass' th Hxapoda (six lggd)'within
.Whil
lutldgad Sons. t suphylum thy oupy with th myriapods. ah of th arthropod
-B.s.G.
suphylahas n quit sussfulin numr of speis,th prolifration and
varity of hxapods ar approahd only by th rustaans.
Th haratristi faturs of insets inlud having thir sgmntd odis
orgaIrizdinto tr majo divisions (had,thorax' admn);thr pairs of
lgs mrging irom th ntral division (th thorax); an xoskiton (a hard-
lud any singl group mor
ar staggring:ovr 820,000 nd outr shll that is not lways so hard, as in atrpillas);a vntral nrv
t h a r u ln u m r o f s p i s ord; and a varity of distint morphologial nd physiologial faturs.In-
till bing disovrd.|n 22, stshav no lungs, rathing instad through spherils,whih ar tus in
ik instswas dsribd, and th ody that open at th ody surf.Lik all arthopods, instsgrow by
molting' shdding thir trnl skin and xosklton priodilly and -
r trpis vry yar.
panding into nw skin and xoskieton.
Instsoupy vry haital
,t xpt th opn sa, and thy on of th striking pattrns in th divrsity of insts is that narly all f
th 1 tant ordrs ar old; individuals that rprsntmost of ths ordrs
Jing nih. any spis uild
an b found in vry old fossil dposits. This is quit diffrnt from narly
;and and dris onstruted by
tnl sysrmsof tropial trmit vry othr major animal group, in whih many ordrs ar known only from
fossils or in whih ordrs rang widly in ag. Th arlist inst fossil is a
ay livs,thris a sptrum of
tions in atrpillrs' small fam- springtail that is stimatdto e 96 million yas old. By th nd of th Pr-
mian, aout 250 million years ago' most of th knwn ordrs had appard
asps,and highly rfind, om-
r,ant, and trmit olonis. Th and fossils of thm are widspred from that point onward. Th only major
rdators,nd gardnrs; som ordrs that had not apprd y th Prmin ar th toparasitsof irds
and mammals, whih appard nly aftr th apparnof irds and mam-
ons and othrs aptur ants of
mals in th fossil rord. o alirat ths osrvations'rmmbr that th
ir own olonis.
olaanths,Ltimri spp.' arc alld ..living fossils'' bausthr ar r-
rular phnomna and display
h bilial plagu of lousts r- ognizalmmrs of this group psrvdas 380 million-yar.old fossils.Bv
this standard, w might all most inst ordrs living fossils.
tion and oganizd movemnt
Th only inst order tht has gon xtint was a goup alld th Palo-
:histlr gr gri. Priodial
ditylptra, whos mmrs rsmld drgnflis in thir apparan.
\mria y th millions on 13-
nd di within wks. onarh Unlik dragonflis, ths insts wr hrbivrous' rrging in th mid-
aronifrous (aout 30 million yrs go), not muh latr than man
rdrds of mils twn Nrth
othr insts,and disapparingat th nd of th Prmian. Th striking aspt
d f y i o l o g i a l o n v n t i o n s :
of th Palaodityopter'sdisppaanis that thy appr to hav n
ad mny spis an produ
,m a singl gg. Insts hav so- normously sussful,ornprising aout 50% of th known spis of in.
stsfrom thos priods. Rsarhrshav no ida why s sussfula goup
l r h s t h . l th u m a n s a n n t ' s u h
lat humans annot. Insts ar disappardso suddnly.
.shil
narly all of th ordrs and many f th familis ar vry old' thr
similat a vait of toxi mol-
;s omponnrsand somtims hav bn four priods whn th divrsity of spis within familis inasd
dramatially. In th fist of thspriods, th rly aronifrous (abotrt380
i dtr prdators.
658 Ists
million yrs ago), wingd insts appard and gan to multiply. ln th s- ago), nd th arlistnl
ond, toward th nd of th Prmian, th arlist holomtaolous insts ap- t y o p t r i d a 'w h i h p p
pard, and this group gan divrsifying soon aftrward. Holomtaolous sil plant lvs from th
insts hv a omplx lif yl of gg, larva, pupa, and adult; th larva and damag from hrivors
adult ar dramatially diffrnt in apparan and lifstyl, and thy rpr- t h m a j o r r d i a r i o n so f J
snt almost two distint organisms. Holomtaolous insts inlud th famil- major radiations of flow
ir groups of flis, uttrflis,tls' ants' nd lawings. In ontrast, th This jint radiation
insts w all hmimtaolous, whih appar in th fossil rord for plants volve dfnssa1
holomtaolous insts, hav a simplr lif yl in whih th organism that om thos dfnss,an
mrgsfrom th gg, alld nymph, rsmlsa simplr and smallr vrsion mrge. Th vidn f
of th dult. ommon hmimtalous insts inlud grasshopprs, rikts, al studis. Instsat a
gnuin ugs lik stinkugs, and roahs. Holomtabolous insts hav n vidul platstht hav
normously sussful,rprsnting aout 9Yo of isting inst spis. hrivors or highr o:
Th third priod of inst xpansion ourrd in th rtaous,frm 130 al to surviv, grow lar
to 65 million yars ago, whn aout half of th modrn familis appard fnss at as agnts of t
and inst spisdivrsity inrasddramtially. This pansion gan im- om dfnsive strutur
mdiatly aftr th initial radiation of flowring plants. Th fourth priod of As a plant spisrfin
pansion, whih som sholars onsidr as a ontinuation of th rtaous tion, only spifi grou
radiations, was in th Trtiary, th priod that gan 6 million yars ago. through volution. As al
Th ontinud radiation of insts with phytophagous hits, maning that a s p i so r g n u s o f p l
thy fd on plants in one Way or anothr' was rtainly a ontinuation of th that mploy diffrnt <
rtaous pansion. As witnss to this, aoot 9 5" of th rnt Lpidoptr- p l n t s t h t p r o d u n w
ans (buttrflis,moths, and skipprs),narly all of whih ar phytophagous' p o p u l a r i o n s .o f t n i n
mrgd in th Trtiary. But th Trtiry radiation also inludd th mr- slvs ttling with diff
gn of th diddly nonphytophagous ants, s, wasps' and highr tr- twn ths plant pop
mits with thir sophistiatd soial systms as wll as th ordrs of li, pross of spiationha
whih ar toparasitsof irds and mammls. ra.
wo tis ind th volutionary historis of insts and flowring plants to Through this pross
ah othr. First, th ovrwhlming majority of flowring plants ar polli. othr riproally towar
natd y insts, and inst pollination srvi hs promotd divrsifiation rsult is that most plant
and spiation. Instshav n ating polln for quit som tim. Polln is phytophagous insts ar
rognizal in th guts of som fossil insets in th Prmian' ovr 250 mil- soiations t broad lv
lion yars ag. Th striking hang that oinidd with th mrgn of lis ar found only on
flowring plants was th apparan of spializd faturs of th had and finr lvls. onarh
mouth that failitatd not th ating of polln ut the transport of polln as a n d p i p v i n s w a l l o w t a
th insts fd on ntr. Ths first appar in som flis in th arly rta- tolohi); in ah s
ous. By th lattr half of th rtaous' 90 million yars ago' most sp- narly all oth insts.
ializd pollination systmsknown tday had volvd and ar larly sn in Th study of insts
th fossil ord. And should on doubt that inst pollination has on. volutionary iology. Th
triutd to th suss of flowring plants, it is worth rmmring that rgardd as on of th ;
groups of plants pollinatd by animals hav divrsifid at twi th rat of natural sltion, and sl
thos groups that ar pollinatd y wind or watr. anon of arly xprim
Sond, phytophagous inst divrsity has followd th divrsifiation of th fruit fly (or, mor
flowring plants. To sur, many insts hav n fding on plants for Thoms Hunt Mogn
most of thir xistn.Fossil plant tissu shows vidnof ing pird y ism that has n usd i
mits and ollmolans sin th lat Dvonian (aout 400 million yars tion and spiation. Th
Ists 659
J
thory of kin sltion.Kin sltion now undrlis muh of our undrstanding
of soiality and many pattrns of havioral volution throughut th animal
world. Studis of individual vriations in moth and uttrfly wing pattrns y
. B. Ford and his ollagus Wr on of th foundations of ologial gn-
tis, th xprimntal study of natural sltion and volutionary hang out-
sid of th laoatory.
Mor rntly, many diffrnt inst groups hav n usd as modl sys.
tms for volutionary studis. Th flour tls, Tribolium spp.' hav n
Jalonski, David (
usd in pathraking studis of group sltion. Studis of Watr stridrs
David Jlonski is amon
hav illuminatd th onflit twn th ss ovr mating, and how that
gists who, hving ntr
onflit has shapd th physiology nd havior of th ss, ttr than any
othr systm studid to dat. Th dmslfly spisthat sgrgaty typ of that took pl in th lat
larg-salpttrns nd
lk (with or without fish) hav n usd to illuminat how rapidly a ]roup
nring palontologists
n divrsify in novl haitats and how adapting to a novl ologial ir-
S t v nS r a n l y .J a l o n s k
umstan oftn oms with th ost of losing th aility to prsist in th
tintions and thir rsr
original onditions.
has approahd this su
|LIoGRAPY sis, whih uss omputr
Brrnaum, . R. 1995.Bgsitt thSstm:lstsd Thir Imptrl um of data drawn from th f
ffirs. Rading,A: Addison-\/sly. i n t i o n si n t h g o l o g i ;
isnr,T. 200. For Lluof Ists.amridg,A: BlknpPrssof Hrv:rrd undrstanding of th mo.
UnivrsitvPrss. iodivrsity risis.
Grimaldi, D., and . S. ngl. 205. uolutio of tb Ists.Nw York:
o n o f J a l o n s k i .ps r i
mridg Urrivrsity Prss. - l I
661
662 Jeki
Prhaps Jalonski's most signifiant rsarh involvd studying th dynam- rviw is oftn suppos
is and pattrns of tintions. In this ara, Jalonski has ontinud to tst of hrdity, suh as
volutionary hypothsswith quantitativ analysis. On qustion h has ad- om a plausil thor
drssdis whthr survivorship during mass xtintions is pditalor ran- rviw, othr sltionr
dom, and whthr th volutionary trnds that follow major tintions populations, not indivr
display a uniform pattrn. A rlatd study askd whthr th survivors of nism of hrdity.
major tintion vnts (for ampl, th mass tintion that killd th di. Jnkin was orn in I
nosaurs 65 million yars ago) r in fat sussfulvolutionry omptitors family movd to Italy
aftrward. H found a numr of possil routs gnra may follow aftr Gnoa in 1851. H t
suh an vnt, ranging from thriving pansion and divrsity to stgnarion dsign and mnufatur
and vntual xtintion. H onludd tht whil thr wr som pr- h i m s l u p a s o n s u l t
dital fators involvd, th paths appar to vary asd on rgional nvi- som with his frind \
ronmntal diffrns, an osrvation that supports th hypothsis of Jnkin wrot ports fo
randomnss. n's ommitt on lt
Surpisingly, Jalonski onfirmd thr was oftn littl orrlation b- was aptdas an as
twn tintion survival and posttintion volutionary suss.In many Jnkin had wid si
ass, formrly struggling gnra thrivd, whil dominant organisms Wr him to ommnt on D
..Dad lad Walking,'' dsrisgn-
wipd out. A third altrnativ,alld rviw, pulishd in 1
ra that wr not immdiatly killd off ut wr doomd to vntual d- arth must hav old
..rippls'' tintion oftn follow millions
lin, suggstingthat sondary of dud th amount of p
of yars aft mass tintion vnts. ovrall, ths studis and othr work H also rpatd th tr
hav stalishdJalonski at th forfront of fforts to us nw mthodolo- r s t r i t dw i t h i n d f i n
gis (and oftn th latst thnology) to answr many of th qustions sltion ould aum
out th auray and usfulnssof th fossil rord that hav vxd pl- variation was a distin
ontologists sin harls Drwin's day. In gnral, Jalonski's work sup- would happn to th n(
ports th onvition that th fossil rord is rlial and thus an usd to sumption that hrdit
infr th rlativ importn of diffrnt vlutinary mhanisms pattrn- parnts. H ntd that
ing th history of lif on arth. By ttr undrstanding th dynamis of x- onntratd within a
tintion' Jalonski has also highlightd th ways xtintions, whih r ft. This had happn
normally sn as ngativ prosss' ontriut to th ovrall dynamis of th farmr had rd or
volution. isti. In a wild popula
with unhangd mmb
|BLIoGRAPHY
halvd in ah gn
Jalonski,D. 1999.Th futurof th fossilrord.Siece284 21,14-2116. Darwin had nvr tl
2001. Lssonsfrom th past:volutilnary impatsof massxtintions.
think of thm as indrv
l,rrledigsrlf th Ntirlnl Acdm lf SiesUS 98: 5393_.5398.
2004. tintin: Pst and prsnt.Ntr 427: 589. -D.Se. Jnkin,s rviw, modif
prosss Wr at work
pointd out' howvr' t
diffrns within a p
Jnkin,Hnry CharlsFlming(1833_1885) favord individuals ar
Hnry harls Flming (pronound Flming) Jenkin made major ontri-
butions to physis nd ltrial nginring in mid-nintnth-nturyn- BIBLIOGRAPHY
gland. Rmmrd y volutionists as th author of a rviw of harls
Bowlr,P. 2003. uolut
Darwin's o th rigi of Spies' h agud tht natural sltion would aliforniaPrss.
not wlrk if hrdity Ws a pross that lndd prntal haratristis. This D w i n , . l l 8 5 9 ] l 8 " 2 .
Jnkin 66
involvd studying th dynam- rviw is oftn supposd to hav shown that th mov to a partiulat modl
lonski has ontinud to tst of hrdity' suh as ndlirr gntis'was ssntilif sltion was to -
ysis. on qustion h has ad- om a plausil thory. In fat' although Drwin was imprssd y Jnkin's
:intionsis prdital or fan- rviw, othr sltionists ralizd that if variation Ws sn as a proprty of
lat follow major xtintions populations, not individuals, sltion was plausil whatvr th mha-
kd whthr th survivors of nism of hrdity.
i xtintion that killd th di- Jnkin was orn in Dungrnss,Knt, ngland, on arh 25, 1833. His
' s f u l v lu t i o n a r y o m p t i t o r s family movd to Italy and Jnkin rivd an MA from th Univrsity of
uts gnra may follow after Gnoa in l85]. rturnd to ngland nd spnt 10 yars working on th
' n n d d i v r s i r yt l s t a g n a t i o n dsign and manufatur of sumarirr tlgraph als, aftr whih h st
whil thr wr som pr_ himslf up s a onsulting nginr in London. H took out numrous patnts'
.sillianr
vary basd on rgional nvi- som with his frind Thomson (latr Lord Klvin). In th 1860s,
supPots th hypothsis of Jnkin wot rports for th Bitish Assoiation for th Advanmnt of Si-
n'sOmmitt on ltrial Stndards, s a onsqun of whih th ohm
ls oftn littl orrlation b- was aptdas an asolut standard of rsistan.
:volutionrysuss.In many Jnkin had wid sintifi intrsts,nd th Nort Britis Reui, asked,
il dminant organisms Wr him to ommnt on Darwin,s thory fom a matlrmatial viwpoint. In his
lad !7alking''' dsris gn- rviw, pulishd tn 1867, Jnkin ndorsd homson's argumnt that th
wr doomd to vntal d. arth must hav oold down too rapidly for natural sltion to hav pro-
(tintionoftn follow millions dud th amount of progrssiv volution osrvd in th history of lif.
thsstudis and othr work H also rpatd th traditional viw that within ah spies,variation is
ffortsto us nW mthodolo. rstritd within dfinit limits. or originally, h ana|yzed how natural
sW mJny of th qustions sltion ould umulat favoral variations. Arguing that ah suh
l rord that hav vd pal- variatiorr Was a distint salation, or sport of natur, h xplored what
nral,Jablonski's work sup- would happn to th nW hratr as rh individual rprodud, on th as-
lial and thus an b usd t sumptiol1 that hrdity blnds th haratristis of th mal nd fmal
Jtiony mhanisms pattrn- parnts. H notd that in artifiial sltion, suh nw haratrs ould
rstanding th dynamis of x_ onntratd within a small popultiorr and ould thus hav signifiant f-
ways xtintions, whih ar ft. This hd hppnd in th as of th shot-lggdAnon shp, whr
lt to th ovrall dynamis of th farmr had rd only from th singl ram orn with th nw haatr.
isti. In a wild population, lrowvr, th favord individuals would rd
with unhangd mmrs of th popultion, and th hatristi would
halvd in h gnration until its fft was too small to ount'
Sci284:.21|4_2776. Darwin had nvr thought of variations s larg-salsports, ut h did
impatsof massxtintions.
think of thm as individual haratr hangs. was thus imprssd y
: e sU S 9 8 : 5 9 3 - 5 3 9 8 .
427:589. -D.Se. Jnkin's rviw, modifying ltr ditions of his b<lokto mphsizthat othr
prosssWr t work. othr Darwinists, inluding Alfrd Russl !7alla,
pointd out, howvr, that if on thought of variation as a rang of haratr
diffrns within a populatiorr, Jnkin's argumnt was infftiv baus
3-188s)
favord individuals ar oitiful rathr than r.
g) Jnkin mad major onti-
n mid-nintnth-nturyn- BIBLIoGRAPY
Luthorof a rviw of hals
Bowlr,P'200 . t , o l u t i o : h i s t o r lnf l d . 3 r d d . B r k l y : U n i v r s i t y o f
l that natural sltion would California Prss.
d parntalharatristis.This Darwin,. [1859l 1872.o th rigi of Specis.6th d.London:John Murray.
664 Jnki
Hull, D. L', ed. 197. Dr,i d His ritis' amridg' A: rvard Univrsity
Prss.
Jnkin, F.1867. Th origin of spis.North British Ruie 46:277-3I8.
-h
Rus, . 1,999. Duii Rurlltio: Si Rd i Tllthd l.2nd d.
hiag: Univrsity of hiago Prss.
-h
Vorzimmr, P. J. 1970. hrles Dnui: Yrs of otrlurs. Philadlphia:
mpl Univrsity Prss. -P.J'B'
Kttlwll, H. B. f
Hnry Brnard David l
atd group oftn rfr
ntis' whih inludd s
S h p p a r d .W h i l r h r
markd y a ommn
populations, and, in pa
ural sltion as th m
was a giftd naturalist \
oyhood lov of ntor
s a r h i n . B . F o d . s
Kttlwll is st kn
mlanism, whih rfrs
in many moth spisin
of th fisr larg-salar
ngland and ontinnt
trnd ourrd in th p'
spiswas fist disov
50 yars it m so .
1 8 9 1 i s t h l s s i n i n
Kttlwll am fa
arly l950s doumnt
ommon. In pollutd a
s o o t - d a r k n dt r t r u I
mnt ondutd in th .
m k dp a l a n d d r k
tmptd to aptur as n
hind his invsrigti
for th two forms shot
Was at an advantag'o
p th rapturrt
vivd during th intrv
what Kttlwll found.
unpollutd wood in Do
ln this wood h ws a|
t I w l ls u p p l m n t d tI
amridg,NtA; arvard Univrsity
lrs of otror:ersy.Plriladihi:
-P.l.B.
Kttlwll,H.B. D. (1907_1979)
nry Brnard David Kttlwll ws a founding mmr of a loosly assoi-
atd goup oftn rfrd to s . B. Ford's Oxford Shooi of ologial G.
ntis,whih inludd suh luminris as A. J. ain, yril lark, and Philip
Shppard. !7hil thr was nvr any formal strutur' th mmrs wr
markd y a onrmon appoah to the study of gnti variatin in natural
populirtions,and, in partiular, y a ommitmnt to th prvasivrol of nat-
ural sltionas th most important agnt of volutionary hang. Kttlwll
was a giftd nturalist who lft mdial prati at th ag of 41 to pursu his
oyhood lov of ntomology. H gan working full tim as a snio r-
srhrin ' B. Ford's nwly foundd suunit of gntisat oxford in 1951'
Kttlwll is st known for his rsarh on th phnomnon of industrial
mlanism, whih rfrs to a rapid inras in th frquny of rar drk forms
in mny moth spisin th viinity of mantrfaturingntrs as a osqn
of th first larg-salair pollution assoiatdwith th Industrial Rvolution in
ngland nd Continntal urop. Th first and most famous ampl of this
trnd ourrd in th ppprd mot|7,isto betulri' Th dark trm in this
spiswas fist disovrd nar Manhstr in 1848. .$ithinth sp of only
50 yars it am so ommon that th pal frm was loally xtint. (Tutt
1891 is th lassi nintnth-nturydisussion of th topi.)
Kttlwll am famous f a sris of fild primnts ondutd in th
arly 1950s doumnting th rason why th dark forrn was oming mof
omlnon. In pollutd aras, dk oloration prottd th moths, rsting on
soot-darknd tr trunks, against ird prdation. In n lgant fild xpri-
mnt ondutd in th su1mrof 195, Klwll rlasdlarg nrrmrsof
markd pal and dark moths into a pollutd wood nar Birmingham and at-
tmptdto aptur as many as possil ovr th nt svral nights. Th logi
hind his invstigationwas that, all things ing qual' th raptur rats
for th two forms should th sam. If' on th othr hand, th dark form
was at an advirntag'owing to its rlativ inonspttlousnss'on would -
pt th raptur rt for it to highr aus mor of th drk form sur-
vivd during th intrvl twn rlas and rapture. And this is prisly
what Kttlwll found. Kttlwll ondud ompanion xprimnt in an
unpollutdwood in Dorst in th summr of 19S5 whr h found th rvrs.
In this wood h was a|to raptur mlr of th pal form thn th dark. Kt.
tlwll supplmntd ths primnts with a film rod doumnting that
66.t
666 Kimur
whn irds wr givn a hoi, thy passd ovr inonspiuous moths in DNA. DNA onsistsl
favor of th moth that was mor onspiuous in its akground rsting sit. sin, guanin, and thy
(S Kttlwll 1955 and 1956 for th original rports and Kttlwe|\ 197 of a fw thousand as
for a summation and ovrall disussion.) qun of ass in tha
Whil ttooks oftn portray Kttlwll's invstigations as unprolmati' th DNA has no knov
rsarhers (inluding Kttlwll himslf) rogniz that th initial invstiga- A mutation is a prma
tions involvd numrous assumptions tht ar opn o qusrion (.g.,th rSting to a T. If th hang l
sit of moths). This has ld thos ritial of volutionary thory to laim that th fft on th individua
phnomnon is no longr an xampl of natural sltion, and furthr that osrval fft. If a l
thre is a onspiray y ttbook writrs to hid this fat. (Hoopr 2002 js a frquny in th popuI
.Wlls
snsationalisti aount of th work y a journalist and 2000 is a svr' nti drift.
rligiously motivatd ritiqu by a iologist who is a followr of th Rvrnd ln t968 Kimura put
Sun Myung Moon, foundr of th Unifiation hurh.) At last ight studis di- ulk of hangs in th
rtly addrss privd problms in Kttlwll's initial invstigation.Th stud- random drift. Kimura
is hav indpndntly onfirmd that ird prdation is th most important th population and ulti
fator in th sprad of th dark form. Prhaps th most onvining vidn rsult of natural sl
that it is an ampl of naturl sltion' howver, oms from th prditd ssnof Kimur's id
rapid dlin of th dark form sin th introdution of ..lan Air'' lgislation ory of volution; Kimu
in th Unitd Kingdom and lswhr. (Majrus 1998 givs a full ovrviw of ovrd indpndntly
modrn rsarhand Rudg 2002,2005 addrssth lims of frud.) most of th susqu
narily attriutd to hin
|BLIoGRAPY
It oftn happns in t
Hoopr, J.2002. f oths d n: Th Untold Stlrlf Sicd th Ppprd its own sak' turns ou
of. Nw York: Norton. stantial ody of thor
Kttlwll,. B. D. 1955.Sltionprimnts on industrialmlanismir-rth
was fortunat that th
lpidoptra.rdit 9: 23_342.
1956.Furthrsltionprimnts on industrilmlanismin th nturis latr, ut wi
lpidoptra.Ilredit 10: 287-301. at studnt, Kimura s
1973. Th uolutilof lism,Oford: larndon. d o m p r o s s s ,s u h
Majrus,M. . N. |998. elism:uolutili Atio.Oford:Oford Univrsity inras in th popula
Prss. gnrations it would t
Rudg,D. !. 2002. rypti dsignson th ppprdmoth. Itrtiol lourl of individuals that woul
Tropil Biolog d oserutio(Ruistd Bklklgi Tropil).50,no. 1: 1-7.
2005. Did Kttlwllommit fraud? R-xaminingth idn'Pbli Kimura l'vorkd out t,
Udrstndig lfSi14, no. 3:249-268. ial, or dltrious.Su
Tutt, J. !. 1891. lism d elohrlismi British Lpidlptr.London: in gologial tims.
Swan Sonnnshin. It turns out tht th
.!lis'
J. 2.los of uolutio:Sior th?.Washington,D: Rgnry. for anothr is simply
-D.vR.
rasonaly wll know
an hkd agains
squnsof this dis
Kimura, otoo (|924_994) w know th mutatiol
Motoo Kimura is st known for th ida that most volution t th DNA tion. For studis of
lvl ours y random prosses'rthr than y Darwinin ntural sl- lok, whih depnds'
tion. Prior to this, volutionists had attriutd ssntiallyall of volution to It is now gnrallyt
natural sltion. Kimura argud tht for most volutionary hangs' han mals and othr vrt
is mor important. is still ontrovrsial, h
Within a ll, a hartristinumr of hromosoms an found (for It is lar that in som
xampl, 46 in humans). h hromosom is a long, tightly oild thrad of ltion. Th answrsv
Kimur 667
669
670 Lck
..gntidrift'' modls pro-
was shown how his osrvations fit wll into th adaptd to diffrnt hl
posd in th 1930s y Swll Wight and fashionalin Amri. A thnial vrgn) Lak suggs
monograph (Lak 1945, writtn in 194) dtaild his onlusions drawn tan; thy offrd a s
from this influn. wild. Inspird y this
Lk rvrsd his viws shortly aftr rturning to nglnd and to Huxly's (1944), and of th or
influn. ..Unptdly, a ronsidrtion of th original matrial ld to Dri's Fihs
markd hang in viwpoint rgarding omptition twn spis and th natural sltion into
ak diffrns twn th finhs'' (Lak 1947' 1).|n Dr,i's Fihs whn volutionists w
and many popular artils thraft, Lk mad two points aout natural ..artifiial'' (laorato
sltion in Gal6pagos finhs. prinipl of omptit
..
First, h onntd finh aks to dit, arguing tht thy wr wll Gaus in th 190s.
adaptd to spifi fding styls: larg sds, small sds, insts, and fruit nglish fforts to d
(s figur). Natural sltion within ah spis rinford spializatin . B. Ford nd rinfor
and ratd highly adaptd forms. Sond, Lak fousd on ass whr sv- land snail, p, a
ral spisoupid idntial ologis in th sam loation. In ths ass, Bisto. Although th
..Sin
h said, natural sltionwas spiallyintns: th han of thir - Rort Bowman and
ing qually wll adaptd is ngligil, on of thm should liminat th othr sity f omptition' i
ompltly'' (Lack 1947,62)' n small islands, Lak found only on spis undr way y Ptr
of ground finh, suggsting that omptition ld on spis to dominat and onlusions.
othrs to loal xtintion. on islands whr two or mor spiswr found, Io 1945, Lak b
Lak rlizd that ths spis W nvr found with atly th sam nithology' part of th
ologis.Natural sltionws ausing spisto highlight thir diffrns, institut had two go
thry rduing omptition. This ausd divrgn, as diffrnt spis data and ovrsaw n.
Trust for Ornitholog;
al ommunity. S
ornithology. Lak fo
lation of population I
ontrolld populatior
ators. H fousd t
took pla. Lak's th
..
dwards and thr
i Birds (1'966).Hts 1
sizd ologia1isola
Not a strng admi
in th dirtor's hait
too narrowly. Lak'r
his waknsssas ar
stdnts.
Lak onvrtd to
th Anglian Churh
uolutior hor1
and volution ould
han prosss's
Figur 1. ampls of Darwin's finhs.Not t vry diffrntak shaps, natural, not vil. v
adaptd to diffntfeeding stuffs-ig strong aks for hard nuts and th lik, squnt history. Th
fin aksfor smll sdsand th lik. an of th first h
Lk 671
h ..gntidrift'' modls pro_ adaptd to diffrnt haitts. hanging dits was th usual avnu for this di-
lnalin Amria. A thnial vrgn' Lak suggstd.In this intrprtation, aks took on spial impor-
ltaild his onlusions drawn tan; thy offrd a sptaular, nuand ampl of natural sltion in th
wild. Inspird y this suss'Lak mad similar studis of passrine irds
rg to ngland and to Huxly's (1944), and of th ormorant and th shag (1945).
. th original matrial ld to a
Druin's Fibs mad Lak famous. H had transformd a as against
ition twnspis and th natural sltion int a prmir xampl for it. His rsrh am at a tim
947, vt). | Dri's Fibs ..natural'' rathr than
whn volutionists wr alling for mor studies of
d two points aout natural ..artifiial'' (laoratory) xampls of sltion.It did th sam for ology's
prinipl of omptitiv xlusion' mad thortially important y G. F.
arguing that thy wr wll ..Lak's finhs''' many ltr alld thm, also oostd
Gaus in th 1930s. as
small sds,insts' and fruit nglish fforts to dvlop volutionary olgy, hampiond y oxford's
is rinford spilization . B. Ford and rinford y Arthur ain and Philip Shppard'sstudisof th
:k fousdon asswhr sv- land snil, p, and Brnad Kttlwll's work on th ppprd moth'
sam loation. In ths ass, Biston, Although th finh as was ritiizd in th 1950s' spially y
..Sinth han of thir -
: Rort Bowman and Danil Simrloff, for ovrstating th tnt and intn-
rm should liminat th othr sity of omptition, it nvr lost its allur. By th 1970s, long-trm studis
;' Lak fundonIy on spis undr way y Ptr and Rosmary Grant onfirmd and tndd Lak,s
,d on spisto dominat and onlusions.
lo or mor spiswr found, In 1945, Lak am dirtor of th dward Gry Institut of Fild or-
found with xatly the sam nithology, part of the Dpartment of Zoologial Fild Studis at Oxford. Th
s to highlight thir diffrns' institut had two goals. First, it was a ntr for hang of ornithologial
ivgn.as diffnt spis data nd ovrsaw nationl nsus work in ollaoration with th British
Trust for Ornithology. This put Lak in a ntral position in th ornithologi-
al ommunity. Sond' th institut sponsord original rsarh in fild
ornithology. Lak fousd this rsarhon population ology and th rgu-
lation of population numrs (Lak 1954). H argud that only thr fators
ontrolld population sizs:disas,food availaility' and parasits or pred.
ators. H fousd th institut's rsarh on prisly how this rgulation
took pla. Lak's thoris w svrly ritiizd, spially by V. . Wynn-
..group
dwards and othr sltionists.''H plid wtth Plpultio Stdis
i Birds (1966). His final ook' ologil Isoltil i Birds (1971'),mpha-
sizd ologil isolation as th driving for of divrsity.
Not a strong administrator, Lak Was mor omfortl in th fild than
in th dirtor's hair. H was ritiizd for fusing th institut's attntion
too narrowly. Lak's strngths as n ornithologist, howvr, far dd
his waknsssas an dministrator. H was inspiring and mntord many
studnts.
.!Var
Lak onvrtd to hristianity aftr \World II. H was onfirmd in
th Anglian hurh in 1951, and h rmaind dvout throughout his lif. In
uolutionr Thor d hristi Belif (1957), Lak argud tht sin
and volution ould hlp to plain mny unrsolvd thms in rligion.
han prosss' suh as natural sltion, ould rativ. Dath was
vry diffrntak shaps, natural' not vil. volution did not spak to th origin of lif, only to its su-
..Th tru signifi-
ksfor hard nuts and th lik, squnt history. Th Bil and ntur did not ontradit:
an of th first haptr of Gnsis is to assrt that God mad th univrs
672 Lmrck
and all in it, that H saw that it was good, and that plad man in a sp-
Lamark initially ml
ial rlationship to Himslf'' (Lack 1957,34). Lak also sought to tterun-
te 177s with th ir
drstand aras of gnuin onflit. For instn'many asptsof man's moral
in Paris. His rr l
bing smdto him to unplaind y thorisaout th moral volution
was ronstitutd as th
of animals.
on of th profssors
Lak's ..y Lif as an Amatur ornithologist,, (797| is an autoiography
was givn instad th 1
to 1,945, writtn whn h am dirtor of th dward Gry Institut. For
..amtur''to ..profssional.'' mals.'' H soon hara
him this markd an important transition from
mals without vrtr
!. . Thorp (1974) provids th kind of knowldgal mmorial only a
initially to what h kn
los frind ould podu. Lak was awardd th Godman-Slvin Gold
am ovr th nt twr
dal of th British Ornithologists Union in 1958 and th Darwin Mdal of
zoology.
th Royal Soity i 1972.
lassifying th inv
BlLloGRAPHY thm wr not Lamar
onward, h had spu
Grant, P., and R. Grant. \986. olog d ullutilof Dri's Fihs. pls of physis, hmi
Pinton,NJ: PrintonUnivrsityPrss.
mbridg:amridgshirBird lu. think aout th histor
Lak' D. t934. The Birds of mbridgshir.
-.7943. The Life of tbRobi. London:Withry. rlatd to his studis
-. 1944.ologialasptsof spis-formation in passrinirds.Ibls 86: similaritis and diffr
260-286. of his ontmpraris
-. 1945.Th Gal6pagosfinhs(Gospizina): A studyof variation.sioI ampls of unprodutiv
Pprsof th lifori dm of Sis2T: 1-|59. ..small fats
-. only with
1947. Duin's Fihes:A ss o th GrlBiologil Thor of
uoltion.amridg:amridgUnivrsityPrss. livd) wr afraid t
-. 1950.Robi Redbresl.oford: Oxford UnivrsityPrss. Lamark first st fo:
-. 1954. Th Nturl Rgultioof iml Numbrs.oxford: lrndonPrss. trodutory ltur h
-. in our.London: Mthun.
1956. St,uifts lassin 1800. By 180
-. -hor
|957. ullutilr d hristiBlif:h Ursoludo|lit. arth, from th simpl
London:thun.
-, dud y natural aus
1966. PopultionStudiesof Birds. Oxford: larndonPrss.
-, 197I. ologil Isoltili Birds. London: Blakwll. mous xpositions of h
-' 1973.y lif as n amaturornithologist[withappriatins y si (Zoo\ogia| Philosop}
ollagus]. Ibis 115: 421431. svn-volum Work ol
Prrins,. .2004. Lk' David Lamrt(1910_1973).In H. . G. Matthwand sns urtbres (Naturt
B. Harrison, eds',ford Ditior of Ntiol Biogrph.oxford: Oxford
Lamark sought to
UnivrsityPrss.
Thorp, \/.|7, 1974. David LamrtLk, 19|0-197. Birlgrphilmlirsof volutionry thorizir
Fellos of th Rol Soitof Lodo 20z 271-29. th sussivprodu
.Winr,
J. 1994. Th Bk of the Fi: Storlf utllutioni ur im.Nvl th diffrnt faultis
York: Alfrd Knopf. -]..
maintaind that th v
to b' gnratdspon
sifid, h laimd, as t
,.po
Lamark, Jan-Baptiste (|7 4 4_|829) powr of lif'' or
omplx'' (Lamark 1
Jan-Baptist Lamark Was th first iologist to st forth road, ompr- partiular irumstan
hnsiv thory of organi volution. Othrs for him had raisd th possi-
ntural ation of flui
ility that nw organi forms had appard in th ours of th arth's
said, for th ordrly sr
history, but Lamark was th first to dvlop at lngth th ida that all th
ths auss, in ontr
diffrnt forms of lif on arth had n produd sussivly,ginning
th min sal.It invo
with th vry simplst forms and poding gradually to th most omplx.
most famous: th id
Lmrk 67
that H plad man in a sp-
Lamark initially mad his sintifirputtion as a otanist, intratingin
L,kalso souglrt to ttr ur-
the 7770s with th irl f otanistsat th Jadin du Roi (th King's Gardn)
' many sptsof man's moral
in Paris. His arr hangd dramatially io 779 whn th Jardin du Roi
risaout th moral volutior-l
Was ronstitutd as th National usum of Natural istory. H was namd
on of th profssos of th nw institution, ut not to a hair of otany. H
t " ( 1 9 7 )i s a n a t o i o g r a p h y
Ws givn instad th profssorshipof .,insts,Worms' and mirosopi ani-
h dwzrrdGry Institut. Fo
..matur''to ..profssional.'' mals.'' H soon haratrizd this part of th nimal kingdom as th ..ani-
mals without vrtra''' Although his xprtis in this ara was limitd
owldgablmrnoril nly
initially to what h knw from having n an avid olltor of shlls, h b-
[d th Godman-Salvin Gold
am ovr th nt two dads th world's lading authority on invrtrat
958 nd th Darwin dal of
zoology.
lassifying th invrtrats and dvloping a ours of instrution aout
thm wr not Larark only onrns in th 1790s. Fror te Late 1770s
onward, h had spulatd roadly aout what h took to th asi prini-
toof Drtui'sFinhs.
pls of physis, hmistry, and mtorology. In th 1790s, h also gan to
ridg:mridgshir Bird lu. think aout th history of th arth. This nw gologial intrst was dirtly
y. rltd to his studis of shlls and to th mging prolm of plaining th
in passrin
irds.Ibls 86: .Whil
similaritis and diffrrrstwn living and fossil shll typs. many
of his ontmporaris wr inlind to s his road spulativ fforts as x-
t studof vrition.sionl
arnplsof r.rnprolutiv..systrrr_uilding,''
h omplaind of ritis who dalt
, 1 :1 - l 5 9 .
rcrIBilk;gilTbor of only with ..small fats,'' who faild to addrss important issus, and who (h
)rss. livd)w afraid tht thir rputationswould hrrrt y his idas.
rrsityPrss. Lamark first st forth his rvolutionary notion of spis hang in an in-
nbers.ford:lrndn Prss. trodutory ltu h delivrd to th studtrts in his invrtrat zoology
lass in 1800. By 1802, h was arguing that all th diffrnt forms of lif on
|'if:
Ursolt',d
o|lit.
arth, from th simplst to th most omplx, had n sussivlypro-
lrndonPrss. dud y rraturalaussopratingovr immnspriods of tim. is most 1-
BIakwll. mous positions of his idas appard in 1809 tr'hts Philosophie zoologique
ithappriations
y si (Zoolgil Philosophy), and thn in 1815 in the irrtrodution t<lhis grat,
svn-volum wok on invrtrat zoology, Histoir trell ds tm
973).|nH. . G. atthw and
sns urtbres(Natural istory of th ]nvrtebrats).
l Biogrph.oford: oxford
Lamark sought to aount for mor than hang at th spis lvl. His
973.Biogrphilemoirs of volr"rtionarythorizing addrssd in ddition th natur and origin of life,
1-29. th sussiv prodution of tlr diffrnt forms of animal organization' and
uolutiotti our Tle. Nw th diffrnt faultis to whilr ths forms of organization giv ris. H
-J., maintaind that th vry simplst fornrs of lif had bn,and wr ontinuing
to , gnratd spontanously. Ths forms thn am sussivly divr-
sifid,h laird' s th sult of two vry diffrnt sorts of auss:(1) ..th
powr of lif'' or ..powr that tnds unasingly to mak organization mor
ompl'' (Lamark I815-t82z 1: \4) and (2) th modifying influn of
to st forth a broad, ompr-
partiular irumstans. plaind th first of ths as a funtion of th
: f o r h i m h a d i s dt h p o s s i -
natllfal tion of fluids nrving through living tissus.It was rsponsil,h
l in tlr ours of th arth's
said, for th ordrly sal of progrssion of th animal lasss.Th sond of
at lngth th ida that all th
ths auss, in ontrast' Was rsponsil for th latral rmifiations from
' d u ds u s s i v I y. g i n n i n g
th main sl.It involvd th part of Lamark,s thoy that has sin om
'aduallyto th most omplx.
most famous: th ida of th inhritan of aquird haratristis.
674 Lmrck
to dvlopnw haits whn rpportt; prd d' ilttrodtio offrnt l dtrtitio ds rtres
ronmnts.hangs in haits ssentilsd l'iml, s distitkl du ugtl et ds utrs orps turls;
fi, l'positirl ds prinips fdmttl d l 7oologie.7 vols. Paris:
and oth organs ing usd _R.vB.
DtrviIl.
lnirrgth lattr. Diffns
ltion, lit too grdually to
'riods of tim, howvr, this
Land,Russll(b. 1951)
l was onfidnt that th dis-
giraffes,mols, storks, and Russll Ld is widly rognizdas th rhitt of modrn thorisof ph-
rsult of th inlrritan of notypi volution nd as an innovator in onsrvation iology. Born in 1951
l haits. Although th asi in Jakson, Mississippi, Land spnt th formativ yars of his youth in
' was not novl with him' h Shrvport,Louisian; Roswll, Nw io; and Sal Bah, alifornia.
;rh that th pration of this Th son of iology tahr nd a ptrolum gologist, h plord th
spishang. swamps of Louisiana and ody surfd on th bahsof southrn alifornia.
,h ntury, wll aftr harls H playd Littl Leagu and rd hoodd rats as a studnt in grammar
on, that th ida of th inhr- shool, and h Was a memr of math and honor soitisin high shool.
tan,''am tO widly r- Land's intrstsin ology w awaknd whn h was an undrgraduat
pis hang in ontrast to at the Univrsir1,f alifornia at lrvin. A tlntd studnt in mathmatis
This took pla vn though and th physial sins,Land lookd for pportunitis to tak a quantitativ
'd approirh in th iologil sins.That opportunity was providd in a wkly
i u s to n p a t o f h i s t h o r i z -
- i n h r i t n a s a n r h a n i s m rading lub organizd y graduat studnts Td s and Mihal Gilpin.
)ross.Th first srious hal- hir influn llourgd L:rnd to apply to graduat shool t th Uni-
,m Grman iologist August vrsity of hiago and to wrk with profssors Rihard Lvins and Rihard
sistd into th twntith n- . Lwontin, minnt pratitionrs of th modling appoah in ology and
it faild to gain primntal volutionary iology. Shortl aftr his arrival at hiago, Lvins and Lwon-
l y i t d i n i t s s r r p p o r tw a s i n - tin movd to Hrvard' Land followd suit.
)ns. As a graduat studnt at avard, Land fousd on dvloping thory
ris ontriutions to botanial for th inhritan of quantitativ haratrs nd using it to modl phno-
ringwork in invtbratpa- typi volution. A han rmak y Lwontin hlpd st him on this ours.
.Whil
houglrh is primariiy rmm- lturing on ndlis-r,Lew<lntinturnd from th halkoad nd
ngr apt-th ida of th quippd, ..Of osall ologially important haratrs ar quantitativ'
.thlssstands out in th his- polygni traits.'' Thr paprs (1976a, \976, 1977| tat ar now lassis
a omphnsivthory of or. in th filds of quantitativ gntisand phnotypi volution onstitutdhis
dotorl thsis.
Land ontinud to dvlop his novl approah to phnotypi volution as
a postdotoral studnt with Jams row at th Univrsity of lWisonsin at
tmrhd ullutionr .!right,
Madison. H also lrad rgular onvrsationswith Swall thn rtird
riversity Prss. .Whn
from th Univrsit of hiago nd living in adison' Land showd
horis i Fr,1790-180.
Wright th drafr of is 1"979papr' anothr lassi, Wright said that h did
]aliforniPrss.
not ar for th approah baus it did not modl th frqunis of th
lo sition ds osi drti os
diursitd lur orgltistio gns that undrlay th quantitativ traits. A yar latr, !right voluntrd
ohsiqs mintienl|tn
1u,i that h might hav n hsty in his judgmnt'
s t,ttt; fi,h llsqui Land's dvlopmnt of quantitativ gnti thory ontinud whn h
itIlig d u 1ui sont joind th faulty at th Univsity of hiago. During his Chiago priod'
volution. ah of ths ontributions stands out aus novl theortial if thr wer qualifid
formulations ar ombind with insightful litratur rviws and appliations C a m r i d g d u l y g a v l
to atual data. thy disovr that h w
Bginning in th 1980s, Land's interstsshiftd to onsrvation iology, thn rqustdro tin
as h took th lad in haratrizing th dmographi plight of th Northrn qunt|y rivdtop h
Spottd owl by using a omination of lassial' modrn, and original d- ors in his major filds
mographi thory. During this ongoing priod, Lnd joind th faulty at ..d
C m r i d g a l l d
th Univrsity of orgon and latr movd to th Univrsity of alifornia at ship from St. John.s
San Digo. Land is urrntly a Royal Soity Rsarh Profssor at Imprial Laky had n a
ollg, London. th summr aftr his s
pursu work in th ar
B|LIoGRAPHY
tory. Th first two a
Land, R. 1976a.Th maintnanof gntivariailityy mutationin a polygni d o u m n t t h s i sf o r
haratrwith linkd|oi.GtilRsrcb26:22I_24.
tumn of l930, h wa
1976. Natural sltionand random gntidrift in phntypivolution.
uolutio:14_34. asis of The Ston g
1977. Stattstlaltstsfor natural sltionon quantitativharatrs' u n i v r s i t i st' h r w r
uolutio7:442444. t o a r .T h s o l r q u i
1979.Quantitativgntianalysisof multivarit volution,applidto p u l i s h d o o k o u n r
rain:bodysizallomtry.uoltion3: 42-416. -cI
His Afianarho
tools. Baus Laky
s i s t a n o f S i r A r t h u r
Laky,Louis SymourBazett (t903-I972)
th Hunrian usu
Louis (pronound Lwis) S. B. Laky, th son of an nglish missionry t was to hav a prfoun
Knya in ast Afria, Was orn at hurh mission station t Kat, north- dspit dlaring his L
wst of Nairoi in Kikuyu ountry. It was th poliy f th hurh ission- yars in a hous on th
ary Soity to rotat its missionaris ak to ngland vry fourth yar so (Kith l 9 l5 ). H ws
that thy would not forgt thir nglish roots. Th Lakys wr rought human volution in r
ak in 194 a.d stayd for two yars' rturning to Knya in 1906. Thir dnfor humn vol
nt nglish sojourn was twn 1910 and 191'3, at whih tim Louis I n l 9 l , L a k yl
bgan formal shooling. In twn thos tims, thy livd t th missin sta- visit to th sit that wo
tion at Kabt. Thr young Louis joind a pr group of Kikuyu boys and susqunttim: oldr
am flunt in thir languag. Adoptd into th group' h ws formally ini- stanof Laky'sin
tiatd as a Kikuyu. Although h oftn said that h thought of himslf as a vai for mor than a th
Kikuyu, h atd and soundd lik a good British olonialist for th rst of ston tools dating a<
his lif. v o l v di n o t u h u m
\World .VarI intrruptd plans to Snd him to shool in ngland. .sfhn th i a l f v i n a m p ' h i s
family rturnd thr in 1'919, Leakey was duly snt to a publi shool in thos tools (M. Lak1
Dorst. Th phnomnn rfrrdto in ngland as ..puli'' shool is rally tition' but ir lkd tl
a privat institution. \/ith th id of a govrnss' h was instrutd in math- walking bipd. l
matis' Latin, and Frnh. Although of a longtim nglish family, Laky's ustrlop ith cs' pre
fathr had n orn in Fran, so Frnh was oftn spokn at th Laky forms hat did volr.
.!7hil
dinnr tal. Louis ould spak th languag, h vidntly did so with a nw gnri nd s
a pronound nglish ant' as did many of his ountrymn. not stik' lhoughin
Laky ntrd St. John's ollg, Camridg' as a frshman in th au- first ast Afrian and
tmn of 1922. He askd if h ould offr Kikuyu as on of th two modrn tivs frrdto s hol
languags rquird, and th univrsity turnd to authoritis in London to ask had n dvlpdt
Lke 677
)ut usnovl thortial if th wr qualifid Kikuyu spakrsin ngland. Tlr answr Was ys' so
t u r r v i w s n d a p p l i a t i o n s ambridg duly gv Laky th permission h had rqustd.Onl lat did
thy disovr that h Was on of only two Kikuyu spakrs known, so h was
ftd to onsrvation biology' thn rqustd to train th prson who am his own xaminr. I I sus-
raphi plight of th Northrn quntly rivd top honors in his languag xams. H also rivd top hon-
:al, modrn, and original d- ors in his majo filds of anthropology and arhology' giving him what
, Land joind tir fauity at amridg alld a ..doulfirst'' in 1926'This arnd him rsearlrfllor'i.
l U n i v s i t yo [ a l i f o r n i a t ship from St. John's ollg for tlrr yars' rnwd again for thr mor.
lsarhPrfssorat Imprial Laky had n ak to ast Afria as part of a dinosaur-hunting group in
th summr aftr his sophomor yar, ut with th fllowship h was al to
pursu work in th :rraof his training and intrsts,namly' Afrian prhis-
tory' Th first two ast Afrian arhologial pditions allowd him to
ilit by mutation in plygni doumnt th asis for an arhologialsqunof ast A1ri. In th au-
6:22|_24.
tumn of 1930, h was awardd a PhD from amridg Univrsity on th
i drift in phnotypi volution.
asis of Th Sto Age ulturs of Ke olo (1931). Unlike Amrin
n quatitativharatrs' univrsitis,thr wr no oufss or xams rquird in ordr to an do-
toat.Th sole rquimntwas n aptaldotoral dissrttion,nd a
virritvlution,pplid to publishd ook untd.
t6. -s./.. His Afrian arhaologialwork yildd human skltalmatril s wll as
tools. Baus Laky was not traind in skltal ilnalysis, h souglrt th as-
.1e72) sistanof Sir Arthur Kith, a highly rsptdantomist and th urator of
th Huntrian usum at th Royal Collg of Surgons in Londorr' Kith
n of an nglish missionary to was to hav a profound fft on Laky's views on hurrrnvolution. Kith,
ision station at Kat, north- ..Darwinism''
dspit dlarirrghis lif in and spnding narly th lst 25
poliy of th Churh N4ission- yars in a hous on th Darwin stat'was in fat pofoundly anti-Darwinian
ngland vry fourth yr so (Kith 1915). H was largly th rason why many profssional studnts of
s. Th Lakys wr rought human volution in ngland and Amria rfusd to apt th vilal vi-
ning to Knya in 1906. Thir d n f o r h u m a n v o ! u t i o n .
I |9\3, at whih tim Louis In 193i' Laky ld his tlrird st Afrian arh:rologialpdition tllr e
; , t h yI i v da t t h m i s s i o n s t a - visit to th sit that would idntifid with Laky's impat on th fild for all
)r group of Kikuyu boys and susqunt tim: olduvai Gorg in what is now alld Tanzania. Th su-
h group, h was fomally ini- stan of Laky's intrntional rputation was groundd in his work at oldu-
at thught of hirself as a vai for mor than ir third of a ntuy. In th first yar, h found quntitis of
itish olonialist for th rst of ston tools dadng ak to th vry alist mad y t anstorsof w,hat
volvd into tr humans. In July 1959, whil Lor.risws suffring from a trop-
l shool in ngland. Whn th ial fvr in amp' his wif' ary Laky, found tlr first fossil of makr of
ly snt to a publi shool in thostools (M. Laky 1984).It h:rdan ap-sizdrain and an normous dn-
..puli''shool is rally
d as tition, ut it lakd th projting anins of a tru ap and it was an rt-
'ss' h was instrutd in math- walking bipd. larly it ws latd to thos rtrmrs of tlr gnus
ryim nglish farrrily, Laky's Austrlopithl,ts, prviousiy known only frorn South Afriil and inluding
ts oftn spokn at th Laky forms that did volv into th genus Homo. As h did so oftn, Lky gv it
;uag,h vidntly did so with a nw gnri and spifi name, Zinjthrops blisei' Th gnus nam did
lls ountrymn. ,,Zinj.,,
not stik' although informally it is still known as Not only was it th
Jg, as a frshman in th au- first ast Afrian and th most sptaulr mmr of th arly hurnn rla.
yllas ol1 of th two modrn tivs rfrrd to as hominids, ut th thniqus of radioativ isotop irnalysis
o authoritisin London to ask had n dvlopd to th tl1ttht it ould givn an asolr.rtdat of ovr
678 Ldrbrg
1.5 million yars. Previously, th Australopithins known only from South ss in atri that r,
Afriir wr assumd to approximatly half a million yars old and muh that th prossLdr
too nt to hav playd r<>l irr l.luman volution. vn though Ztni was a or plasmid, that mdia
sidlirr,llot o th pth toward tu humans, its grar ntiquity nd primitiv othr' This pss,kr
hominid traits providd sr.tstan for Charls Drwin's (1s71) suggstionthat nbld sintiststo rr
Afria was th originl lol for th volution of hr-rnr:rn
form. soms' in a mannr lo
Laky took full advantag of th opportunity provid ed Zinj. H wrot y Thomas Hunt or
a sisof thnial as wll as popular dsriptiv artils. vn mor ff- Sond, working
'sisonsin,
tiv was his prsnon th ltur pltform. A harismati and olorful pu- Ldrrg
li spakr' his frqunt intrnirtionalapparansrought him arningsthat as vtors to mov g
supportd furthr fildwork, spially at Olduvai Gorg nd lswhr in this virus-mdiatdp
ast Afrir.This also nour:rglothrs to n.agin th sarh for arl ho- rol irr failitating ya
nrinids and thir tools lsrvhrin ast Afri. Hrdly a yar gos y now atil gntis.
without furthr disovris. Third, Ldrrg :
Laky also promotd fildwork on living grat aps-himpanzs and rplia-platingpri
gorillas in Afria and orangutans in SouthastAsia-y a sris of prot6g6s rial mutations-in par
who hav grtly pandd what w know aout t lif and ways of hu- ourrd prior to th
mans' losst nonhumn rlativs. mutations ould not
rtd mannr to th t
BIBLloGRAPY
at ando, with sl
ol' S. |975. Lk1,.sLk:.l'h l-if r'tfLouis S\'tlt<lr
tszttLk'Nrv York: nti dapttion. In o
HarourtBraJvaIrovih. winian mhanisms i
D:lrwin,. 1871.Th Dstttof , d Sltil i Rltitl/rl Se.Lor.rdln: Salvador Luria and tr
John Murra. sion, th rplia-plati
ith,A. 1975 . h tiquittlf . London:Williamsand Norgat.
Lky,L. s. B. 19I. Th StonAg ltrsof llon.amridg: it providd a striking
ambridgUnivrsityPrss. on a mastr ptri disl
1974.B thuide:loirs,19.]2_19.'1' Nll,Y.ok:arourtBra mutnt dsndantsi
Jovanovih. .!idnfld popultion, using a v
Laky'N,I'l984. Dislosingtha Pst:lt utobiogrD. Lodn: and sujt to sltionfo
Nilson.
lv{orll,V. 1995. Astrl Pssios:b Lk Flill'd th Qst for from th orrspond
Humnkid'sBgiiltgs. Nw York: Simlnand SlrLrstr. -.L.B. is<llatmutations witl
ditions that favord tl
At th tim that L
Joshua(1925_200s)
Ldrbrg, primnts, littl wa
Joshua LdrrgWas on of th lading miroiologistsand gntiistsof th ads, th fild of mi
twntith ntury. His primntsplord th natu of ftrndamntalgn. pralrs to undrst
ti nrhanismsin rria:rnd thus slrdlight on prosssthat hlpd shp attntion to volutio
th volution of th rrrostaundnt organisms on ilrth. For his ontriutions, work on mirobil v
lr rivd a Nol Priz irr 1958, at th ag of 3. Ldrrg also advisd prativs.First, th fi
poliymakrs on suh mattrs as mrging inftious disassand iologial trial spisto or
warfar, and h was awardd th Prsidntial edal of F.rdomin 2006. ry y arl !7os of
Ldrrg mad thr major ontriutions to undstanding atrial gn- girnismsthat ar dist
tis' First, as a graduat stulntworking with dwrd Tatum at Yal Univr_ of molular voluti
sity, Ldrrg showd that batria ould romin thir gnsto produ lrang within and a
nW gnotyps.This primnt dmonstratdth istnof a sual pro_ of th gntimhar
Ldrberg 679
importan in natur. Sond, owing to thir rapid gnrations and othr ompr (Lssos il
tratabl faturs, atria ar now oftn usd in primnts that sk to thr mor volums'
undrstand th dynamis of phnotypi and gnomi volution as wll as p u l i s h d y 1 8 0 5 . V
to tst partiular hypothss aout volutionary phnomna. Finally, thr th organs of snsatio
is growing attntion to th rol that vlution plays in th mrgn of and th organs for pr
nw inftious disass,inluding th aquisition of viruln fators that and srtion. uvir
allow pathogns to inft thir hosts as wll as th sprad of antiioti r- ..th ono
tions on
sistan. Th mhanisms of gnti hang disovrd y Ldrrg prfa to th work, l
hav n found to vry important in th mrgn of many nw a- ing dstroyd''(uv
trial pathogns. uvir quotd Immar
..us'' of vry sing
ILIoGRAPHY
organism in itslf. Dr
lna, S. F., and R. . Lnski. 2003. volution xprimntswith miroorganisms: notably Antoin-Laur
Th dynamisand gntibassof adaptation.Ntre ReuitusGetis4 tion of parts" (1789\
457469.
a n a t o m y ( 1 8 0 5 ) ' u v
Ldrbrg, I. 1997.Inftious disasas an volutionaryparadigm.mrgmg
IfctiosDisss: 417423. tion, nutrition, vision
..laws of oxistn'
Ldrbrg, J., and . M. Ldrrg.|952. Rpliaplatingand indirtsltionof
btrialmutants.Jourl lf Btriolog63: 99406. tion of on organ or
Luria, S. .' and M. Dlriik. 1943. utations of atriafrom virus snsitivityto th whol organism a
virus rsistance.Getis28 491-51\.
funtions of intrnal
National Lirary of diin.N.d. Profiisin sin.Th JoshuaLdrrgpaprs.
http://profi ls.nlm.nih.gov/BB/. divisions within th l
Tatum, . L., and J. Ldrrg.|947. Genrminationin th trium tiv anatomy provd
shrihioli. Jourl of Bteriolog5 673-684. sifiation.
.Wos,
. R., O. Kandlr,and . L. !hlis.1990.Towardsa naturalsystmof uvir plad stron
organisms:Proposalfor th domainsArha,Btria,and uarya. tion y insisting that
Prodigsof th NtklI Adm of SinsUS 87 4576-4579.
isms. Lif, h agrd
Ztndr, N. D., and J. Ldrrg.1952. Gntihangi Slmoll.lourl of
Bteriolog 64: 67 9-699. -R..L. proprty to tmpora
agnts. H did not dt
tim unvil th intim
..animal mahin.'' Fi
Lessons i Comprtiue Antom (Gorgsuvir)
tn of a hain of
In his vr-usylif, th yars 1795 to 1802 wr partiularly hti ons for isms to man. Gradua
Gorgs Cuvir. on July 2, 1,795,th profssors' assmly at th Mus6um intrnal organs, and
national d'histoir naturll approvd his nomination s assistant to Jan- twn ah of th ma
laud N4rtrud, th anatomy profsso, and h startd in arnst tO arrang anatomy was disov
th omparativ anatomy olltion. H gav lturs at th museum and
rad sintifi paps to a varity of soitis (th Soit6 Philomatiqu and |BLIoGRAPH
th Natural istory Soity in partiular) and at th Institut d Fran (insti- C o l m a nW , . 1 9 6 4 .G
tutd in 7795 to rpla th old Aadmis, inluding th Aad6mi frangais ambridg,MA:
and th Aad6mi ds sins).uvir took full advantagof th musum li- uvir,G. \80.Leqo
rary that nfitd, as did th musum itslf, from important additions . Baudouin.
q u i r d f r o m t h o s u o p n o u n t r i s o n q u r d y t h F r n h a r m i s . 185.Lgosd
Baudouin.
During 1798 and 1799, assistd y Andr6.Mari-Constant Dum6ril
J u s s i uA, . L . d .1 7 8 9
(1774*1860), uvir workd on th nts Dum6ril had takn at his lturs O u t r a m ,D . 1 ' 9 8 4 .G
in th prvious four yars. Th first two volumS of t'eLos d'tomie ruolutiorFrn
Lssonsin omparativAnatomy 681
Rudwik, N{. 1997. Gorges uuier, Fossil Bons d Gologil tstrophs: Th findings of L
Nlu Trsltios d Itrprettils of th Primr xrs. Chiago: Univrsity
Huy, suggstd-gl
of hiago Prss.
Viq-d'Azyr, F. |794 [1805]. uurs reuillis t publies ue ds ots t u|1 vry gn lous is po
discours sur s ui t ss ouurgs |)r . Jqus L. oru d l Srth. 6 (Huby and Lwontil
vols. Paris: Baudouin. -P.. This might sm to
viw of volution (i..
ural populations, mai
Lwontin, Rihard (. 1929) and against th
..las
(i.., that natural sl
Rihard Lwontin has provd to vry it as influntial in population g- '
ntis as his mntor, Thodosius Dozhansky. Lik Dozhansky' Lwontin within populations).
is wll known for artiulating a vry dfinit prsptivof th fild and th th rsults wr parad
most important prolms to solvd-inluding, lik Dozhansky' an s- hr was just too
pially pozz|ing..paradox.'' Lik Dozhansky, Lwontin is wll known for stiv' Lwontin ar1
a highly original approh to masuring variation at th gnti lvl and Kimura and Jams
for his own viws of how st to plain that vriation. Lik Dozhansky, a htrozygot ross (
periority at a partiul
Lwontin twakd molular iologists for suggstingthat mlulr pro-
sssWr mor fundamntal than th volutionary prosssthat atually rior homozygots and
gav ris to thm. Lik Dozhansky' Lwontin has n dply onrnd of th population. Tht
with th roadr moral and politial signifian of gntis and volution. suprior' th gratr
population, until th I
ary biology. And, lik his tahr, h has mntord sors of graduat and
postdotoral studnts. But ths ar similaritis at a vry high lvl of Morovr, Lwontil
gnrality. othrwis, ths two influntial population gntiistsar vry ttr snsof doum
diffrnt! variation'' tht thr
Along with Dozhansky' rnst ayr, Gorg Simpson, and othr volution- tion, no mattr whth
ary iologists' Lwontin grtdth fanfar surrounding th advnt of molu- th alan thoristsl
lar gntis in th 1960s with a ationary rmindr to kp in mind th proposd y th lass
xplaind in trms of
distintion twn ffiint and final aus in iology. Th molular tion, a position that L
onfigurations of living organisms ar th ffiint auss of iologial What is/was to d
phnomna ut not thir final auss. That is, xpt in a trival sns' tial tt, he Geti
th laws of gntis ar not th rsult of th strutur of DNA, ut rathr suh a rih thorti
DNA has bn hosn y natural sltionfrom among an immns va- op with th ody oJ
rity of moluls prisly aus it fits th rquirmnts of an volvd ary insight that in a flr
gnti systm. DNA is only th tati adoptd in th ours of working tivity did for th ont
out an volutionary stratgy.That is why som orgnisms n gt on th prolm mor pr
without it. (Lwontin 1964, 566| n? I livit is th
on rason for th
But h saw onsidralpromis in th nw fild and was at th vanguard of not ..mpirially suffi<
applying molular gnti thniqus to volutionary issus. In prtiular, b masurd dirtly,
h hlpd to pionr th us of gl ltrophorsis to doumnt variation twn altrnativ at
los to th gnti lvl-namly, at th lvl of amino aid diffrns rn pistmologial on
protins (du to nulotid diffrns in th gns that od for thos pro- infamous-for prsist
tins).In so doing, h hlpd stalishth fild of molula population gn- iologists an possib
tis. His studnt, Martin Kritman, WaS th first population gntiist to know. His ritiqus o
study variation dirtly at th DNA nulotid lvl, using mor rnt DNA drls of Sn aro an
squningthniqus. Gould) and gnti d
I'luoti 68
d G llogicl tstrop h es : Th findings of Lwontirr and his arly ollaorator in this work, Jak
m r t s . h i a g : U n i v r s i t y
Huy, suggstd-givn th inhrnt iass of tlrir mthods-tht n.ly
'ublis u des otas t Im vry gn lous is polymorplri, and that a third of all loi ar htrozygous
s L. oru d l Srth. 6 (uy and Lwontin 1966; Lwontin and Hr"ry|96Q Lw>ntin|974).
-P.. This might sm to b strong vidn in favor of Dozhansky's ..alan''
viw of volution (i..,that thr is onsidralgntivariation within nat-
ural populations' maintaind largly y sltion irr fvor of htrozygots)
and against th ..lassial''position of Dozhansky's arhrivl, . J. ullr
(i.., that natural sltion in favor of optimal gns liminats vri:rtion
; influntialin population g-
within populations). To his tahr'slrilgrin, Lwontin argud instad that
L i k D o z h a n s k y ,L w o n t i n
th rsultswr paradoxial' and not t ll sy to plain.
rsptivof th fild and th
Thr was illst too muh variation to mak sns from th alan pr-
ling, lik Dozhansky' an s-
., Lwontin is wll known for sptiv'Lwontin argud, iting similar onsidrtionsproposd y otoo
Kimura and Jams row' On major prolm is tlrat hlf of t offspring of
ltion at th gnti lvl and
a htrozygot ross (x) ar hornzygots (, ) . Heterozygot su-
variation. Lik Dozhansky,
priority at a patiular gn lous thus ntails th prdution of many inf-
lggstingthat molular pro-
rior homozygots and a orrsponding drasin th rprodutiv apatty
ionary prosssthat atua1ly
of th ppultion. Th gratr th numr of loi t whih htrozygotsa
n hs n dply onrnd
srrprior' th gratr th damag don to th rprodutiv apaity of th
r of gntisand volution-
population, until th population would b quit unal to prptuatitslf.
ttordsors of graduat and
orovr, Lwontin argud, no altrativto tl.lalan viw mks any
itis at vy high lvl of
ttr snsof doumntd pattrns of gnti variation. It is a ..parado of
pulation gntiistsar vry
variation'' that thr is too muh to plaind in trms f natural sl-
tion, lo mattr whthr it is variation-maintainingsort of sltionsuh as
Simpson,and othr volution_
th alan thorists had proposd, or a variation-rduingsot suh as that
oundingth advnt of moiu-
proposd y th lassial thorists.Tlrr was also too muh variation to
ndr to kp irr mind th
plaind in trms of th sltiv insignifian or nutrality of th vria-
: in iology. Th molullr tion' a position that Lwontin dud ..no-lssia1.''
.Wht
ffiintauss of iologial is/was to dorr?or as Lwontin psd th qustion in lris influrr-
is, pt in a trival sns, tial txt, Th Geti Bsis of uolutilnr bge (7974,267),*ov an
trutur of DNA, ut rathr suh a rih thotilstrutur as population gIltisfail so ompltly to
rom mong an immns va- op with the ody of fat? Ar w simply missing som ritial rvolution-
l rquirmntsof n volvd ary insight that in a flash will mak it ll om right' as th Prinipl of ltla-
:din th ours of working tivity did for the ontraditory vidn on th prpagatiorr of light? or is
som organlsms an get on th problm mor prvading' mor dply uilt into th struturof our si-
n? I livit is the lattr.''
On rason for th impass,h argud, is that population gntithory is
:ld and Was at th vanguard of not ..mpirially suffiint.''For aml, it inluds paramtrsthat annot
'utionaryissus' In partitrlar' masurd dirtly, or with suffiint aury' to distinguish larly -
t o r s i st o d o u m n t v r i t i o n twen altrnativ ausal aounts. his rflts Lwontirr's mor gnrl
l o f m i n o a i d d i f f r n si n pistmologial onrns-indd, his skptiism. is w|l known-vn
gnsthat od for thos pro- infamous-for prsistntly qustioning whthr gntiists nd volutionary
of molularpopulation gn. biologists an possily know what thy want to knW and oftn laim to
f i r s t p o p u I a r i o ng n r i i s t o know. His ritiqus of adaptationists(inluding his I979 arti|e..TlrSpan-
lvl,using mor rnt DNA drls of San aro and th PanglossianParadigm''' oauthord with Stphn
Gould) and gnti dtrmirristsar largly pistmologial.Adaptationists
684 L,llttin
ar-rdgnti dtrminists do not know nough aout th inhritan of th as]in th way that m
traits thy study and th volutionary fors ating on thos trits. Morovr, ally ror a as of sho
no on knows. This is a ntral motif of Lwontin's Geti Bszl and his livrs in th gnti as
stibsquntrviws of tlr stat of population gntis'Th titl lf on suh Lwontin's pisrmrl
rviw wll illustratshis disontnt vn with mtlrods of his oWIl dvising: matrialism, whih play
..ltrophorsisin
th Dvlopmnt of volutionary Gntis:ilston or
Millstn?'' (1991I. D i . r | t i e lm r r i l i
A sond rson for th filur of thory to mk snsof data in popul- mtlrod for solvir-rgpar
tion gntisis that pattr.nsof gnti vriation an du in lrrgpart to sis provids an ovrv
..history''-6or instan, not just whih nvironmnts a population pri- forms of dogmatism a
ns' ut th ordr in whih thos nvirtlnmntsare nountrd(Lwontin tht history ma' lav
|967). Population gnti hory,howvr, is an quilirir.rmthory tlrilt dis- rrrring ar dul as1
oLlntshistoiirlontingnis.Lwontin's disussiorrof th ahistorial har- ard that th onditil
atr of popultion gntis is partly pistmologial: history is ignord dstroyd y th
largly eausth past is so oftn unrovral.But it is also partly politi- ojts in tim nd sp
al. In population gntis,as in othr :rrilsof th rraturl nd soil si- tiorrs.Rmmbr tlrt
ns,quilirium thoris prvaild in tlr twntitl.lntury in part aus lost whn phnom
..prouptionwith tht all th othr avt
of a vry gnral staility'' (Lwontin 1974,269).
For Lwontin, th way forward in sin involvs, in part, posing th appliation to diffr
idologis that hlp to prptu:ltinorrt viws of h world' For ampl' ( L v i n s n d L ' r l l l r i
h rgud tn cnetic B-.istlrt th lssial/alanontrovrsyould not
understood without taking into aount th onfliting politial idals of B|LIoGRAPHY
Mullr and Dobzhansky. Gould, S. J.' and R. -.Lw
A ommon mistak of Mullr nd Dlzhansky' ording t<l Lwontin, Pr-rglssin palrdigm
Was thir gl.ltidtrrrrinistassumptiol-tthat xisting pattrns of gnti di- l|th I|ol SlittI
vrsity hav moal and politial impliations. For his own part' h hs laimd, Huv,J. I-.'and R. (-. [-
..I htrlzygosity in natu
do not liv that th ultimat issus dpnd on how muh divrsity -
in Drosophil psud
ists'' (Lwontin t al. 2001,4).lnstad of trying to did how o organiz I . v i n s 'R . ' n d R . [ , r r ' . l l l
soity on th ll,sisof gnti findings,w shold Llsour st undrstanding R' I-w<lntir-r. ds., h
of gntis' psyhology, soiology, and so forth to dtrminwhat intrvn- Harvrd Urrivsity Pr
tions would rsult in th sort of soity w aspir to (Lwontin t al. 2001, l,wontin, R. . 1964. A m
1 45 : 5 6 6 - 5 6 7 .
4344i Lwontin t al. 1984). Along th sm lins, Lwontin has lftnn
| 9 6 7. T p r i n i p l
onrnd to rr.rtth harg tht his skptiism onrning gnti dt-
M . . K p l a n , d s . ,
minism rndrs him ultural dtrminisr instad.As h quots Karl Mar' l 1|trp rttiI1of u olu
..Th matrialist dotrin that mn ar
th produts of irumstansand up- l972. h pti
ringing, and tht, thrfor, hanged mn ar prodts of othr irum- 8l-398.
sns and hngd upirrging, fogts that it is mn that lrang |974. h Glttt
Ullivrsity Pss.
iumstnsnd that th duator himslf nds duating'' (Lwontin t
1991. ltoph<l
a I .1 9 8 4 , 2 6 7 ) . ilston r millstl:
Th on as whr Lwontin has laimd politial signifianfor his own Lwotirr,R. ., and i. L.
findings in populzrtiongntishas to do with th sorts of studis that h ini_ hrrzygsity in n:lttlr
..Th Apportionrnnt of Huma htrozygosit],, i nrur
tiatd in his lassi 1972 ptlper Divrsity,''
54: 59-5-609.
in whih h argued that th proportion of variation within so-alld ras is
Lwontin, R. ., D. Pul, J.
far gratr (mor than 10 tirns) than th variation twn ras (s also
Llr.tlntin. ln R. S. Sing
Lwontin t al. 2001, 43). Raial diffrns.and all th othr soil nd po- h i kig but L-t'll ut
litial diffrns that lrirv aomp:rnid thm, ar thus not iologially f 2 . 6 l . C e r t l l ' j t l (:. .
Lotin 685
about th inhritan of th sdin th way that mny h:rd supposd.Aording to Lwontin, this is -
ing on tlrostraits. Morovr' ally more a as oi showing that gntisis not as politially signifi:rntas -
ontin's Geti Basis and his livrsin th gnti sis of raial diffrnshad imagind.
Th titl of on suh
4r-rtis. Lwontin's pistmol>gial and politial intrstsmrg in his dialtial
mthodsf his own dvising: mtrialism.whih plays a autionirry,huristi rol in his work:
ionary Gntis:Milston or
Dialtil matrialisrn is not, nd tlv lras en, 21programmati
mthod for solving partiular physial problms. Rathr, dialtial analy.
mk snsof data in popula_
sis provids an ovrviw nd a st of wanirrg signs ginsr patiular
)n an du in lag part to
) n m n r sa p o p u l t i o n x p r i - forms of dogmatism and nrrownss f thought. It tils us, ..Rmmr
(Lwontin that history my lv n impotarrt tra. Rmm that ing and
1 t sa r l ] o u n t r d
oming r dual asptsf natur. Rmrnbr that onditions hang
rn quilibrium thoy that dis_
and that th onditions nrssary to th initition of som pross may
:ussionf th Lrhistorialhar-
dstroyd y th pross itslf. Rmmr to pay ttnrion to ral
mologial: history is ignord
ojtsin tim and strra nd nt los thm in uttrly idalizd astra.
l. But it is also partly politi-
tions. Rmmbr that qualitativ fftsof ntxt and intration may
of th ntul and soial si-
lost whn phnomna ar isoltd.''And aov all ls' ..Rmmr
)ntithntury in part aus
'' (L,r,ontin1974, 269)' that ll th thr avats r onl rnrindrsand warning signs whs
appliation to diffrnt irumstnsof th ral world is ontingnt.''
nvolvs,in part, xposing th
(Lvirrsand Lu.ontin 198.5,|91-|92)
ws of th wold. For ampi,
[anontrovrsyould not
onfliting politial ideals of B|LIoGRAPHY
Gould,S..|..ant1R. . L,ontln, 1979.Th spandrlsof Sn o arrdth
nsky, aording to Lwontin, Panglssin paradigm:A ritiquof th adaptationisr programm.Proedigs
istingpilttrnsof gnti di- lftll Rry'lSlitl|.Lodl,Srists 20.5:'581_598.
Huy,.|.L., arrdR. . Lwontirl.1'966.A mlularapproahto th studyof gni
lr his own part' h has laimd,
htrozygosity irrnaturalopulations.l. h nLrm f lllst diffrntloi
n d r l r th t l w m u h d i v s i t y x - in Drlslphilpsudllr'lbscut.. Getics54:577_594.
i n g t t l d i d h o w t o o r g n i z Lr.ins,R.' and R. Lrvontirr. 198.5.Th prolnrof l,ysnkoism. In R. Lvinsand
uld us our st undrstanding R. Lwontir-r, ds.,}c Diltiltsiologist, \6-196. arnridg, A:
.h to dtrmin what intrvn- HarvardUnivrsityPrss.
lpir to (Lwontin t al. 2001, Lwotrtin,R. . l964. A moltrlar mssih: Th nw gosplin gntis? Sic
1.45566-567.
lins, Lwntin has oftn n
|967.Teprir-ripl in volution.In P. S. lorlradilnd
of histliit
;ism onrninggnti dtr- . . Kpln, eds.,themtilhllgsto th No-Drilti
.Wistr
:ad.As h quots Karl r, Intrprttit,l <l|-u<lllttil,81_88.Phildlhi: InstitutPrss.
cutsof irumstansand up- |972. Th apportionmntof human divrsity.ultionrBirlllg6:
r r p o d t r t so f o t h r i u m - 3 81 - 3 9 8 .
1974. h Geti l3sisof ulllutilrhg.Nw York: olurr-ri
rhat it is mn that hang
UnivrsityPrss.
r d s d i r a t i n g '(. L w o n t i n t l991. },ltrlphorsis in th dvlpmnt of volrrtionrygntis:
ilstonr millston?Ctis128:657_662..
litial signifiane for his own LwontiIl,R. (].'ntl.].L. u1'.1966.A nlolularapproahto th stud)'of grri
th sorts of studisthat h ini- htrozygsity in ntr.lralpopultilns. II. Amuntf varitinand dgrof
onlnnt of Human Divsity,''
htrlzygosity irl rlatualp1rrrlatinsof Drosophil psudolbscur. Gtics
54:595-609.
i a t i o nw i t h i n s . a l l d s i s
l-wot-ttit-t,
R. ., D. aul, |. Batty,ad . B. Krilr.rs.2001.Intrviw of R. .
'iation twnras (s also
Lwlntin.In R. S. Singh,. B. Krims,D. B. Paul,ndJ. Batty'ds.,
rnd ll th othr soil and po_ hiking llout |,Itttitl: Histlril,PbiloslphiId Politi,lIPrsptiL's,
?m' ar thus not iologially 22_61.mridg:()rnridg UnivrsityPrss.
686 Lineus
Lwontin, R. ., S. Rs' nd L. J. Kamin. 1984. No, i r Ges: Biollg, that inludd whals z
Ideolog, d m Ntur. Nw York: Panthon.
inal nomnlatur, th
Singh' R. S., and . B. Krima, ds. 2000. ullutior Gtis
fr-lmrlluls to gnri nam plus a m.
orphollg. amridg: amridg Univrsity Prss.
Singh, R. S., nd . K. Uynoyama, ds. 204. Th ullutio of Popultitl and Homo spies (
Billog. amridg: amridg Univrsity Prss. -I.B. ordr to fit th thous
arviations, omitt
l. H inludd itat
liographi guid or in
Linnaus,arl (or arl von Linn) (|77_|77s)
Linnaus,s systm f
Born and raisd in Swdn, arl Linnaus was passionataout otany from sophial foundations
his youth. By th tim h wnt to Holland tn |75 to finish his mdial d- larnd aout rnt
gr, h possssdan imprssivknowldg of th known kinds of plants and stamn and pistil, as .
animals. Thr h was hird to manag nd atalog th hug privat mu- vntd lasssasdt
sum' Zoo' hrarium, and otanial grdn of Gorg lifford' a walthy asd on dtails of th
mrhant. By th tim LinnaLrsrturnd to Swdn in 1'78,h had pu. asd on only a fw
lishd numrous works, all in Latin, that st out th ruls, framwork' and otanists ould us (
atalogs of ll th plant gnra thn known. H was a foundr and th first forward to th day v
prsidnt of th Swdish Aadmy of Sins.In |741 h was hird y th natural ons. At th l.
Univrsity of Uppsala, whr h taught for th nt 3 yars. rting a of faturs' inluding
dominating influn upon natural history (otany and zoology), h at- sion of th whol pla
tratd studnts from ross urop, som of whom h snt on ollting a onfusing mixtur '
trips ovrsas.H hangd lttrs with distant naturalists who snt him spis.In his lassifi
spimns.Most important, h frquntly produd nw ditions of his ata- vr smd to work
logs of plant and animals: Sstem Nature (12 ditions twn 1735 and insts' and fins for fi
1768)' Gener Pltrm (six editions from |77 to 1764), and Speis naus aimd at th lo1
Pltctrm (thrditions from 175 to 1764). Ths works providd natu- terms gus and s1l
ralists of all nations with on ntral, wll-organizd rgistrof nams. Lin- ill-foundd laim tha
naus'satalogs wr th sarh nginsof thir day. his aims and produ
Linnaus workd in th ontxt of th worldwid olltion of natural his- At th start of his <
tory spimns that aompanid th ploration, trad, nd olonization in latr yars h saw
arrid out y uropan govrnmntsand ntrprnurs.His partiular tal- think that God had
nts, whih inludd unloval traits lik gotism and an osssion with arising latr. ost of
making lists, quipped him to produ an pandal rgistrof th divrsity Baus visil varia
of living things. Although h ptd th nt gnration ro omplt th th ntir rang is n
projt, iologistsin latr nturis,inluding our own' ontinu to nountr sult of nourging
spisnot prviously dsrid.Th projt of making an invntory of lif is spis.Linnaus ha
still unfinishd. his studnts to study
Linnaus's systm ws stritly hirarhial, gouping similar spisinto a at what dt diffrn
gnus' similar gnra into an ordr' ordrs into a lass, and lasss into a alan of natur.
kingdom. Latr taonomists addd familis (low ordr) and phyla (low Th most rmark
kingdom), as wll as suordrs' suprordrs,and th lik; this pandd the Was that th strutur
hirarhy without hanging its prinipls. Linnaus gan th onvnint to ongnial to th
prati of attahing a propr nam to ah individual group, using a tradi- groups. Th old ida
tional on whr possil (suh as Avs and Piss' Latin for ird and fish, did th prdition
rsptivly)ut oining a nW on if ndd (suh as mmali for th lss Baptist Lamark th;
L'l|1,7us d/
ur Ges: Bioltlg1,, that inludd whals and humans). H also introdud what is alld inom-
). ial nonrnlatur' that is, always using two words to nam spis, th
Gtis from tlluls tl-l gnrinam plus a rnodifir, suh as is lupus (wolf), is fmilirls (dog),
'f.,ii,,,,,, and I7oml spies (human). (urrnt prati is to italiiz suh nams.) In
o|.Popultit
-J.ts. odr to fit th thousands of spis into mangal volums, h invntd
rviations,omittd illustrations,and mad dsriptions as short s possi-
l. H inludd itations of othr authors' so his works funtiond as a bi-
liographi guid or irrdx to th sintifilitratur.
)7-r778) Linns's sysrm fo flowring plants stimultd disussion of th philo-
ssionataout otany from sophial fundations f lassifiation. At th start of his arr h had
35 to finish his mdial d- larnd aout rnt thoris intrprting th ntral parts of a flowr, th
: known kinds of plants and stamn and pistil, as organs of mal and frlsul rprodution. H in-
talog th hug privat mu- vntd lasssasd on th numr and arrangnrntof stalns and ordrs
Gorg lifford, a walthy basd on dtails of th pistil or th fruit. Ths groups Wr artifiial, ing
d ni n 1 7 3 8 , h h a d p u _ basd on only a fw hartristis,ut thy gav ky tht vn inprt
: th uls' framwlrk, and otanistsould us (as long as th spimn was in flwr). Linnaus lookd
ws foundr and th first forwrd to th day whn ths artifiial groups <luld rplad y mor
7741 h was hird y the natural ons. At th lvl of gr-rra
and spis,Linnaus usd a widr rang
n t . ] 3 y a s . t i n g of faturs,inlLrdingth lvs and sds,and llowd his gnral imps-
ltany and zoology), h at- sion of tlr whol plant to affthis judgmnt.His ..sxualsystm'' Was thus
vhom h snt on o|lting a onfusing mixtur of artifiial lasss and ordrs with natrrral gnra and
tt nturlistsrvho snt him spis'In lris lassifiationof animals, he usd variolts harateristis,wat-
: dn w d i t i o n s f h i s a t - vr snrdto Work for ah group' suh as tth for marnmals, wings fo
ditions twn 175 and insts,and fins for fishs.Mny othrwis rlial historians stat that Lin-
77 to |764I' and Spcis naus aimd at th logial dfinitions of nrdival ..Aristotlians'''ut th old
hsworks providd natu- terms gt|us and spcies furrtiond vry diffrntly trom Aristotl's, nd th
rizdrgistrof nams. Lin- ill-foundd laim tht Linnaus Was an ssntialistadly misrprsntsoth
du. his ims and produrs.
id olItinf ntal his- At th start of his rr Linnus insistd that spisannot hang, ut
o n , t r a d , n d o l o n i z a t i o n in latr yars h saw vidn of hyridism and lnutation, whih ld lrim to
prnurs.His partiular tal- think that God had ratd only on spispr gnus' with th othr spis
ism and an osssion with aising lat.ost of his followrs prfrrdto stik with t fixity f spis.
l a l r g i s t or f h d i v r s i r y Baus visil variation within spis is oftn vry slight, speially whn
gnrationto mplt th th ntir rang is not sampld' th assumption of fiity had th positiv r-
.own, otltinuto nountr sult of nouraging th disovry and dsription of prviously unknown
laking an invntory of lif is spis.Linnaus had a livly intrstin th dyrranrisof n:rtu,nlurgirrg
his studrrtsto study whih kinds of plants various animals prfrrd to at'
luping similar spisinto a at wht dat diffrnt migrating irds rturnd, and how insts fit into th
l lss, nd lsss into a balan of natur.
w ordr) and phyla (below Th most rmarkal dvlopmnt toward th nd of Linnaus's arr
l th lik; this prndd th was that th strutur h had imposd in his lassifiationsmdvry oftn
l u s g n t h o n v n i n t to b ongnil to th pattrns of similaritis and gaps of pprntly natural
vidul group' using a tradi- groups. Th old idzrof a hain of irrg,a linar sl,did not fit' ut nithr
s, Latin for ird and fish, did th prdition of Gorgs-Louis Llr, omt d Brrffon and Jan-
:has anrmlia for th lass Baptist l,mark that onntions run irr all dirtions. Instad, pattrn of
Liuig ossl/-.
lumps, sts Within sts! smd natur's own. This unptd and gradually
mrging pitur Was th gat phnomnon lft to Linnaus's hirs. Unlik
Lamark's thory, whih grW Out of th l,rain of eing, harls Darwin's
thory dirtly addrssd th hirrhial pattrn of Linnaan taonomy by
positing a high rt of xtintion and a prinipl lf divrgIr.
BlLIoGRAPHY
r 25 49-54.
1u/pr/l/pro j.lin.html.
. lsof th issori
n t i l i s t Living rlativsof animals long rognizdas flssils,ut thought to -
-.P,w. tint' ar oasionally disovrd.Thy ar most apt tl rfrrd to as liv-
ing fossils, spially in th popularizatitln of sin.Th most familiar of
ths is th olanth.Th first fossil olaanth, of Prmian ag (255 mil-
lion yars ago), was dsridy Louis Agassiz in 1839. Today, 70 or mor
os not hav a pris thni- fossil speis rprsnting 27 genera ar known, ranging from th Lat D-
v hangd vry littl in form vonian (365 million yars ago) to th l-at rtaous (80 miliion yars ago).
jtors' r said to living fs- Th living olaanth' Ltimri, rtains haratristilim-lik stalkd fins
h mor divrs far ak in th and a joint within its rlativly primitiv brin:rsthat sparatsth nasal
ts volutionary hang tnds organs and ys from th ars nd rain. I'timri was first disovrd off
a t i o n l y i n t h x p a n s i o n South Afria in 198. Many additional spimnstlf th sm spishav
rgly ovrlap. sin bn aught off southastAfria and adjant t() th omoro Islands,
y prssion ..living fossil'' to nar Madagasar. Rntly, on spimn of a dif{rntut vry losly r-
|| as Polptrus, t|1eAfrian latd spishas n found on th othr sid of th Indian Oan, off th
mals a1lrtain primitiv har- lbs,in Indonsia.
intrmdiat twn typial Ltimri has not n found in th fossil rord, so th vllutionarytim
ving ony fishs' as Darwin span of th spis Ltimri hlm is unknown. It diffrs in only vry
ung.It has hair and is warm modst ways from its narstrlativ' a Lat rtaousolanth assignd
aturthan typial mammals. to th gnus ropom. Stritly spaking,it is th onsrvtivharatris-
s a fid st of sparatly r- tis of Ltieri that ar unquivoally living fossils, and not th spisor
alis.To Darwin, and now to gnus-a lustr of losly rlatd spis-in whih thy our.
s n d n tosf r m o t , t i n t Thr smsto nl singl planation for th long-lntinudsurvivl of
:ritillinks in th ranh- small groups of losly rlatd volving lirrags,with limitd spiatin and
plants. littl hang in form. Arthur onan Doyl imagind dirrosaursnd ptrodatyls
690 Liuig Fossils
wid gogrphi distriuttons. Nutilus poryilius, th mst ommon and r-lnifotrrand tlnstn|
varial of thsphalopods with lganthmdshlls,rangs from th Nonthlss, it has m.
Grat Barrir Rf to Nw Guina, th Philippins, and oani arhiplagos lLrsk, Neoplli, wit
as far out in th Paifi as Fiji. Fiv distint spisof Nutilus ar known. othr organs, is a d
Typially, ths animals liv at dpths of 100 to 400 mtrs or mor' on during th arly Pal
stplysloping sa ottors,los in gainst oral rfs.opulations living o{f among ivlvs at al:
isolatd oeani islands display limitd variation. Som onstitut distint largly n displad
spis,whil othrs ar rgardd as loal suspisof Ntilus pompilius' fdillg adaptatilns.A
This has promptd th rnt infrn that Ntillts is urrntly panding thir skltons from g
its divrsity, in a modst burst of spiation.To put this in ontt' not that dpths, low th zor-r
th spisand gnra of anint lozoi nautiloids hav n assigndto sort still our in slral
ovr 125 familis, with an traordinar varity of slill shpesand mods of rrrillirryars :rgo)rrl1
lif. Th prolifi divrsity of nautiloids was rpatdlyut ak in mjor Pa- ar th stalkd hino
lozoi tintion vnts.A furthr shrp rdution ourrd nar th nd f prominnt of all ;rnr
th Triassi (210 million yars ago). Sin tl.ln,th group has dwindld to spis of stalkd rin
th singl gnus that survivs today. n d a t a y s s a ld p t h s
Th horssho ras hav nvr n vry divrs, thoughout thir long and th olaanth,o
history. Fortns not dissimilar to th living animals first appard during th displmnts towrd
S i l u r i a n ( a p p r o i m a t e | 4 2 5 m i l l i n y r s g o ) . H o r s s h o r a s ' x t r n a l faun.r and tlrn tlrs
skltons ar not as asily prsrvd as mollusk shlls, so thir fossil rord i r l n e s h o r s t r i n g 5
may b lss omplt. Owvr, no mor than ight gnra an urrntly T varity of irtl
infrrd to hav livd t any on tim. Thr gnra ar living today. that thr is nl on
On fmiliar spis,Limlus polphmus, is aundant and widly distri- plas irr whilr to n
utd, from }ylain to Yuatan in th wstrn Atlanti. Irr th Irrdo-Paifi, tion. Living fossils do
two spis rprsnting two diffrnt gnra ang from th Bay of Bngal th ft that thy tn
to th Philippins' and third spis rangs from Borno to Japan. Lik major groups to whiL
Limuls polphemus, this last spis has a larg limati rang. It is also has bn a struln (
tolrant of varyirrg salinity and othr nvironmntal varials. In this r- spis,grotrps of v
spt' th horssho rabs ar quit unlik Nutilus, whih is tmpatur- from th stal dpt
snsitiv nd annot surviv in th wrm surfa watrs of th tropis, u i l r l .T r i l o i
rm;.rin
whr it livs. ut thy finally suu
Among living plants, th spor-aing1utnosssand horstails ar r. Thi |ong Jlin t
lits of groups tht dominatd vgtation of th late Palozoi. Both ar from hangs in omt
widly distriutd gogrphially. Th horstils, also known as souring Chn, otrldda
rushs, ar limitd to aout 30 spis of a singl gnus. Typially, thy ar thir ultimat dmis
oppotunisti olonists of wt slops and rivrbanks that ar frqr-tntly dis- ris and futLlrprosp
Liuig Fossl/s 6L)1
(nowldgof th inssil turbd. A similar hitat was formrly prfrrd y th gingko tr, whih
l Guyarr.Sorliving fossils may th oldst living spis' Fssil lvs nd wood up to 100 rniliion
y hav avoidd intrtion yars old ar morphologially indistinguishal from th singl surviving
d its spiny, hdghog-lik rl_ spis, Gingko bilrlb, of this anint group. This plant is so distintiv
dontids,lativlyul-lspial- tlrt it rrrrainsunrtin whthr it is nror losly rlatd to onifrs o to
ratl divrs zrnd widly th tr frn_lik yads. Orr widsplrd,th gingko survivd only in th
,lsozoi.oday. two spis
Yangtze Rivr sin in hin until its domstiation. Now, its auty and
.stritdto small islands off
tolran tlf nvironmntl insults hav rought th gingko to many ity
stts.
rnd th horsshoras hav Th dp ;anfloor is nithr as low in divrsity of arrimal lif rror as
'lius'
te most ommon and uniform and onstnt in its physil onditions as Was on supposd.
rrdshlls, rangs from the Northlss,it lras nlor than its slrar of living fossils. lr rping ml-
ts,and oani arhiplags lusk, Neoplli, wlt its ap_shapd shll and srial sts of musls and
isof Nutilus ar known' othr organs, is a dp-sarlit of two familis that livd irr shllow watr
to 400 mtrs or mor' on during th early Pirlozoi. Protoranhs with simpl gills prdominat
rl rfs.Ppulations living off among ivalvs irt ayssal dpths; orr th ontinntal slrlvs thy hav
on. Som onstitut distint largly n displad y mor advand forms with vrid urrowing and
isof Nlltilus pompilius' fdingadptations' Arong singl-lldforaminifra, only thos that uild
utilus is urrntly panding thir skltons fom grairrs of pristing sdir-rrnt an liv at th grtst
put this in ontt, not that dpths, low th zon t whih aliurrrronat dissolvs.Forams of this
tiloids hav n assignd to sort still ur in slrallow Watr haitats, ut in ambrian tim (aor"rt510
of shell shirpsnd mods of million ys ago) tlrrl1,this prinritiv cr()up was prsnt.ost striking of all
:atdlyut ak in major - ar th stalkd lrindrmspopularly known as sa lilis. z\mong tlr most
tion ourrd nar th nd of prominnt of all nimals in shallow-Watr Palozoi faLrnas, about 80
l, th group hs dwindld to spisof stalkd inoids suviv today, worldwid' on Ontinntal slops
and at yssal dpths. Ths distriutions' togth witlr tlros of Nutilus
ivrs'thrughout thir long and th olaanth.orroorat vidnfrom th fossil rord of sussiv
rals first appard during th displalntstoward dp Watr' first of dsndants of th amrian
,).Horsslro ras' xtrnal fnand thn th<lsf a lirtr lozoi fauna, ils nW orrlnruniisvolvd
< shlls,s thir fossil rord in narshosttin8s.
n ight gnra an urrntly Th varity of iumstansin whih living fossils hav survivd indiats
lr gna ar living today. tht thr is no on aus of tlris phnomnon. Tlrr ar lan ways and
rundarrtand widly distri- plasin whih to marginlizd witlrtlut ing liminatlby naturl sl-
Atlanti. In th Indo-Paifi, tion. Living fossils do not ollstitut a spial ast of organisms' pt for
.angfrom th Bay of Bngl
th ft that thy tnd to mong tl.rlst spializd mmbrs of th
from Borno to Japan. I-ik maior groups tl '"vl.rihthy long. At zrnygivrr tim in arth's histor,v,thr
rrglimati rang. It is also has brra sptrum of living things: rpidly volving groups of phmral
Lmntalvarials. In this r- spis'goups of vragstaility and tnur, anl groups that dpart littl
tttillts' whic is trprature- from th stal .ldaptations lf thir anstors, as long s thir nrods of lif
rfa watrs of th opis, rmin vil. Triloits Wr th prrninntliving fossils of Prmian im,
tthy finally suumd in th grat tintion irt th nd of th Palozoi.
mosssand horstails ar r- Thir long dlin fom urant divsity in th mid-Palozoi rsultd
th lat Palozoi. Both ar from lrangsin ommunit strutur tht ar not yt urrdrstoodin dtail.
i l s .a I s o k n o w n s s o u i n g han, outmodd daptation, and omptition all surl ontriutd to
gl gnus.Typially, thy ar thir ultimat dmis. Tody's living fossils hav similar volutionary lristo-
n k sr h t r f r q u n l yd i s - ris rrd futur prospts.
692 Lorez
,ies(1859),it am appar- umans, apal of rading and writing, ntrd urop around 40,000
yars ago. Somtims known as r-agnons' thes human anstors had
rl tht it was apal of lui-
originally volvd in Afria' and thir ultural and intlletual apaitis ld
lo. Furthmor,it plind
to th dmis of sr.rhspis as Nandrtals (0,000 yars ago) and ulti.
on rth.
matly to th popling of th ntir glob.
ilar to h othr as of .W
hav om 1rom huml ginnings. ah and vry fatur that d-
niqulydistint aus th
fins us as human-ipdalism, tool making, nlargd rains, rdud tooth
to at in mannrs rflting
siz, odis of modrn proportions' prhaps vn languag-first md thir
Darwin (1871) and his los
apparanin Afri. Ys, Lu hzis lft a lgay that unites us all s humn
to tk tstlprognosti-
ings with a ommon ginning in Afia, and it is my hop that this n-
Afriarr aps look alik, thy
largd, nlightnd, and profound insight into th qustion of ...!fhr did w
iffrnsar rlatd to how
om from?'' will unit all humankind into a fruitful nd lons-livd ommon
th ground).If th thory was
futur.
, as wll as dsntwith nrod-
)moton ll fours whn thy |LIoGRAPY
lhumans sprang from a long-
Drwin' . 1859. th rigi of Sples.London:Jhn urray.
f humanity rsid in Afria. 1871. Th Dstof , d Seltili Rltioll /o Sex. London:John
uy, and thousands of othr urray.
storsfirst mrgd in Afria. Huly,T. . 1863.ides trl ris Pli Nture, London:Williarnsand
and humans still luds - Norgar.
rnavatd Afrian statum. Johirnson,D. . 2004. Luy, thirty vrslatr:An -ptrndd
viw of
Austrlopitbas frsis.Joull of nthropoklgilRsrh60' no. 4:
ry'sspisas having livd at 465-486.
tion. Prior to Luy, th fossil Jhason,D. .' and . A. dy.1981.Lu: Th Bgiigsof Humkid.
r oming to light at 4 nrillion Nlv York: Simlnnd Shustr.
dipithus rmidus), and p- Johanson,D. .' nd B. dgr. 2005. Frlm l-u1,16 Lgug,Nr,r'Yrk:Sirrron
(Shlthr opus tch desis) . nd Shustr.
Kinrl,!. H., Y. Rak, and D. . Johnso.204.Th Skll of Australoithus
ory of volution, ths oldr
farnsis.Nw York: Oxford UnivrsityPrss.
imila to th prsumd mor Tttrsall,I., and J. Shwartz.200,tict ums' Bouldr,O: !stviwPrss.
-D..J.
sdinrntsw se vidn of
e. t )-.6 million yars' ston
nd, ar found assoiatd with Lyll,harls(t797_\875)
at had ntrd th human's
Invariably rmmrd'vn on his gravston'as th author of Priipls of
douling in brain size (Homo
Golog (firstditiotl in thr volunrs:1830, 1832, and 1833), harls Lyll
n irr largr skulls. h mod-
Ws th first hild <lfan nglish mthr' from lndd Yorkshir (Swaldal)
y short arms) appars shortly
fan"rily,and of a Sottish fthr, lird of a larg Angus ounty statought
is doumntd in th Rpu-
only 15 yars arlir y th oy's grandfathr, a farmr,s lad who had md
tus).
his fortun whil srving at sa in th Nw World' From infany, howvr,
hih hurnan spis migratd,
Lyll livd in southrn ngland, always in London itslf aftr graduatingwith
mor nrodrrrhumnlik n-
his BA from oxford in 1819. xtnsiv travls, largly in gology's srvi,
rgesis (500'000 to 1 million
took him oftn to ontinntal urop and four tims' in th 1840s and
ling glimpss of art, ngrvd
:ul manufaturof bon tools 1850s, to North Amria. H workd sporadially for a fw yars as a lawyr
in th 1820s, and rifly (181_1833) hld a profssorial hair in his sin
s go) two to thr tims ar-
at th nW King's ollg, Lndon. Othrwis, h dpndd on his privat
d t h o u r r n o f a n t o l n i -
..wis man'') at los to mns supplrnndy prods from writing and, in th Unitd Stass-
;dly
pially, from lturing,sO dvotil-lghimslf almost lusivly t his sin.
698 Lll
vntually knightd, a frirrllf Prin Alrt, .1ndn advisr to thr govrn- with what h had
mnt, h ilm a prominrrt r-riltionalfigu witlr ir h<luson rly Strt, affiliations.
.s7stminstr
fmous for its mdial grnds. His finl rsting pla is in That form, as
Ay. onform gology t
Th yars' frorn mid- l 82 to mid-1826' Wr th most formtiv for th xplaind by al
viws uphld y I,yll in Priipls. arlir, as an undrgraduat,t-yll had fid y Nwtonin
attndd !7illiam Bukland.s gology lturs nd thn don fildwork with thory-a nW vr
puliatiorr in mind, whil rr-rraining unitial of oxford-taught gology. onformal' pot
By mid-1826, howvr, h was pulily gr"ring for a furrdamntaland om- a v o i d i n g d i v i s i v 'i
prhensiv altrntiv to th tahings of th )ford shooi of gology and sin fom os
to th tahingsof ParisiirrrGlrgsuvir, th shool's primary mntor (u- ould only mt i
vir 1813). In mking this shift, Lyll was aov all intgratingth Huttonian roks wr all prol
thory of th arth, as dfndd yJohn Plyfair (1802)' whom Cuvir had nd with th sam i
onsistntly opposd, with th proposls of orrstant Prvost (1823, 1825), futur' th sam so
who wirs lpposinguvir's orvrrintrprtationsof tlr Trtiry formations of insistd, ntitldto
th Pris Basin. Lyll's ngagirrgwith la,vfairin his writings arrdwith Pr6vost by nrost gologists
irr prson wr prol,vokrd y William Fitton, a sretaryt this tim, knw nough of pr
long with Lyll, of th pstigiousGologial Soity of London. Fitton, an fts. By forming
Irish-born' Sottish-taindphysiian' Ws a suppotiv frind of th Frnh- his prim amition
man and an admiring sussorto Playfir as th main writr on gology for urrnt low rput
te dibrgb Reuitu. B1, lat 1823, Fitton lrad rjtd Buklnd's vatd status ompa
Noahian liluvial gology ln thc diburgb RcL,,itL, (s lso Fittorr 1824, takn to th hig
1828), whil it rvas nlltdin th rival Ttlr1,,Anglian Qttrtrl Ruitu Sottish gntry, Lyt
y dward oplston. Coplston, a futur ishop, was Oford's potry pro- th sinthat h ]
fssor, longtim ruttor of Playfair's ritiisrrrsof his univrsity,and Buk. proSPtivprofss
land's oyhood frind (Buklnd Was vn rumord to hav hlpd litist anxity Was
oplstlnwith th writing of his rviw). Th ultimat sta
Th t]iburgh Ruictu oftr-rdfind itslf y its liral, Sottish, r/hig, transndntani
forming opplsition to Tlry, Anglian, nd Oorri:rrrhgmonis in th (a livr in Gd l
n a t i o n ' s p o l i t i s , r l i g i o r l ' l t t r s ,a n d s i n .D t r m i n i n g L y l l ' s v i w o n and govrnmntn
any issu in gology or yond an start fro.r his family's and his own matrial soul om
.shn
alignmnts with th diltburg Rui,s stl1. Lyll wrot in Jan-BaptistLam
1826-1827 on duation and sintifi institutions as wll as on gology rjtd in th arl1
for th Qurtrl Reuita, lr did so s a onsious dissidnt intrudr. It mitmnts bout m
Ws as :r lissidnt intrudr too that h tauglrt :rt King's ollg. With rord s progrs
oplstorr ils n ovrsr' King's ollg ws dsigrrdt Toy, Angli- suh as Lamrk's
n for'rndation, out-ttrr,vight to th young Univrsity ollg, London' am thn to join
whih had n foundd y liral Sottish Whigs and hdd y Lyll's progrssiv nssi
own fathr-in-law, a rothr of an diburgh Reuit,tlfoundr. Lyll's nw tory of lif on arth
r f o r m i n g a m i t i o n i n t h m i d - 1 8 2 0 s w s t o r f u t a n d r p l a w h o l s a l many adaptiv on1
th Oford shool's n<l_(]ttr.irian gllogy, jr-rstwhn that shool had ing suh ontrsts\
ahivldorninan in amridg, in th mtrop;lis,and so in the ntion. orizing as raising,
Argualy, rhrfor' tis anrititll-lis pli;rlirs th mition of a young thology and mor
man wl.rohad om to think that starting ltlt in th Oxford shool was not hildlss and d
a pl whr a liral, Sottish, Whig lawyr lik himslf should vr' g- hugly inf]untial
ologially spaking, hav gon. onvrslv, h now had to at in aord any shol of om1
I,ll 699
'nd an advisr to th govrn- with what h had om to appriat as his authnti familial and prsonal
ith a hous on Harly Strt, affiliations.
i n g p, | e i s i n W s t m i n s t r That rform' as pounded in its fullst xtnt in Pricipls, ws (1) to
onform gology to t'eur us eidential-xplanatory idal (fftsar to
r th most formativ for th plind y auss known to ist from indpndnt vidn)xmpli-
an undrgraduat, Lyll had fid y Nwtonian lstial mhnis; (2) to giv th sin a systmi
nd thn don fildwork with thory- nw vrsion of th aosmogonial Huttonian thory (itslf now
l of oford.taught gology. onformal, potntially at last, to that idal)-that Was thortial ut
for fundamntal and om- avoiding divisiv, inonlusiv' osmogonil spulation;and (3) to fr th
ford shool of gology and sin from osai ilial assoiations.Ths nds, as Lyll saw thm'
;hool'sprimary mntor (u- ould only mt if gology prsumd that th past hangs rordd in th
a l l i n t g r t i n gt h H u t t o n i n roks wr all produd y uss still working in th sam irumstans
i r ( 1 8 0 2 ) ,w h o m u v i r h a d and with th sam intnsitisand so produing, in th psntnd on into th
n s t a n tP r v o s t( 1 8 2 3 ' 1 8 2 5 ) , futur, th sm sorts and sizs of ffts. It was indd a prsumptin, Lyll
of th Trtiary fomations of insistd,ntitld to a priori prfrnovr its dnial and so wrongly rjtd
his writings and with Pr6vost y most gologists beaus thy had prmaturly didd that thy alrady
i t n .e s r t a r ya t t h i s t i m , knw nough of prsnt auss to judg thm inadqut for thos past f-
ioityof London. Fitton, an fts.By rfoming gology y giving it ths prinipls, Lyll ould ahiv
)portivfrind of th Frnh- his prim amition for th sin and for himslf: to sav gology from its
Lmain writr on gology for urrnt low rput and vn ridiul y giving it sound foundtions and l-
ln had rjtd Bukland's vatd status omparal to th foundations and status of lstialmhanis,
ui (seealso Fitton 1824, takn to th highst sin of all. As th son and grandson of arrivist
, Anglian Qurterl Reuietu Sottish gntry' Lyll worrid intnsly aout th foundations and status of
op' Was oxford's potry pro- th sinthat h had mad his avowd avotion and vn, sin 1827,is
Lsof his univrsity'and Buk. prosptiv profssion in pla of th law. Notorious fr his snory, Lyll's
L rumord to hav hlpd litist anxity Was at on soial and ognitiv.
Th ultimat status of mankind-our filiation' dignity' and fat-raisd
ly its lirl, Sottish, Whig, transndntanxitis that, Lyll insistd, nly rligion an addrss. A dist
oxonian hgmonis in th (a livrin God ut not in th Bil), h hld to a Divin ration' dsign,
Dtrmining Lyll's viw on and govrnmnt not only for ntur ut lso for man, whos immortal, im-
'm his family's and his own matrial soul oms undr God's moral law in this lif and th rrt. H saw
n. Whn Lyll wot in Jan-BaptistLamark's (1809) systm of zoology, tnsivlydisussdnd
.rtionss wll as on gology rjtd in th arly ditions of Priiples, as dirtly thratning ths om-
tsious dissidnt intrudr. It mitmnts aout man. Aordingly, h rjtd any rding of th fossil
\i7ith rord as a progrssion,from low lif to high, so as to disrdit any shm
3ht at King's ollg.
lsigndto a Tory, Angli- suh as Lamark's that has humans produd from animal anstors.Lyll
Univrsity llg' London, am thn to join uvir' Lamark's othr opponnt' in rplaing any unifid
Whigs and hadd by Lyll's progrssivnssitatingplan with ontingnt' fitting adaptations in th his-
Ruilu foundr. Lyll's nw tory of lif on arth. harls Darwin,s thorizing rtaind this favoring of th
rfut and rpla wholsal many adaptiv ontingnisovr th on nssitatingplan. Whil rogniz-
just whn that shool had ing suh ontrasts with Lmark, Lyll alwys rad this nw Drwinian th-
r o p t - ' l i sa.n d s o i n t h n t i o n . orizing as raising' if xtndd to man' th sam old intolral thrat to
as th amition of a young thology and morality.
n th oxford shool was not hildlss and dstind not to inhrit th family stat' Lyll, although
lik himslf should vr' g- hugly influntial ovr svral dds f gologial sin,nvr foundd
L now had to at in aord any shool of omprhnsivlyonsnsual disipls. In his gology' Darwin
I.,h
was vry muh Lyll's most loyal protg6, vn aftr dvloping lris wn dis- Lynh gan his ar
sident iology. Lyll's no-uttonin s,vstmof th arth, itiizd from th of zoopl:rrrktn as a sl
start ovr its laims for a onstant' undelinirrg thrmal onomy for th n s o t a . l n a s r i so f 1
plant, Was mor widly thought implausil from th 1850s on aftr artiulatd a omprh
William Thomson (Lord Klvin) and othrs raisd thir novl thrmodynam. orporating disparat
ial o1tionsto ths lims (Burhfild |975). Th intriat omplxity of gtis, and ulminatin
.Walla's (Lynh 1980).His os
th young harls Drwin and young Alfrd Russl agrmnts
and disagrmnts With Lyll, and with Lyll's agrmnts and disagrmnts rprodutiv sussll(
with Lamark' put th proximat and ultimat dbts of volutionary iology tional wisdom of lif-h
to Lyll yond quik-and-asy summary gnralization. Thos dts ar sking th simplst x1
mor athr thn lss dp ausof that vry omplity. put him J th nt
quartr ntury.
|BLIoGRAPY Th 1970sand 1980
Burhfild,J. D. I97 5. Lord KIuid tb g of th rth. New York: Sin ary iology: th mrg
Histor,vPulitions. uration of th fild of
uvir,G. l813. sso th.heorof the rth.R' Krr, trns.dinurgl-r: rinvntd imslf s a
\fl/.Blakwood.
talished himslf as ]
Fitton' !. H. 1824. Inquirisrsptingth gologilrlationsof th dstwn was instrumntal in .
th hlk and th PurkLimstonin th South-rrst of ngland' Als of
gnetis (.g.,Lynh an
P hillso1lh
8: 365*383.
1828.Addrssdlivrdon th nnivrsary, Fbru:rry1828. Prldigs of Qutititiu rits, l
th GerlltlgiclSoit1,of Lodo 6: 50-62. standard for th fild ol
Lamark,J.-B. 1809.Philoso1lhie zoologiqu.Paris:Dntu. voiutionry quantit.
Lyll, . 1830_183.Priiplesof Golog:Big ttmpt to pli the lizing or positiv dirt
Formr hngesi tb rth's Surfb Rfretl usesNotu i
Kondashov, Lynh was
prtiolt.3vols.London:John urray.
Lyll, K. ., d. 1881 [1970].Life, Lttersnd Jlureils of Sir hrlsLlI, Brt' dltrious mutations t.
2 vls.London:Jolrrrurray. Rpublishdin fasimil,Farnoror-rgh, U.K.: Wilfid Garil, Rinh
Grgg IntrnationlPublishrs. ati study of th ffts
Playfair,J. 1802. lllstrtit.lsrlf th Huttt'lihor oi th rtb. dinurgh: tion xtintion (Lynh t
!illiamrh. ..mutationalmltdo
th
Prvost,. 1823.D l'importand l'tuddsrpsoganiss vivntspur l
th lvanof th r
gIogipositiv.moirs d SoitD'Histoir Nturelld Pris 1z 259-268.
1825.D la formationdstrrainsdsnvironsd ris.Bulltide l mind-st of iologists or
SoitPhilomtiqud Pris:74_77,88_90. In rnt yars, in o
Rudwik, .J. S. 2008.'X/orldsbefordm: Th Rlstrtil of Gllhistori A I l n F o r 'L y n h h s
th ge of Rform. hiag:Univsit,vf hiago Prss. prat ftursof gn
Wilson' L. G. 1972.rlasLll:Th Yrsto 784/. Nw Havn,CT: Yal
transposal lmnts(
UnivrsitPrss.
1998. Lll i mri:TrstltiGolog, 1841*1B5.1. Baltimor: 7999 Ln and Fort
Johns Hopkins UnivrsityPrss. -].H. first prinipls of popul
slightly dltriousmut
nomi faturs that dist
Lynh, ihal (b. 1951) from multillular orga
!7hn onsidring th impat a sintisthas had' on must tak into ount vrsial xplanation in
th radth :rnd dpth of h intllualontriutions.By thos ritri, vry ook The rigis o| ()
fw of his prs hv hd a grt irrrpat On volutionary iology than s y n t h s i so f r h i s w r k .
ihal Lynh. Furthr, his arr hirs n markd y that most valud of volution maks sns
..stady Kimura's lassi Th |
aadmi ahivmnts: progrss.''
Lh 701
r aftrdvloping his own dis- Lynh gan his arr as an ologist' studying th ommunity strutur
f th th' ritiizd from t of zoplankton as a studnt of Josph Shapiro at th Univrsity of in-
ing thrmal onomy for th nsota. [n a sris of paprs stmming from his dissrtation rsrh, h
ll from th 1850s on aftr artiulatda omprhnsivmodl of lif-history volution in ladora, in-
isdthi novl thrmodynam- orporating dispart lmntsinluding prdatirln, omptition, and nr-
i). Th intriatomplxity of gtis, i:rnd ulminating irr th first f nrany influntial synthti works
l Russl !alla's agrmnts (Lynh 1980). His osrvation that a tad-off twn prsnt and futur
agrmntsnd disagrmnts rprdutiv suss nd not b nssary was a hallng to th onvn.
, dtsof volutionay iology tional wisdom of lif-histor thory. Th undrlying philosophy of lwavs
nraliz:ltion.Thos dbts are skingth simplst plantion has n th hallmark of his arr and has
y omplxity. put him at th ntr of ontrovrsy irlmost unintrruptdly for th past
quartr ntury.
Th l970s and l980s witnssdtwo imprtant dvlopnrntsin volution-
thrth' Nw York: Sin ary iology: th mrgnof volutionry quantitativgntisand th mat-
uration of th fild of molular volution. By tlr mid-1980s' Lynh had
R. Krr,trns.dinurgh: invntdhimslf s qllantittiv gntiist and ovr th nt dad s-
talishd himslf :rs a ladr in th fild. Along with Russll Land, Lynh
.lrlationsof th ds twn
was instrumntal in odifying a nutral thoy of volutionary quantitativ
luth-astrfngland.ls of
gntis(e'g.,Lynh and HilI |986). Th txtook Gnetis d nlsis of
, Fruilry|828. Prldigsof Qutititiu Trits, oauthord witlr Bru Wlsh (1998), is th industy
standard for th fild of quantitativ gntis.
: Dntu. volutionary quantitativ gr-rtis historially fousd on traits undr stai-
|1|1
tt|1\ptto pli th lizing or positiv dirtion:rl sltiorr.Along with Tonroko Ohta nd Al
rarc t uses Nr>tui
Kndrashov, Lynh was among th first to ogniz th importan of slightly
rIsof Sir hrls L,ll,Brt, dltrious mutations to th volutionary pross. Along with ollaorators
tsimil,
Fanorough, U.K.: Wilfrid Garil, Rinlrrd Burgr, nd John ony, Lynh gan sstm-
ati study of th ffts of dltrious mutations on th proility of popula-
or of th rth' dinurgh: tion tintion (Lynh et a|. 199, 7995). Th rsr'rltingsynthsis stalishd
th ..muttionalmeltdown'' in th lxion, and, mor important, stalishd
rs orgarris6s
vivntspour la
irNturIId Pris |:259-268. th rlvn of th rlatiorrship twn fitnss nd ppultion siz in th
ironsd Paris.Bullti de l mind-st of iologists outsid th ralm of thortial population gntis.
In rnt yars' in ollaoration with onry and dvlopmntalgntiist
Rslttitl of Cois!or ilt Alian For, Lynh hs forgd markbl s},nthsisof svrl smingly dis-
go Prss. parat faturs of gnom volution' inluding dr"rpliatgns, intotrs, and
4 / . N w H v n , : Y l
transposallmnts (Lynh and onry 21,2003a, 2003; For t l.
,og, 1999;Lynh and For 2000a,2000; Lynh t l' 2001). oprating trom
1841-1 85. Baltimor:
-1.H. first prirripls of plpulation gntis' Lynh showd that population siz and
slightly dltrious mutations may intrt in suh a Way as to produ th g-
nomi ftllrs that distinguis prokaryots from rrkaryotsand singl.lld
from rrrtrltiliuiirr<lrganisms.ru to form, Lynh providd a simpl, ontro-
ld, on must tak into aount vrsil planation in liu lf a ompliatd' ontrovrsial on. His nt
, u t i o n sB. y t h o s r i t r i a ' v r y ook The origis of Geome Arcbiteture (2007) provids a highly radal
on volutioary iology than synthsisof this work. It is s an-ritilus in sop nd mssg(..Nothing in
narkd by that most valued o{ volution maks sns pt in light of populzrtion gntis'') as otoo
Kimura's lassi Tbe |'leutrl Tbor of oleculr uolutil (198).
702 L1,h
BlLIoGRAPHY
Fr, A., M. Lynh' F. B. Piktt, A. Amlrs' Y.-l,. Yn, nd.|. Postltlrr,vit. 1999.
Th prsrvatinof dupliat gns r, omplmntlrrydgnrlltivmutartions.
G tis 15 1 : 1.531_l 545'
Kimlrra, . 1983. Th Netrl Thor of rllulr ullutil, amridg:
mbridg Ut-rivrsity Prss.
Lvnlr, . 1980. Tlr volr.rtinlf 1adrnlif histlis.Q'LtrtrIRuie of
Biolog 55:2342.
althus, Thomas
207, The rigis of Grl Arhittr. Sundland,A: Sinaur Thomas Rort Malthu
Assoiats.
omist. H ws an ordal
Lynlr, ., R. Birrgr,D. Buthr' and W. Gil. 1993. Tlr mutati()1llmltdwn
in sul plpultions..|ttrlof 17ardit84 9-344. yars taught politial
Lynh, N4., andJ. S. onry.2000. h vlutionary fat and Irsqttns of ivil srvants dstind t
dupliat gns.Si290: 1 1 .5l-1 154. frindly and muh likd
2001. Gn dupliation nd vlluti<ln: Rspons tl Long nd Thornton nd His laim to fam is
Zlr:rrrgt aI.Sic 29: 1.5.5 l.
P rinipl of opultil
23a.Th volutionllry dmogirplrof dr"rpliat glls.In A. Myr and
Remrks o te Spul
Y. Van d Pr, eds.,Gom ullutillt,544, Dlrdrht,Th Nthrlands:
Kluwr Aadmi Pulishrs. ers. Malthus was dism.
2003. Tlr origins f gnom omplity. Sci 2: 141'-|404. writrs and thrfor tr
Lynh' Nl.'J. nry, ar-rdR. Burgr. 199.5.utatitlt umrtlationar:rdth mnt ar impossil.Ht
tintion of small popultilrrs.merin Nturlist 146: 489*.518. arlir argLlmnt of B
Lynh, M., and A. Flr.2000e. Gn dupliatior"r nd th origill lf intrspifi
ppulation numrs t
gnomi inrnptiilit,Amri Nturlist 156: 590_60.5.
2000. Th prlailityf duplit lln prsrvltion y supplis an at maximu
..struggl
sufuntior-ra lization. G ctis | 5 4 : 4 59_47. ar invital
Lynh, M., nd W. G. ill. 1986. Phnotypi vlution [y nutral mutation. at th prolm (Mlt
u o l t i o 4 0 : 9 l 5 - 9 3 5 . Malthus's hif intn
Lynlr, M.' . o'ly, B. Y/lslr,nd A. L.lr.2001.Th proaility of
how God had rrangd
prsrr,atinof rrwlv ais gn dupliat. Gatis l59: 1789-1804.
idl tim way aimlssl
Lynh, ., and B' Wlsh. 1998. Gtis rld nlsis of Qutititiu Trits.
Sundrlnd, A: Sinur Assoiats -'B, his work Was takn up
alulations 21sth as
workhouss wr mad(
sty out of thm. on
through wht Malthus
laus that Mlthus ad
spons to ritiism tht
ut loody intrhuman
lt was th sixthditi
nd of Sptmr1838
lizing that thr uld
gud that tl"rrwill b
motiv for hind a n
.Wlla
frd Russl war
s i s w i t h s l t i o t i t s
nd mad it a ruial l
.iutions on tpis s
divrs as
lluti<lrrof prthnogrrsis, th
iill ptlpulatiln gntis.
. t r I . u 'i u t i l t . t h i t l g :
tisttlris.Qurtrl Rui lf
althus, Thomas Rort (|7 66_834)
..Sutlrlarrd,
MA: Sirlr'lr
Thomas Rort lthus, known as Bo to his intimats)Was an nglish on-
l993. Th muttrtitln:rlmltdown
ol-nist. was an ordairrd lrgmar irr tlr Anglian Churh and for many
|:9_44. yastaught politil onomy at Hailyury Collg, shool that produd
'ry ft and ons1usof ivil srvants dstind to work in India. As an individual h was warm and
frindly and muh likd; h was an fftiv spakr dspit a lft lip.
spollsto Llng nd Thorntn nd His laim to fam is a ook that first appard in 1798, ss on tbe
Priiple of Populti>s It Affets th Fture Improt,ement of Soiet, ith
luplitgns.In A. iVIyrnd
[4. Dildrht,Th Nthrlands: Rmrks o th Spultions of r. Godi, . odlrt d thr Writ-
ers. Malthus ws dismayd at what h flt Was th fals optimisnr of many
' S i J 0 2 : l 4 0 | - l . 1 0 4 . Writrs and thrfo trid to show that grandios plans for human irnprov.
i o n t l t t m t t I l i onn d t h mnt ar impossil.H did this through a famous inquality, drawing on an
turllist146: 489_.518.
rli argumnt of Bnjamin Franklin and arguing that although human
nd th rigin of intrspifi
' 1 5 6 :5 9 0 - 6 0 5 . p o p u l t i n n t l m b st n d t o i n r s g o m e t r i a l l ( | , 2 , 4 , 8 , . . . ) , f o o d
lrsrvirtiny s u p p l i s a n a t m a x i m u m g o u p a r i t h m t i a l l (y1 , 2 , 3 , 4 , . . . ) . H n t h r
..strr'rggls
ar invitabl for xistn,'' nd plans fo stat aid only ar-
: t i o n v n u t r a Im u t l t i o n . at th prolm (Malthus 1798,14).
althus's hif intnt was natural thologial. H was onrnd to s
)01.h proilit of
how God had arrangd that humans do somthing hr on arth rath than
c tis 1.59:1 789_l 804.
. idl tim away imlssly.Th lthusian inquality is th answr. Howvr,
l sis f Q tititi rits.
-.B. his work Was takn up with nthusiasm y soial formrs' wh usd his
alulations as th asis for drlnian rforms of th Poor Laws' wh
workhouss wr lnad so unplasnt tht th porr would do nything to
staY out of thm' On wy of voiding suh dgrdation would hav n
through what althus trmd prltdtil rstrint (forn rding), e su-
laus that althr"rsaddd to latr, muh-nlargdditions of his ook in r-
spons to ritiisr that h smd to hav implid that God lft n option
but loody intrhuman strif.
It was th sith dition of althus's ook that harls Darwin rad at th
nd of Sptmr188. Gnralizing to th animal and plant worlds and r-
lizing that thr ould no prudntial rstraint in thos aras, D:lwin ar-
gud that thr will a univrsal struggl for istn'and that this is th
motiv for hind a natural form of sltion. Twnty yars latr, whn Al-
.$7alla
frd Russl was indpndntly formulating an volutionary hypoth.
sis with sltionat its ntr' h tlo rmmbrdth Malthusian inquality
ad rd it a ruial lnrntof his thirrkin.Lik althus. th vlutionists
70
704 mmls
th Prmian (z99_z5\
larg lnd vrtrats,spiall prominnt during
vior'rs,whr organisms at- plyosaurs, whilr spannd
million yars ago). Th arlist forms wr th
anl.Walldiffrd in thir dogs today and inludd arrri-
aout th ody-srzrng sn in dmsti ..sils'' on
inlindto think that it is b- had distint
vors, hrbivo..,, nnd fiih .nt..,. Som plysurs
oW thirt somtims it ours hat xhngrs.
thir aks, whih wr praly usd as nvironmntal
that wr in tropial or
Piyosaurswr mainly io..nd in gographi aras
Amria.
sutropialZonst that tim, suh s North
wr largly xtint' rpld y thir
y tir. Lat Prmian th plyos:rurs
lpLitttk. Llndon: Printd for thir lgs mor undr-
{w Ytlk: N4rillan,1966. d.*..,,d"nt,, th thrapsids. Thrapsids had movd
upright thn th sprawl-
optt|titltl. 6tlr d. l-ondon: nath thir body, with a stan that was now 1or
tl run and ratlr t
ing, lizardlik postur of plyosaurs, naling thm
of a highr mtaoli rat.
|i Vitorion ,,'',',,'-'.n' th sam tim; this is suggstivof th voltrtion
linags) volvd vrious fa-
Latr thrapsids (onvrlJntly n*n.,g diffrnt
rat, suh as inrsdvol-
,u'., ,ugg.,.iv of inr]si,rglvls f mtaoli
um in th skull for iaw -i,.l., and mor omplly diffrntitdtth,
Th thrpsids iIrludd
indiativ of inrasej amounts of food prossing.
smallinstivrsndmdiumtolarg-sizdarnivors(sornasigaslions)
snddfrr rptils, ut al-
andhrbivors(smasigasbison,withhornlikknobsonthirhdsthat
aior divisions of irmnilts,th Bt in ontrast with modrn
might hav n usd in iitrspifi omat).
aurs),mammals nd thir fos. tlrlvr no glidrs,
nrammals or vn with ontmporanoussauropsids,
th synirpsids.Nonmammalian
swimmrs'oripdalrunnrs.LatrinthPrminthrapsidsamom-
i' a fatf rnisladingnam - this tim. This hnging go-
mon in South Afria, whih had old wintrs at
rosmodrn forms tht w all for thrmorgulrion.
g r a p h i d i s t r i u t i o na l s s u g g s t sa n i n r a s da h i l i t y
two main linags of tlr.
Aftr th dvstating .'1J-'..Ini*'1 xtintiOns
rany uniqu fatures' suh as hrivors' and th yn.
apsidssurvivd:th diynodonts' pig- to ow.sizd
ls hv also rtaind numbr latr volvd largr
odonts, initially small and instivorous. ynodonts
)st or modifid in sarrropsids. non Was muh iggr than a
hrivoousand rnivorous forrs' although
: volv th thikr sals with
Labradordog.ynodontswranstraltomammlsandhadntlmrous
psids (s rvll as th mor gn- rat, whih suggsts som
anatomial faturs suggstivf a high rntaboli
thy hav n abl to volv
degrofmammal.lik-ndothrmy.ororrrpltthandvidnofn
-hat prodtrvarious odifrous w prossingror food.
inrasdvolum of jaw musls show that thy
. r.lrivdeility to xr ui vidn of a mr.rsulardi-
An inrasdrt of lung vrrtilationis shown y
smisolidWast' synapsids x- y vidn o{ turbinat
aphragm (rdution oft-th postrio ris) and
in a urinry laddr (whih is to Wrl inlming
(sroll-lik)ones in th. .,n,ai avity. Turinats at oth
; l l sm m m l st o t ' t s u r i n aus wtr loss fronr th lungs
air anl to rlaim wat from pird air,
Tlr typiall mammalian us vntilation.
n b a signifiantprolm with a high rat of
ftrr sprm tansnrission'is also thrpsids startd to fa omptition from th radia-
Irr th latr rissl
rp. Diffrns in th intrnal (anstorsof dinsaurs).
tion of largr sauropsids, inludirrgth thodonts
st that vrious fatllrs volvd
Gloalhangsinvgtation,suhasthrplamntoftlrmlrrhai
artiular, ndothrmy (warm- volutionary trnds.An-
sdfrns by th onifrs,my hav influndths
irds and mamtnals. that tmosphri oygn
othr iss that latsto this rplamntmy
luring th L,atarbonifrous, in rh Prmin, and it an
lvlswr apparntly lowr in t Triassi thn
.h liegslookd rathr |izard- thodonts (rodilsand
infrrd from th bioiogy of th living rltivsof
th skull hind th y sokt, providd thm with ompti-
birds) that suplor lu,ig fun.tion may hav
L attarnnt.(ost sauropsids mor highly r-
tiv lg.Additionally,1h. .gg, of irds nd roodils
:ull, ut with two hols on ah this {aturrny hav givn
sistant to dsitronthn thos of othr mrriots;
;ids wr initilly th dominant
706 mmls
ditary itms. Thrians also had a mor hinglik ankl joint and a shouldr
ntlydrir onditions of th
ivly smallr' prhaps down_ lad that luld now swing frly, ontriuting to th strid lngth and mak-
ing possil th haratristi ounding gait of many modrn mammals. Both
h Trissi (around 200 mil-
modrn marsupials and p|antalsar also viviparous (giving irth to ais
Js rmaind. Two rat-sizd
rathr than laying ggs),although thy do it in somwhat diffrntways' and
urassi, along with th first
it is not lar if viviparity Was prsnt in thir ommon anstor or if thy
this tim, although a rlit
,known from th arly r. volvd this ondition indpndntly. rtaous mammals wr divrs, srm-
ilar in numrs of spis to th rtaous dinosaurs, ut wr still small,
Lammalsof th Jurassi wr
non biggr than th siz of a small dog. Although th nozoi ra is om-
: instivorousdit. Th ma-
mnly alld th ag of mammals, it in ft rprsnts only a third of mam-
s, a ondition that ontinud
malian volutinary history, whn mammals raditd out into largr ody
'ainsthan ynodonts' as wll sizsto fill th typs of larg land vrtratnihs prviously hld by th di-
nosaurs.Th volution of nozoi mammals is st undrstood in th on-
lxion up nd down. In latr
tt of th hanging pattrns of th arth's ontinnts and th subsqunt
aturallowd for ounding
.tantto allow mammals to li hangs in gloal limat. Although the suprontinnt Pangaa startd to
lf mammals now intrlokd rak up during th sozoi, th southrn ontinnts Wr still unitd in
Gondwana at th start of th Cnozoi, and th northrn ontinntal loks
ln) of th food, and formd
had not yt rahd thir final positions. Movmnt, fragmntation'and oa-
as in humans).Th volution
lsnof ontinntal plats ratd hangs in oan urrnts and pisods
tion was proaly important
of mountain uilding that rsultd in gloal limati hangs.To a first ap-
r rplingtthould not b
proximation th world hangd during th nozoi from initially gloally
loth rplamntis probaly
warm' tropial-lik onditilns to a world that am inrasingly oldr at
l u s t h r u p t i o no f d n t i t i o n
th higher latituds and inrasingly drir throughout.
solid fod. Th volution of
Ths hanging glol limati onditions onsquntly inflund th
rld form a sal and musular
typs of mmmals that volvd. For mpl, th ologial nihs now
s of th omplx sris of fa-
filld y nimals suh as polar ars and amls wr not in istn until
tans now us for sph and
around 5 million yars ago' with th dvlopmnt of an Arti i ap and
.rms(platypussand hid-
th dsrtifiationof tropial rgions. AdditionalIy' th isolation of diffrnt
ls, thsarly mammals must
'ls and an xtrnal ar, and sd groups of mammals on diffrnt ontinntal bloks (lthough this
mainly <rurrdvia migration rthr than th mammals ing arrid on th
sparatingontinnts)rsultd in a grat amount of onvrgnt volution of
:s of aly mammals volvd
mammalian typs. Th st known of ths r th xampls twn Aus-
orous in thir dit. Fw wr
tralian marsupials and plantal mammals lswhr(.g.,plantl wolf and
int by th rtous. our
marsupial ..wolf,'' or thylain),ut mny othr ampls ist twndif-
rrsdin th past fw yars,
frnt groups of plantals. Fo xampl, th Afrian goldn mol ts mor
3, and somwhat largr (opos-
losely rlatd to lphantsthan it is to th tru mols of th Northrn Hmi-
y of known forms. In th Lat
sphr.
lnivrous/hrivorous linag
of th various mammalian omorphologial typs (partiular ody typs
:aus of thir ompl hk
adaptd for spifiologial rols), two typs that hav rpatdlyvolvd
rssfullinag of rodntlik
at diffrnt tims and plas in th past ar missing from today's world. On
ntil around 40 million yars
is th sar-toothdprdator. This typ of mammal volvd twi within tru
h volution of tru rodnts.
ats, within a now-tint family of ..fals sar-tooth'' arnivors, and in
' a nW type of mammal ap-
South Amrian marsupils. (Australian marsupials produd a similar typ
rsupialsand plantals.(lntr.
of havily uilt, musular prdator, th marsupial lion, whih had volvd
ris tim.)
from hrivorous forms and so lakd th distintivanins.)A sond typ
:s of hk tth (triosphni
is th larg (ison-sizd)lawd hrivor that ould ris on its hind lgs to
roadrsptrum of availal
mmls
pull down ranhs. ammals lik this volvd among th hlrs-rlatd rn Hmisphr. Th l
haliothrs in th Northn Hmisphr and Afria, among nativ ungu. onfind to th trtlpis
lats and giant ground sloths in South Amria, nd among womat-rlatd ats) mad an appar
forms in Australia. hrivors)Wrom
A rif history of nozoi mmmalian distriution is as follows. At th rathr than fruit atrs
start of th nozoi, Afria was isolatd from th othr ontinnts and took ovr som of tht
volvd its own spializdfaun (.g.'lphnts);primats wr also known rly ion, around
arly on in Afria, ut othr northrn immigrants (suh as arnivrrsand un- and drying during th J
gulats, or hoofd mmmals) did nt rah thr until th arly ion, th sprd of savann
around 20 million yars ago. Th monotrms Wr th oiginal Australian Long-1ggdungulat
inhabitants; marsupials (immigrating from South Amria via a thn-i-fr in North Amria (hor
Antatia) Wr not known until th rly on, around 55 million yars tiv forms). Howvr, 1
ago. South Amria was isolatd until th middl Plion (2.5 million yars Wr not in vidrru
go), whn th Isthmus of anama formd. Its original fauna onsistd of ing in th lat nozt
dntats (sloths' armadillos, and antats),marsupials (lpossums nd transformation of ths
orhyanids, now-xtint prdatory forms), and nativ ungulats (now all lss prairi' with th l
tint). Around 40 million yars ago it gaind its stoks of primats and million yars ago) oo
aviomorph (rltdto guin pigs) rodnts' most likly from nimals rafting Northrn Hmisphr
ovr from Afria. Alut 2.5 million yars ago many frms migratd from dr tod and wolly
North Amria, nd now ovr half th South Amrian fauna is of rnt nw high-latitud ha
northrn origin (.g.,ats, foxs' tpirs, dr, and mouslik rodnts).North and drying rsultd in
Amria and urasia hav long had fairly road faunal onntions,ut today ial Afria and Asia, rv
their mammal faunas ar mor similar than in many Past tims. For ampl, that was formrly om
North Amrian dr and ison ar reent (Within th pst 3 million yars)
|BLloGRAPHY
ursian immigrants.
During th arly nozoi (th Plon and arly on pohs, from Agusti,J., rrdM. Ant6n
65 to 50 million yars ago) th world was in gnral warm and qual and' Ycrs f mmlit
with th rowsing pssur of th hrivorous dinosaurs movd, was largly Prss.
Bnton,. J.2004, Vr
ovrd in forsts;topil-lik forsts tndd vn within th nfins of
A: BlkwllPu
th Arti irl. Th typs of mammals Prsntwr largly roral inst iflli, R. I-.2001.rly
and fruit atrs'with a fw trrstrilomnivors nd hrivors.Small prd- 12\4-1226.
ators inludd rly mmrs of th modrn ordr arnivora, ut largr J a i s , . . 2 0 0 1 .V i t o
prdators wr not divrs and blongd to groups now tint. Most of d.,Th Book f Li
Krnp,. S. 2005./:e(
ths mammals hav n trmd rhi typs, maning that thy longd
UnivrsityPrss.
to linags that did not prsist until th prsnt day (indd,most Wnt x- Kilan-Jwlrowsk, Z.,
tint for th nd of th Palon).Th start of th on saw th first lf Diosurs:rigi
apparanof primat and modrn ordrs of ungulats(with th first rodnts UnivrsityPrss.
apparing in th latst on) and also th highly spializd ats nd Plugh,F. Ll., . . Jn
whals. Th first known fossil ats ar alrady larly spializdflirs, and SaddlRivr'NJ: Bt
Thwissn,J. G' ., n
w hav littl information on thir anstry. Howvr, in th past dad a
marovolution. Bir
sptaularsris of arly fossil whals hs n olltd that larly shws Turnr,A', and . Antt
th transition from a land animl to a srndrilyaquati n. uolutio of th f
Around 50 million yars gO tmpraturs in th highr latituds startd to UnivrsityPrss.
fall, rsulting in inrasd sasonality zrnd wintr frosts. By th start of th Vughan,T. A.' J' A. Rv
P h i l d l h i S :a u n
oligon (33 million yrs ago) th tropial forsts wr onfind to the
qutorial rgion, and diduouswoodland sprad aross muh of th North.
mmls 709
lvd among th hors-rlatd rn Hmisphr. Th arhai manrmls Wr now all tint, primats wr
d Afria, among nativ ungu- onfindto th tropis, and mor modrn typs of larg prdators (dogs and
a, nd among womat-rlatd ats)mad an apparan'Rodnts and raits (or quivalnt typs of small
hrivors)Wr omlon, and th ungulatsWr largr and mostly laf atrs
triution is s follows. At th rthr than fruit atrs.In South Amria larg rldnts (lik ttlday's apyara)
.om th othr ontinnts and
took ovr som of th small ungulat nihs. A warming trnd during th
rts);primatswr also known arly Mion, around 20 million yars ago' followd y susquntooling
nts (suhas rnivrresand un- and drying duing th lt ion (12_.5million yars ago), rought with it
:hruntil th arly Mion, th sprad of savann-typ haitats (trd grasslands) at highr ltituds.
|sWrth original Australian Long-lggdungulats'with tthmodifid for ating grass,amprvalnt
rth Amria vi a thn-i-fr in North Amria (horss),urasia (antlop),and South Amria (tintna.
'n'around 55 million yars tiv forms). Howvr, th quivalnt typs of marsupil, th largr kangarols,
dl lion(2.5 million yars Wr not in vidn until th Plion in Australia. Furthr ooling and dry-
[ts original fauna onsistd of ing in th lat nozoi (from around 8 million yars gl) rsultd in th
), marsupials (opossums and transforrationof thspodutiv savanna haitats into lss produtrvtr-
rnd nativ ungulats (now all lssprairi' with th loss of larg mammal divrsity. Plio.Pliston(from 5
rd its stoks of primats and million yars ago) ooling rsultdin th i ags (larglyprindy th
rostlikly from nimals rafting Northrn Hmisphr),and spializdmammals (suh as musk o and rin-
lo mny forms migratd from dr today and woolly rhino nd mammoths in th rntpast) inhaitd th
r Amrin fauna is of rnt nw high-latitud haitats suh as tundra. Howvr, i-ag glol oolirrg
and mouslikrodnts).North and drying rsultdin th dvlopmntof xtnsivsavanna haitats in trp-
l faunalonntions'but today ial Afria and Asia, whih tody prsrvth typ of larg.mammal divrsity
many pst tims. For ampl, that was formrly ommon worldwid.
.ithin th past 3 million yars)
BIBLIOGRAPHY
1859). It was, h told a frind, ..as rsptalto modifid mnky as tht ur souls ar insrt
.ry Huly)
r (186) (th full titl is zi-
.aysknown y th shortr titl,
rglish ook aftr Dwin,s on
ls anatomially and in volu-
ljt).And it did so in punhy
t antitransmutationist Rihard
;orill,disovrdin th 1840s),
rala (rulingrain), in 1858.
misphrs'whih divid into a
..hippoampus minor.''
ltruding
ins, hatd own's hautur Shltons ojtha
..thr is vry lit_ Gro. o.
y stating that Clpz. Gorrrl. trL{N.
Dsmond, ^. 1997, Hul: From Dui!'s Disipl tl uolutio's High Prist. to work as a unifid v
Rading, A: Addisln-'Wsly.
winin struggl than
Gross, . G.1993. Hippompus mirror and man's pla ill natur:A as stlrdy in
th soial onstrution of nuronatom. ippr',m|>s
matril fom n or1
3: 40_415.
uly, T. . 1858_1862. on fossil rmir-rs of mn. Prtlatligsrlf tb RolI tria. his vi. has
I nstitut i l lf r t B rit i tl: 1Ii,420-422. work on th human
-h
1859. Sinnd rligiorr. Buildr,Januarv 1.5. frrd sidways in thi
1861. on th zoologil rlirtionsof man with th lowr nimals. howvr ltaind, it
Nturl Histor Ruilu: 67_84'
as a omplmnt to D
1863. uiden s to 's Pl i Nture. Llndtln: \/illims nd
Norgt. In rnt yars' ais(
L,vll,. 786. Th tiquit rlf . L.ln:Jhn ur,v. N4argulis (togthrw
Lyons' S. L. |997. ovining mn thy monkvs. Irr A. P. Bar, d., Thlms Sagan) lras n a st
Hr ltl's Pl i Sittd Lttrs: lttttr ssays, 95-l l8. arth as i1norganismi
Athns: Univrsity of Gorgia Pss.
so signifintly that it
Own, R. 1858. on th haratrs,prinipls lf divisin, and primry groups of th
lass mammalian. Jourl of tbc Prodigs tlf th I-i Soit (Zotllog)
ptdly hapd r
'l'_7. gists quat th Gaiz
2:
.W.ilson,
L. G. 1996. Th gorilla and th qustion of huml oigins:Th rlin qually xptdly st
ontrovrsy. Jlurl tlf tll is|or ll|-adiinc d llic! ,\is 57: vind of th ightn
184-2()7. -.D.
lLIoGRAPl])
l\4argulis,
L. |970. rtg
argulis, Lynn (. 1'99) for ,hlrtf th
l,ynn Mrgulis, a longtim profssorof gosinsilt th Univrsity of Mss- o tb Prmbri
200Q.Smbiot
ahustts'Amhst, is proprlv lratdfo hr stongildvoay of th oi.
rgrrlis,L., nd D' Sa
gin of ukurryotlls (lls witlr a nuleus) frotn trtorprimitiv prokary<lt ullti.Nw Yo
lls (lls withlut a nulus). Som vrsion of rhis thrlry gos bk to th
nintnthntury' whn it was pushd y th Russian lihn spialistKon-
stantin Srgivih rzhkovsk (1855_1921), and it was advoatd in th
1920s th Swdish Amrin ]vn !alln. But it lvas argulis wh took lvlarsh,othnil (
up th ida, now known aS te dosmbioti thor, offrd mirlsopi othnil harls al
vidn,rrd pushd it (in hr ook rigi oi ukroti lls, 1970) until nintrrtlr_tury An
it am orthodoy. Paody, and his on
A4argulis laimd that rtain funtining p:rrtsof th ll (organlls)ar rih fossil ds of th \
in fat formr prokaryots that hav n takn up and now funtion for th usd many of his fossi
nfit of th whol ukaryoti ll. Inludd hr ar mitohondria and arsh was orn in
hloroplasts' rsptivly'th powr plants of th ll and (in plants) th or- Shffild Sintifi Sh
ganells th2rtpfom photosyrrthsis.Dfinitir,prof of N{argulisposition trrop rturnd to b
am in th 1980s whn it was disovrdthat tlr DNA f thsorgzrrrlls is unl finnd th P
diffrnt from that of th ll's nulus ut ars signifiant similaritis to pditilns that ars
that of various prokaryots' for instan, in ing irular, hving th sm h was politilly astu
size, and ing l to pfrr th sam funtiotrs as th appropriat or- my of Sins(h s
ganllsor funtions similar to thos of th organlls' ogist of th U.S. G
argr-rlis'sthinking is dply inf]ur-rdy plriltlsophy of lif, wlrih onntioll to Lrosth
ss hrmony and mutualisrrrin natur rath rlrn fighting and arrtagonism. fir fud with dwa
Although som hav argud that th ndosymioti thory an st sn th narrringand dsri
in trms of ptur and slvry-th organllsy th main ll-for hr it is in this volr.rm).H dit
rathr a mattr of diffntlif forms oming togtl"lrin a sussfulatmpt Marsh's tams dis
rsb 71
,lF,uriutil'sigh Priest.
to work as a unifid whol. Sh ssvolution lss in trms of a ruthlss Dar-
winian struggl than s a pross asd on t sidways transfr of gnti
plain ntur:A as sttrdy in
()mpL|s matrial from on organism to anothr, arrid via miroorganisms lik a-
3: 40-415.
n. Proecligsrl|.th Rowl tria' This viw has lr rivd with som skptiism, although rnt
work ln th hurrrirngnom strongly suggststhat gnti matrial is trans-
u a r l 5 ' frrd sidways in this fashion. Sin Drwinism works on gnti variation,
''ith th lorr,ranir-rrls.T/.,c
howvotind, it is pobaly rtrto tlrink of this sort of phnonrnon
as a omplemnt to Darwinian volution than s an opposing viw.
. l-rlnln:\lilliamsand
In rnt yrs, :.rlsoguidd hr philsophy of hamony and mutualism,
n Murray. Margulis (togthr with Dorior-r Sagan, hr son from hr mrriag to arl
s. In A. P. Brr, d.,Thoms Saga) has n a Strong dvoat of th Gaia hypothsis, whih ss th
(t1r\'ssv-s,95_l l 8. athas an organism and as slf-rgulating'at last Llntil hLlmansdisruptd it
so signifiantlythat it ould no longr funtion proprly. .I.hisadvoay has
visin,nd primry grorrps of th
'f th I'itl Slit (Zoolog) ptdly hapd ritiism on argtrlis's had-rany onvntionl iolo-
gists quat th Gaia hypothsis with pr-hristian natur woship-and
I humrrlrigiIts:h rain qually ptdly sh is mor tilan prpard to wthr th stom' on-
d llid Sis 51: vind of tlr rightnssof hr aus.
-.D,
|LIoGRAPY
Margrrlis,L. 1970. Origi of uhroti alls:uidd Rsrh]mDlititls
for T|trf the rigi t! uo|utilof iro|lil,PIt,d iml lls
]nsat th Univrsity of N4ss- o th Prmbrirth. Nw Hvn, T: Yl UnivrsityPrss.
20'SmbiotiPlt: Ntu Llok t t,olutill,Nw York: Bsi Books.
hr strong adv<layof th oi-
a r g u l i sI, . . , n d D . S a g a r -1r9. 9 7 . S l t d T r u t h s : s s s l G i , S m b i o s i s d
oln mo primitiv prokaryot uolutit.l.Nr.York: oprnius Books. -.R.
f this thor1'gos ak to tlr
: Russian lihrrspialistKn-
), nd it Ws advoated in th
But it rvas Margulis who took arsh,othnilharls(1s31-1s99)
' it..thllrl,' offd mirosopi
othrril als rsh was t most influntial palontologist in lat-
'f ukrotic lls, |970) unt1| nintnth-ntuyAmri. H xploitd tlr walh of his unl, Gorg
Pody, and his onntions with th U.S. Gologial Survy to plor th
lrts of th Il (org:rrrlls) a rih fossil ds of th Wst. H Ws an nthusi:lstisupportr of Darwinisnr and
3n up nd now funtion for th usd many of his fssil disovristo prvid vidn in support of volution.
:d hr r mitohondri and arsh was rn in Lokport, Nw York, irr 1831' ws rind at th
th ll and (in plants) th or- Shffild Sintifi Shool of Yal Univrsity nd aftr thr yars of study in
v p r o o [ f e r g u l i s . sp o s i t i o n urop rturnd to onrrofssor of plontology at Yl in 1866. His
: th DNA of thsorganlls is unl finand th Pody usum at Yal and providd funds for th -rany
a r ss i g n i f i n rs i m i l r i t i st o xpditions that rsh snt to t !st' Although rsh Was a rsrvdmirn,
ling irular, having th sm h was politially stut and m a powrful figr.rrin th Ntional Ad-
l t t i o n si l s t h a p p r o p r i a t o r - my of Sis(h srvdas prsidntfom 188 to 1895). H was palontol-
:ganIls. ogist of th U.S. Golgial Sr-rrvyfrom 1882 o |892 and xploitd this
, hr philosophy of lrf, whih onntion to oost his wn privatly finand pditions. H ngagd in a
t h a n f i . g h t i n gl ] d n t g o n i s m . fir fud with dward Drinkr op ovr ass to W,strnfossils and ovr
iotithory an bst sn th rraming and dsriptitln of spis (salsrth lphatial ntry on op
:s y th main ll-fo hr it is i n t h i s v o l u m ) .H d i d i n 1 8 9 9 .
ogthrin a sr-tssftrl attmpt Marsh's tms disovrd mar-rynw spis of fossil marmals and rptils.
714 rsh
(wW
RN.
fl tinuous squnof v
ilfl
l{lr modrn hors, a squ
QuUs.
t1 IJ v i d n f v r l l r t t i l l n
E )(
t, t r n d o f i n s ds p
rfi,
^ww
although th tru storv
I{
4Jl(lt'w\
r s h l s <dl i s o v Jt
PLIoN.
in mon<lgraph.dl
PLloIPPUs.
from Grrrnv.ths\\
l_r
,( i d s h a d v o l v df r . , m
t-l \\ \^J
w s s u s q u n l f1l.l u n
)
ffiffi
n a m d e I a r g t t u m r.
il
o f g i a n t r l ym m m |
l ( l 8 8 6 ) h r r a tl h l r
#h t(
PRooIPPUs
(Ei|!4i'. ansto. Lik Huly,
muh fr.rth llirk ill g
N4arsh was n nthl
.RgF
A
'n
o n r n l r i m s I fr r i r h r l
Io,N.
ir
tf \-cY Lg)
natural sltion woul
,t*
rrsintllignWs
loIPPUs
/]'\ The ltrodutit.l d
(hilhcriuni.
t h t p a | o n t o l o g vp r o v
ing rain siz in th ou
il produtof nint
[
/,111
|l ffiF </
l{l \sss}-/ (\J
ILIoGRAPHY
.l
H r t l v . . H . I 8 3 8 .
oEo
t$
sol{IPPus.
l\ N .Y l r k : plrr
M . t s ho, . . | - 7 .I I
tr
ddrss Dliuardba
At
mwffii?ffi
'\itt t Nlshuill,
oN.
Iili
#h
hous& Tylor.]
\t w, |880. dtrlr
A m,ri . Wr shingttt
oRoIPUs"
P r a l l vl ,o l . 7 .
"fJ" |886.Diort
Y/eshingoll.D(,: Mr,
On a ltur tour t() th Unitd Stts'homas nry Lluly was owld ovr
y othniel Marsh,s olltion of hors fossils.To mak th as for volution in
puli letu,h used this reonstrutionof hosvolution(somthlngw
now know to muh mor ranh-lik),prditingtht soon a fiv-td
anstor would disovrd.T his dlight' soon thrftra spimnwith IyIasstintions
rudimntaryfifth tos was disovd.(arsh al[dit ohipps,..dwnhors.''
Mass tintions ar glr
It was thn disvrdtht spimnshad nfound arlir,althoughnot
rs of spis from di
idntifids horss.Th alirname, 1,rotbrium' taks prdn,
althoughoviouslyarsh's nam has stuk.) frotn rrrlrrginai t
o l o g i a l p r o s s s l.- o
ayrr-rnt or a lk m1
sdimnt. In ()ntrast'l
ss titios 715
lww
UPr oj!. Low6r ol..
His olltion of hors flssils(sfigur)gnratdWhat appardto on-
tinuous sqLinof volutin frtlm tlr snrll fotr-todrrrohippus to te
rodrn hors, :l squn that Tlrlms Hnry tlxly alld ..dmonstativ
'? vidnof volution'' (uly 1888,90). Th fossils smd to irrdiata
trnd of inrasd spialization y adaptation to running on th pn plains,
frww
although th tru stoy of th hors fmily turnd out to far mor ompl.
M;rrsh rlsodisovrd toothd irds from th rtatlr.rs,whih h dsrid
in a monograp6, ()dltlritbs\1880). Lik th ttr_knownrlltloptr
from Grmany' ths wr haild zrsimportant onfirmation of th viw that
irds had volvd from rptils. Th had of n of arsh's irds, Ihthornis,
was susquntlyfound to long to marin rptil. arsh disovrdand
ffiffi
nald a larg num of dinlsaursnd also studid th volution of group
of gint aly mamtnls h alld Dinort. In his n-rorrogaph on this group
(l886) h trad tlr origin of tlr Ungulata ak to:r hypothtialrtous
anstor.Lik uxly, h livd that th origin of ordrs and lassslay
muh furthr bak in gologial tim than th arlist known fossils.
]VIarshwas an nthusiastisuppotr of Darwinism, although h did not
sffiffi
()nnhimslf with th dtails of how volution rvlrkd.H livd that
natual sltiorrwould gradLrirllvfavr th irrrasingsiz of th rin -
aus intllignwas an importirnt adaptiv advntag.In his 1877 ddrss
T'h ltrodutir> nd Suessio of Vertbrt Lif i mric h argud
tht palontolog providd iar poof of this adaptiv trnd toward inras-
ing rain siz in th Ollrsof vllution.Mash's law of rain growth is a typ.
ffi8
ial produt of rrintr-rth-ntllryprogrssionisr volutiorrism.
B|LloGRAPY
Huly,T. H. l888. AmriAddrsss, tuith Lturo tStudof Biohg'
Nw York: Appltn.
Nlrsh,o. . 1877. llttrodutilnt!Sttssitl lf Vrtbr,ttLif i mri:
ddrss Dliurd bf<lrtll Atltt,rittsstlititlilr th dmtri.
A U s E so F M A s s x | N | o N s omparisln of tin
Th ida that mass tintions hav puntuatd th history of lif has n larg mtorit rtrl
rognizd xtintion
around flr a long tim. Th rason that most mass-tintionvntsour at
ruption of larg vol
mjor oundaris of gologial tim, suh as etwn th rtaous and
oinids with mass
Trtiary' is that thy mark major disontinuitisin th history of lif and thus
ar usful markrs for th sudivision of gologial tim. Ths tims w
ovious vn whn suh intrvls wr ing dfind in th arly nintnth liquid and last th
ntury. Howvr' th intnsiv study of th auss of mass tintions is thy would stay sus
a muh mor rnt sintht an datd to th puliation of a study y prhaps vn yarsal
.Waltr,
Luis Alvarz, his son and two ollagus in 1980 (Alvaz t al. dvastating for th p
1980). This group masurd th onntrationsof th xdinglyrar tra most osystms.Tl
mtal iridium in sdimntsthat straddld th rtaous-Trtiarytransition. though th food h
For th most part this mtal ours in tiny amounts in th arth's rust, ut whih would di of s
th Alvarzs nd ollgus found a dramati nrihmnt prisly at th snlight. It appars l
lvl of th mass tintion. Thy suggstdtht this iridium am from a nd. This original ki
hondriti nrtorit(on of th most ommon typs of n-rtorit) and got addition of various <
into th sdimnt from fallout in th aftrmath of a giant impat. Th iridium sit is vry unusuai i
anomaly has now n dttdin dozns of loations round th world, and This would hav b
this, togthr with th disovry of th impat ratr in th Yut5n Pnin- ntratd typ of aid
sula of io, has nsurd that thir hypothsis is now widly aptd. this ornr of Mi
Th ratr is loatd nar th villag of hiulu and is urid nath to hit.
youngr sdimnts.It may as muh as 180km in diamtr, making it on Th 1980sand 19
of th largst irnpat ratrs known on arth. impat with etinti<
In th original 1980 artil th link twn th impat and th rsultant plil to all tin
mass tintion was thought to hav om aout from th gloal darknss tintions in whih tl
that would follow suh a larg impat. Th nrgy of suh an vnt would spad vry 26 milli
pulvriz oth th mtorit and targt oks into dust and fin droplts of nation that is unlik
ss titios 7I7
x
lill
i l p I a n t s .I n o t h r x t i n i o n
o
animls. o
qr
various astronomial snarios Wr proposd whry th arth was su- of asalt lavas: arb
jtd to dvastating mtorit and/or omt showrs evr 26 million yrs. dioxid is a grnh
owvr, susquntstudy of th fossil rord and improvd ag dating of mosphr will aus1
th roks has shown that svral of th 26-million-yar tintion vntsdo (y ration with w.
not xist (for ampl, thr was no tintion eent 26 million yars aftr th most notiabl
th dath of th dinosaurs).Non of th tintion vnts that rmain show tion of Mount St. H
a rgular spaing. For mpl, thWas a mass xtintion 200 million yars mat. Howvr, su
ago, known as th nd-Triassi vnt, followd y smallr-magnitud- ar raind out of tht
tintion vnt 20 million yars latr in th arly Jurassi.Th nt xtintion that th ruption of
vnts did not our until th Crtaous' ovr 80 million yars latr, and gloal warming' sug
ths wr all vry minor affairs ompard with th K-T vnt. mor importnt li
Attmpts to link othr xtintion vntswith mtoritimpts hav so far gloal tmpraturl
provd rathr unonvining, mainly ausof a lak of vidn.For am- all lif on ath. Thr
pl, th Maniougan ratr of northrn Ontario, anada' is roughly 70% tion of flood asalt
th siz of th hixulu ratr. It is on of th largst ratrs known ut that lads to a runa!
sms to hav had littl or no fft on tintion rats. Th ratr was formd
220 million yars ago at a tim markd y vry low etintion rats. ow-
oNsoUN(
vr, it is worth noting that a giant impat rat datd to th oundary of
th Prmian and th Triassi has rntly n rportd from th northwst If th aus of mass
Australian ontinntal shlf, thus linking in tim a ig ratr with th iggst mass xtintions ar
mass xtintion. Howver, th laims for this ratr hv n tratd with portant turning poir
skptiism y th gnral sintifi ommunity, whih rgards th strutur vnts is that groups
as mor likly to of volani origin, and it rmains to sn whthr nan in th aftrnra
thslaims gain ptan. mammls in th ft
If th link twn impats and xtintions is gnrally wak' th sam for this mass t
annot said for giant volani provins. Th surfa of th ath is markd ially living in th s
by svral suh provins that ar omposd of layr upon layr of asalt nosaus Wr mam
lava flows that typially d 1 million ui kilomtrs in volum. Thes fornrs w s tody
ruptd from long fissurs or raks, and individual flows oftn dd nosaurs may also ow
1,000 ui kilomtrs in volum' a figur that is two or thr ordrs of mag- pard in th Triassi
.Within
nitud largr than thos sn during eruptions in historial tims. th yrs of thir histor}
past 10 yars improvd ag dating of ths volni provins has rvald trrstrial land ani
two rmarkal fats. First, most of th provins appar to hav ruptd in grops. Only aftr tJ
lss than 1 million yars; a gologial link of an y. hus th nvironmn- dinosaurs quikly ris
tal onsquns of suh ruptions would hav n onntratd into a In fft mass t
short intrval. Sond, many of th volani provins hv n found to into th vaatd ni
prisly oinid with tintion vnts. Indd, it now appars that vry of this volutionary
xtintion vnt in th past 00 million yars, inluding th mass tintions, way from ommuni
oinids with th ruption of a giant volani provin. This inluds th n in th rats of;
K-T vnt, whn th Dan Trap provin of India was formd. Howvr, oans ompard
not vry giant volani provin oinids with an tintion vnt.In othr highly sussfulgr
words, ther ar mor volani provins than thr ar tintions. ovi- shlls. Thy suffr
ously, it would ni and nat if vry provin oinidd with a mss - idly within a millior
tintion, ut th history of th arth is dfinitlynot nat. wipd thm out). I
How an volanism aus tintion? uh rsarh has fousd on th of millions of yars
limati fftsof th two major volani gassassoiatdwith th ruption tintion vnt also
ss tintios 719
l whyth arth ws su- of asalt lvas:rbon dioxid and srrlfudioxid. As is wll known' rln
lowrsvry 26 million yars. dioxid is a grrrhousgas' and th injtion of larg volums into th at-
c and improvd ag dating of mosphwill aus gloal warming. In ontrast, sulfur dioxid forms arosols
lion-1.arintion vnts do (y ration with wtr vapor) that lok sunlight nd aus ooling. Thus
n vnt26 nrillion ys aftr th most notial fft of mdrn lrg volni ruptions, suh as th rup-
.tiOn vntsthat rmain show tion of Mount St. Hlns in Washington Stat in 1980, is ooling of th li-
ss tintion200 million yars mat. Hwvr, suh fftsar dinglyshot livd austh rosols
d y a smallr-magr-ritud - ar rairrdout of th atmosphrin yar or so. Gologial vidn shows
Jtrrassi.T h n t t i n t ion that th ruption of giant volni provins is nrly always assoiatdwith
r 80 million ars l a t r , a nd global wrming, suggstingtlrat th uption of aron dioid may th
h th K-T vnt. mor inrportnt limati fft of volnism. owvr, it is on thing to rais
t mtoritimpats hav so far gloal tmpratur y a fw dgrs and ntirly nothr to wip out narly
a lk of vidn.For xam- ll lif on arth. Tlrrfor,many urrnt xtilltion snarios s th erup-
lrio, nad, is roughly 707" tion of flod aslt prvins s just triggr in a ompl hirin of vnts
th largst ratrs known but that lads to a runaway grnhoussnario.
)n rats.Th ratr was formd
:r low tintion rats. F{ow-
'atrdtd t<lth oundary of oNsoUENEs oF MAss ExINloNs
n rportd fom th northwst If th aus of mass tintions is still ativly datd, th onsquns of
n big rtr with th iggst mass xtintiorrs r aurrdantly lar. hy lrav long n rognizd as im-
rtr hv n traltd with portant turning points in tlr volutiorr of lif. An intriguing aspt of ths
y, whih rgrds th strutur vnts is that groups that wr insignifiant for th tintion ris to domi-
it rrrainsto L]sn whthr nan in tlr aftrnrtlr.Th quintssntialarpl of this is th suss of th
mammals in th aftrmath of th K-T tintiorr. amnrals had a long history
rs is gnrallvwak' th sam for tlris mss tintion' ut thy spnt this tim litrally and mtapho-
surfof th arth is mrkd illy living in th shdlw of th dinosaurs. only with th dmis of th di.
i of layr upon lyr of asalt nosaurs Wr mmmals abl to volv into tlr largr sizs and divrs ody
i kilomtrsirr volum. Ths forms w s today. Somwhat pradoially, th initial suss of th di-
rdividual flows oftn dd nosaursmay also OWsomthigto nlass-xtintionvnt.Dinosaurs first ap-
t is two or thr ordrs of mag- pard in th Trissi round 230 million yars ago' ut tor th first 30 million
.Within
; in historialtims. th yars of thi history thy wr gnrally only a rlativly small omponnt of
,olaniprovins has rvald trrstrialland animal omnrunitis,whih inludd m:rny othr ptil-lik
Ll1Cs appnr to hav uptd in groups. only aftr th nd-Triassi mss tilltion 200 million vs ago did
f an y.Thus th nvironmn- dinosaurs quikly ris to total dominan of trrstrialommunitis.
rav n ()nntrtdinto a In fftmass tintions provid lpportunitisfor the survivors to rdiat
provins hv n found to into th vtd nihs lft hind y th xtirrtspis.owvr, th pa
[d,it now appars that vry of this volutionry radiatilnis highly inonstant and vris in an intrsing
i n l r r d i n gr h m s s x t i n t i o n s . way fron-rommulrity to ommunity' For ampl, thr is a distint diffr-
rri provin.This irlluds th n itr th rats lf ror,ryof rarursthzrtliv in th wtr olumn of th
lf India was formd. Howvr, oans mpard with thos that liv on th safloor. Ammonoids wr a
ith an tintion vnt' In othr highly sussful.roup'distantly rlatd to squid, that had attrativ spiral
.rn thllr tintions. Ovi- shlls. T.hy sr.rffrd nulnrous xtintiolr vnts ut irlwys rovrd rap_
.inoinidd with a mass ' idly within a million yrs or so (xptfor tlr K-T tintion, whih finally
Iynot nat. wipd thm out). In ontrast, lams that liv on th saflor oftn took tns
h sarhlrs forisd on th of milli<lns of ,varsto rovr from xtintion vnts. Th nd-rln -
; sa s s o i t dw i r h t h u p t i o l l tintion vnt also shows n intrstingdihtltomy in th rovry rats of
720 rd Smith
was haratrizd y th us of rathr simpl mathmatial mthods that opd' whih appli
onald a good dal of ingnuity in thinking up approahs to iologially Ths mthods ar :
signifiant problms. This sortimsmt with disapproval from applid mth- ioral ology. ayn
.
matiians' who oftn prfr to us ompl mthods to solv insignifiant uolutil nd th
prolms. Maynard Smith ontributd importantly to arly work on molu- luid and informativ
lar variation and volution, using th thn nwly proposd nutrl thory as Aftr formal rtir
th asis for svral important puliations. Th most influntial of ths is Brian Spratt'smiroL
with his SussollguJohn Haigh on gntihithhiking (aynard Smith on molula variati
and aigh 1'974).This showd quantitativly how th spad of an advanta- work was vidnl
gous mutation rdus varition at linkd nutrl loi' y dragging a hro- atrial lls in natr
mosomal sgmnt alng with it. This onpt has lm vry important in ods for xamining th
rlation to th avalanhof data on ntural variaility at th lvl of DNA s- mmrs of atria
qrrns' sin a vally of rdud variaility an intrprtd as th signal DNA-squn lvl (
of a rnt fixation of an advantagousmutation. rmarkal ft of
Anothr fild to whih h md important ontriutions was th volution rputation as an ld
of sx and gnti systms' starting in th lat 1960s. is work hlpd fr N4aynard Smith al
this fild from its domination y th rathr woolly group-sltionist idas of tionary qustions' in
yril Darlington and Gorg Ldyard Stins, whih had had a svrly tion (th lattr trm
ngativ fft on its dvlopmnt. In partiular h mphasizd th prolm quilirium, and th
of th ost of sx, whry an othrwis sltivlynutral asxual variant wrot a sris of <
rising in a sual population xprinsa transmissiondvantagand rap- ogy, ulminating in l
idly sprads to fiation. Maynard Smith was th first prson to apprit 6rs Szathmiry dv
th diffiulty this poss for plaining th prvaln of sxual rprodution tions in volution, frr
among ukaryots. H mad numrous population-gntimodls of pro- th volution of lang
sssthat ould provid an advantag to s or inrasd romination' fts of msothliom
and h summd up th stat of th fild in his 1978 ook h uolutio of rsarh until his d
Se, whih is still th st survy of this sujt as a whol. Th study of th haratristially sti
volution of sx and rding Systms is now a flourishing disiplin within 2003 mting of th
volutionary iology, in no small part aus of aynard Smith's own work
|BL|oGRAPH
and his nouragmntof othrs.
A4aynard Smith's most influntial singl ontribution waS th dvlopmnt F i s h r ,R . A . l 9 3 0 . e
of th onpt of th euolutioril stbl strltg(SS),initilly in ollao. UnivrsityPrss.
N4aynrdSmith,J. 195
ration with th lte Gorge Pri. This flowd out of his lng-standing intr-
longvitof Dros
st in animal havior and his dsir to undrstand why animal onflits 1 9 5 8 b .S u a
usually do not nd in srious fighting. lt uss th prinipl that, for a trait 20_244.l-ondon
valu to rprsntan quilirium undr natural sltin,a nssaryondi- 1 9 7 8 . T h u
tion is that all possibl dviant trait valus ar sltivlydisadvantagous 1982.L|ulltt
UnivrsityPrss.
whn introdud t a low frquny into a population whr most individu-
1989.uolti
als hav th spifid trait valu. Dtrmining an SS provids powrful N4ynardSmith,J.' ..
mans of prditing th outom of sltion in ass whr frquny- volutionary dyn
dpndntfitnsssar gnratdy th iologial ontt, suh as s ratios ynard Srith,.|.,zln
or soial hvior. Although this pproah had n usd for for daling GetilRsr
with sx ratios' notaly y R. A. Fishr and.s7.D. Hamilton, aynard Smith Mynard Smith,J., an
oford: oford U
dvlopd SS thory xpliitly and applid it to many prviously intratal
volutionary prlms. It is n of th most sintifially fruitful appliations
of gams thory. A hug thortial and mpirial litrtur has sin dvl-
rd Smith 72
mathmatialmrlrlds that pd' whih plis SS mhods tO many diffrnt iologil prolms.
up approahsto iologially Ths ntthods ar a mainstay of muh of th thory that undrlis hav-
lisapprtlvalfrom applid math- ioral ology. Maynard Smith rviwd his ontriutions in his 1982 ook
mthodsto solv irrsignifiant uollltil nd th horl' of GL1ftls'whih is haratristiall rif ut
tntlyt<larly work on molu. luid arrd informiltiv.
vl proposd nutral thory as Aftr trmal rtirmntin 1985, aynard Smith startdto olloratwith
.lr
mtlst influntial of ths is BiarrSprtt'srniroial gntisgroup' thn at Suss.on th analysis of dt
i hithhiking (Maynard Smith on molular variation an]volutiorr in tria. An inrportant rsult of this
how th spadof an dvanta- work was vidn for muh mor hang lf gnti information among
urral loi, y dagging hro- atrillls in ntur than prvior-rsly livd.It lso gnratdnW mrth-
h . r s o m V yi m p ( ) r t a n ti n ods for xamining th ffts of infrqunt rominti<lnalhng among
:iailityat th lvl of DNA s- mmbrsof atrilpopulations On pattrnsof variation and volution at th
:n intprtds th signal DNA-squnlvl (Mynd Smith t l. 2000). Mynard Smith ahivdth
ol1. rmarkbl fat lf oming a ladr in a nw fild wll aftr stiriishinghis
ontriutionsWas th volution rputation as an ldr statsmnof volutionry iology.
l960s. His work hlpd fr aynard Smith lso mad many important ontriutiolrst gnral evolu-
'olly group-sltionistidas of tionary qustions' inludirlg suh topis as group sltion vrsus kin sl-
ins, whih had had a svrly tion (th lattr trm was invntd y him), syrnpatri spiation,puntutd
lar h mphasizdth prolm quilirir"rrn,
and tlr vllutionaryrol of dvlopmntalonstraints' H also
tivly nutral asul variant Wrot a sisof xllnt txtooks on various aspts of thrtial iol_
ansmissionadvantagand rap- ogy, ulminating in his 1989 ook uoltior Getis. Lat in lif h and
, t h f i r s rp r s o n t o a p p r i a t ors Szthm6ry dvlopd a st of spulatividas out th mjor transi-
vlnof sxual rglrodution tions in vllution'fom tlr volution f lif itsIfand th volution of lls to
lation-gnti modls of pro- th volution of languag.Maynard Smith suffrdinrasingly from th f-
. o i n r a s dr o m i n t i o n , fts of nrsothliomain th last two yars of his lif ut ontinud with his
s l978 bok /: uoltio o| rsarhuntil lris dath irr April 2004. is l:rstpuli ltu wils a rif ut
t as a whol. Th study of th haratristillystimulating talk on his atrial work, at th Dmr
.a flourishingdisiplin within 2003 mting of th UK Population Gntis Group in Susse.
of MaynzrrdSmith own work
|BLIoGRAPHY
tion. H also ws
Nlayr, rnst Waltr (7904_2005)
Ornithologists' Uni
rnst !altr N4ayr was a Grman-orn Amrin ornithologist who playd N4ayr's ontriut
a ntral rol in volutionary thory and systmatis during th twntith h fousd on th
ntury. H was a strong advoat of two idas: th thory of spiation thory. Sstmtis
y gographi isolation and th Biologial Spis onpt. ayr also tionally and am
mad important ontriutions to iogography, systmatis,and th study was part of a small I
of whol organisms during a period whn biology as a profssin am r- in population gn
dutionist. volutionary pro
N{ayr was born into a profssional Grman family. His fathr did in 7977, ialty, working los
and th family prind onsidral finanial troul during th 1920s. In approh a
Mayr's intrst in natural history strtd with his fathr. gan univr- H passionatiyadl
sity in 192' studying mdiin at th Univrsity lf Grifswald. Dspit that a population
high marks' N4ayr swithd to zool<lgin |925' H alrady had dvlopd ould our. In this
tis to profssional zoologists, of whih th most important wr with th making. on t|
rwin Strsmann at th usum of Natural istory at th Univrsity of onpt. This pla
Blin. Strsmann am Mayr's mntor. Undr his guidan Mayr om- spis.Th biologi
pltd a hD tn t926 on th uropan distriution and iogography of nisms and arrirst
th srin finh, Sris ri srius. Aftrwrd Strsmann hird ayr mhanisms istd
as an assistant. most important m
From 1928 to 1930 ayr plord Nw Guina nd th Solomon Islands, tio (\96) prsn
ollting for svral major musums. Aftr his rturn ayr sought a prma- mpirial and intll
nnt musum position' ut svralpromising offrs in urop faild to mat- ing volutionary stu
rialrize.Rathr than rmain in Brlin' Mayr aptd tmporary work at th nizd th Soityfo
Amrian usum of Natural History (ANH) in Nw York. H gan in uolutio (\947).
Jnuary 1931. In 192Walter Rothshild sold his orrrithologialolltion to vrsational sps l
th musum: 280,000 ird spimns'among othr things. Mayr was givn a ommon purpos ar
prmannt position to urat th nw matrials.This kpt him in Nw York. i\{ayr's impat or
nvr undrtook major fildwork again. N4ayr promotd vo
Whn th National Soialists ros to powr in Grmany, ayr hos not Arthur ain' among
to rturn hom. In 195 h mrrid Margart (Grtl) Simon (7912_1990), nar th spislv
and thy had two daughtrs. Grtl playd an important rol in Mayr's a- ipl of this approa
rr. Sh srvd as srtaryand assistantand also lookd aftr studntsand tionships and to us
visiting ollagus.In 1950 oth wr naturalizd as U.S. itizns. taonomi dision
ayr ontinud at th AMNH until 1953, whn h aptd an Agassiz dmd suffiint to
Profssorship at th Musurn of omparativ Zoology (Z) at arvard thought to adapt
Univrsity. H srvd as dirtor of te Z fm |961 to 1970. For ayr that mad a group
th arvard appointmnt Was a symoli hang in status. H was now a full mriai taonomy a
profssor, lik his mntor. yr am an mritus profssor at Harvard in Profssionally
1'975 and rmaind in amridg thraftr' with th Z as his bas. ology. On th on
As an ornithologist ayr spializd in irds of th southwst Paifi, struggld to onvin
aout whih h wrot marry thnial studis and svral fild guids. From dat in iologial sr
th 1930s ayr was ativ in th Linnaan Soity of Nw York nd on- struggld to draw tl
triutd to suprvising many dotoral studnts.H als mad onsidrabl luding volution.
ontriutions to svral ditions of Jams Ptrs'shek-list of Birds of th iology in th lat
World. Aftr Ptrs did in 1952. ayr suprvisd the beck-lisf's nxt di- bodis and othr gr
r 725
lLloGRAF
l B Bok, W. 1994.rns
vlution.Bil/
2004.rnst]
12]l:67_65|
ayr, . 1942' Sst
UnivrsityPrss
1 9 5 . t h o
Hill.
1 9 6 . i m
1982.Th G,
5 t r s m a n n 't ' ' 1 9 / 5 '
Hrvard Univrs
}yIm
Inhts Ssttltiltis
"td thtl rigi of Spis(1942I, rnst Mayr malmuh of
suh phnomnaas ..ringsof ras,''whr mmrsof touhingsuspisan Th trm meme \^/
intrdut whn th nd populatitlnsirl k trdttlulr,lh. is ..o- ish G t<ldsri
dutivisolation.Hr w sspisir-rth mkir.rg. If th middl populations r units f iologi
(of th warlr PhIklscopustrohiloids)wr Iiminatd,w w.,ulj huu.
popularizd, as vi
two sparatspis'Th suspisin th ring'ar = uirids;I,=ldlotui;
th spawning of a .
T=trohiloids;:obsrtus; andp:plmbitrl,ls.Th rss.hathdra'
in th distrit twnth wstrnS:ryanuntins irnd tlr YnisiRivr, is graphs. ms an
where ui r i dlzsanl p lu it r sus.,u..lop. lifs, hairstyls,
mms to srv as i
any systm witlr di:
tion), sltion' an(
supportirrgth fild. His vision for systmatisis st prsntdin ethods Lik organisms,id
d Prinipls of S1t57g7nti
Zorlllg1,(195), rvhilr aln standrd train. ally inundatd with
ing manual. (othrwis rv wtlul
In th sotld hlf of his Iif, ayr Wrot tnsivIyon th lristory arrd with h thr fr
philosophy of io|ogy, spially on thms rlatd to natttral history and and get rpliatd,
volutiorr. His ook Th Groth of Billogi"ll Thlght (19s2I was a signifi. through imitation, i
ant ontriution to this topi. Drwir-r m l.]nlmportant sr.rjtfo sions t th dinnr l
ayr' as did many othr ky volutionists of th past 100 yars (.g.'Karl lt is important n
Jrdan). ar's ontritionss histlialrntrdon two thnrs.First, h th idas. For m;
me 727
BlLloGRAPHY
Bok, W. 1994. rnst ayr, naturalist:is ontributionst systmatisand
volution.Billogd Philosopb9: 267-327.
2004.rnstMayr at 100:A lif insidand outsidof ornitholog.Th uh
121:637-651.
ayr, . 1942.Sstmtisd th rigi of Speis.Nw York: olumia
UnivrsityPrss.
1'95.thodsd Priciplsof SstmtiZoolog. Nw York: MGraw-
Hill.
1963.Aiml Spiesd uoltio.amridg,MA: BlknapPrss.
-hoght.
1982. h Groth of Biologil amridg'A: Blknp Prss.
Strsmann, . \97 5. ritholog:From Aristotl to th Preset.amridg,MA:
Harvard Univrsity Prss. -l..
m
12),rnst Mayr mad muh of
Th trm mm was oind y Rihard Dawkins in is 1976 ook The Self-
l r s t l f t t l u h i n gs, u s p i s n
ish Ge to dsri units of ultural volution (as opposd to gnes,whih
lak nd touh' thr is rpro-
l i n g .l { t h m i d d l p o p u l t i o n s
ar units of iologial volution)' In th intrvning yars th trm has n
liminatd, w would hav popularizd, as vidnd y its inlusi on i he oford glish Ditionr,
are = uirids; L: ldlotui; th spawning of a ddiatd (and now-dfunt) journal, and svral mono-
itrus. T ross.hathd ara, graphs. Mms an suh things as songs' tools' styls of drss, rligious
tinsalrd h YnisiRiv, is lifs, hairstyls, aphorisms, and myths. Dawkins originally mployd
..Univrsal Darwinism',-th thought that
mms to srv as an instan of
any systm with disrt units that hiit diffrntial fitnss vals (varl'a-
tion)' sltion,and rpliation (hrdity)will volv y Darwinian mans.
'!7
s is st prsntdtn thods Lik organisms, idas appar to hibit ths haratristis. ar ontinu-
whih ama standard train- ally inundatd with idas. Thankfully, most of this information is disardd
(othrwisw would hav mmory ovrload). Thus idas ar in omptition
xtnsivlyon th history and with ah oth for survival within our minds; th fit ons rsid in our minds
rltdto natural history and and gt rpliatd, whil th unfit ons gt disardd. Rpliation ours
tl Thought (1'952)was a signifi- through imitation' it through th writtn word, dmonstration' or disus-
]am an importnt sujt for sions at th dinnr tal.
lf th past 100 yrs (.g.' Karl It is important not to onfs mms' whih ar idas, with th rfrnts of
:ntrdon two thms. F.irst,h th idas. For ampl, th mm rro|Uhdshrpnig is the id of a mannr
728 dl
of produing an arrowhad, not th rrowhad irslf. his distintion pr- Th stain of studv
srvs th gnotyp/phnotypdihotomy witi iology, or, in Dawkins's ausd outs of sv
trminology, th diffrn twn rpliators and vhils.Rpliators ar This prinlay b
th ntitis that opy thmslvs and od fr struturs that intrat in th of Saint Thomas in I
.,from th bittr strr
world (som thorists prfr th t,erm itrtor to uhile). This :liss sv-
ral points that distinguish mmti volution from ompting onptions 1 9s 4 ) .
of ultural volution. ultual volution is oftn sll s simply improving th At th monastl.
fitnss of th goup or group tnms.For xrlpl,fishhooks ad arrow- oftn-sad tasks of ;
hads volvd to onvy a fitnss advantag to th groups and individuals for.rndhim work as z
who usd suh artifats. ms, howvr, an quit sussfulwithout vr, rquird an
onvying fitnss advantag to thir hosts, ven to th point of birrg dl- oth tims. Aftr tlr
triolls to thir vtors. Dawkins famously lims tlrat rligion is a perriu- Vinna to obtain d
larly sussfulut toxi mm. Mms ar fit wlrn thy hav high fidlity th pratial dn-ron
and high feundity unto thmslvs.Aording to proponents' lms hold lnost rnt wrk in
impliations for our onption of ultur and human minds. N4ndl's talrr
Th srrssof th mme mm' howvr, lis its suss as a trm of art Vinn from athol
in th aadmy. Critis hav idntifid svral problrrrswith mms, nt th ing ndl's tim ir
lst of whih is that it is simply stitlgan old id il-lnw pkaging. Pr- hav krrown of this
aps it is not th mms thmslvsthat ar xplirnatory ut th undrlying trs aquird y plal
psyhologial mhnisms rsporrsilfor th sltion and rpliation of ral loals. Latr
tain mms tht ar usful for xplaining ulturl voiution. Furthrmor, suport for th inh
it is unlar tly what a mm is. It is asy to oll up with xanrplsof l21ntransmutatlons.
mms' ut a lar irnd aptd dfinition rmains lr"rsiv. For xilrrrpl,is possibiiity m11tiO
a mm a part of Bthovn symphony or th rrtirsynrphorry?Until an otanistsJosph KoJ
operational dfinition is found, tl.lfruitfulnssof this onpt rtllins qus- sking to stlish
tional. Finally' som hav argud that th lk of fidlity of mms, as wll hyridization.
as thir smingly nondigital natltr' srrggstsa Lirmarkian rathrr than Having rturnif
Drwinian volutiorrrypross. Th tru impat <lfmms for undrstand. tst varitis of tlr
irrg ultural volutin rmains lagly urrdidd. lryridizatitl11pr
im, h plairrl, u
BIBLIOGRAPHY
hyid progny pp
Blakmor,S. 1999.The m hi.Nw York: Oford UnivrsityPrss. tion with rtainty,
Dawkins, R. \976. h SlfishGe, ()ford: Oford UnivrsityPrss. drtak this progril
Dnntt,D.1995.Dttt'i'sDgrousIt!,I-ondon:Prrguin. -].Z.
hzrratrdiffrn
to hiv stirtistia
..fr.rahing'
was
ndl, Grgor (t822_884) .
portant qlrstion:
..th prist who hld organi forms'' (St
Grgor Johann ndl hs n lld th ky to vo-
lution'' (isly \959'205), ut in lris own day fw iologistshad vr hard th ould forr th
..rvrtd
of him, hrls Drwin inludd. His faril was poor' ut h souglrt sup- turnd or
Port in a monasty whr fortun shon upon him y giving him tlr mans vidntly this rva
to arry out his rmarkal and aritiousxprinrnts. tsting his rsarh
lvIndl, th firstorn of Anton ndl and Rosin ndl (n pulishd. Not unti
Shwirtlih), ws isd in Grmn-spaking Silsizl in th littl vill:rg o{ rigi of Spis.
inzndorf. At 11 lr ttndd slrols far from hom, and at 18 h rrtrd sgs prsntingrr
th Philosophial Institut in Olniitz, ompltil-tgl]is ourss th in 1843. was attaking a tar[
dl 729
ad itslf.T.his distintion p- Th strain of study plus tutoring to provid his own funds had rnanwhil
ithin iology, or' in Dawkins's ausd otsof svrdpssionand tndd tim k honr to rovr.
rrsand vhils.Rpliators ar This prin iay hind his dision to apply to th Augustinian Monastry
|r sttlturstlrat intrat in th of Saint Thomas in Brtinr-r.H knrv, h plaind, that h nddto frd
. t o r t o u h i l e ;T. h i s a i s ss v - ..from th ittr struggl for istn''
that hd bn his lot hithrto (lltis
,n from ompting onptions t9 s4\.
n snas sirnply improving tlr At th monastry r-rdlprt>vdunal to op motionally with th
xampl,fishhooks and arrow- oftn-sd tasks of a prist, ut his aot' th undrstnding yril N"P,
l to th groups and individuals found him work as a sustitut tahr inZnaim. Tah rtifiation' how-
:n b quit sussfulwithout vr' rquird an xamination. This lr took twi but though stssfild
vn to th plint of ing dl- oth tims. Aftr th first attmpt th aot snt Mndl to th Univsity of
l a i n l sr h t r l i g i o n i s p r t i u - Vinna to otairr adqut foundations irr th sins.Thr h assistdin
fit whn tlry hav high fidlity th prtial dmonstrtionsfor th ph,vsisourss arrd was posd to th
ing to prponnts, Inms hold most rnt work in plarrt ytology.
d hu-ranminds. ndl's tahr. th otanist Franz Llngr' Was a vitim of attaks in
lisits sussas a trm of art Vinn from athlli quartrsfo his advoay of spistransllltationdur-
lprolms with mms' not th i n g M n d l ' s t i m i n V i n n a ( 1 8 5 1 _ 1 8 5 3 )a n d s u s q u n t l y
. ndl must
old ida in nw pakging. Pr- hv known of tlris nd of Unge's transplant xpimntsin whih hr-
planatoryut th undrlying trs aquird y plants in novl lraittswr lost on rturning to thi natu-
h sltion and rpliation of ral loals. Latr ndl rpatd suh primnts and found in thm n
;r.rltural volution. Furthrmor, support flr ti-rinhritanof aquird hratrs,thus ruling out Lamark-
iy to om up with ampls of ian transmutations.Spismultipliation y hyridization, howvr, was a
. m a i n s l u s i v .F o r m p l , i s pssiility mntiond y Ungr, for h knw th wok of th Grmn
. rh ntir symphorry?Until an otanistsJosph Klrutr and rl von Girtnr' who had hyridizd plants,
l s sr ' f t h i s o n p tr m a i n s q u s - sking to stalishwhthr spisrmain onstant or an hang through
Lakof fidlit1.of m.ls,as wll hyridization.
its a Lamarkian rzrthr thzrn a Having rturnd frorrrVinna in 185, Mndl gan th following yar to
npat of mms for undstand- tst varitis of th dil pa for onstany of typ in prparation for th
dd. hyridization primnts that oupid him twn 1856 and |862. Ifis
aim, h plaind, was tO ,.dtrminth numr of diffrntforrns in whih
hyrid Progny appar' prmit lassifiationof tlrs forms in ah gnra-
rk:oxiord Univrsit,vPrss. tion rvith rtilinty, and asrtzrinthir numril intrrltilnshi1rs.'' To r.rn_
'fordUnivrsityPrss. drtak this program h hos varital diffrnsadvrtisdy nonlnding
rdon:Pnguin. -.J.Z.
harat diffrnsthirt ould l-llrly distinguishd' Awar of th nd
to alrivstatistiallysignifiantrsults,he grw larg populations' Th task
..far-rahing,''
was ut h warnd tht it was th only way to solv th im-
portant qllstion: ..th
dvlopmntl history ftiklugsgeschihte]of
pristwho hld th ky to vo- organi forms' (Strn and Shrwood 1966, 2). If som hyrids rd tru,
ay fw biologistshad vr hard thy ould form th asis for th dvlopmnt of nw spis'ut if thy r-
y Was po()r' ut lr sought sup_ turnd o ..rvrtd''to th origirrtingforts, no rrvlywould rsult.
on him y giving him th mans vidntly this was th qustion h st out to solv in 1854 whn h gan
prlmnts. tsting his rsarh mtrial. rls Drwin's thory Ws at that tim rrn-
dl nd Rosin ndl (n6 pulislrd.Not until |862 did N{ndlrd th Gman tanslationof th
rg Silsi in th littl villag of origin of Spis. ndl's most frqunt marginali thrin fr to pas-
ror hom, ad at 18 h ntrd sagsprsntingnd opposing tlr dotril of spial rirtin.H Drwin
ltinghis ()urssthr in 1843. Was attaking targt rht Mndl, it appars,did not pt ithr.
70 dl
Th ontt of Mndl's famous primnts Was thus not Darwin's thory prsnt in th hy
of volution. It was instad th dat among Grman-spaking iologists' sv- lls ar ithr of
ral of whom, lik Ungr, had spousdlmntsf th spulativtahing ominations of p
of Nturphilosophi, prinipally th notion lf th sussivdvlopmnt of ny of th two kin
highr and highr forms of lif ovr an tndd priod of tim. On of th tios that h found
Augustinians in Briinn, Matou Kl5l, also supportd a dvlopmntal viw of gg lls of th sar
spisorigins' ut unlik his frind Mndl, Kl:il was a spulativphilos- th old ida of on
ophr who nvr arrid out a rigorous sris of primnts. Mndl was a nw variant will
drawn hifly to th xprimntal litratur of th Grman hyridists. His hif was also wrong. N
rsour was arl von G;irtnr, who, in his ook F,periments d bseru- dirtly rlvant t<
tios of Hbridiztio i th Plt Kigdom (Vrsuh ud Belbbtugen Darwin had found
br di Bstrdrzgug im Pflzereich' |849), found that frtil hyrids rgation is thus s
yildd progny that tndd soonr or latr to rvrt to on or th othr of th Also from sgr
originating spis. Gzirtnr listd oniy a fw hyrids that appard to rmain in ll sorts of o
onstant in thir offspring, nd it is signifiant that ndl rprodud many pndntly of n :
of ths hyrids and found that thy did rvrt. ndl wnt on to hoos tion with this pro
hawkwds (Hirim) aus of th multitud of forms found in th wild. laim that domst
Although th otanist arl Nigli nouragd him in this dirion, it was not orrsponding wil'
h who prsuadd ndl to hoos this diffiult gnus' as is oftn assrtd. lif.'' Instad, h o;
Mndl rasondthat polymorphi gnramight ow thir multituds of spis Th aptan
to hyridization' ut of a diffrnt kind from that h had found in his lassi adhrn to nonb
xprimnts with th dil pa (Pisum). rativ rol of nat
ndl plaind th rsults of is Pisul primnts y suggsting that lnding stp' thr
thr must a pross of sgrgation (s figur) of th diffring potntials frns littl y li
gnetisinto Darw
|BLIoGRA
dtrN RuD Y.{.Lgt,tl{'flNLD
Batson'!' l 909. ,\
UnivrsityPrs
i s l yL, . 1 9 5 9 ,D
London:Vitor
YR q G:irtnr,. von.184
P|lzerih' S
Iltis,rs. H. 1954.(
fl.#w fliP)
2I*24. Reprt
hiago:Univ
oly, R. 1985.orig
orl, V. 7996' Grg
Strn,.' and . Sh
Blo&.Sanrn
,=.
.siling,
F. 1991.il
P.P|f-l
dil Gti
www.Mndlw.
nglishand Gr
othrs'th follo
Pul' D. B., and
lfilliam Batsonws ndl's mouthpiin Britain in th arly yrs of th prati,1900
twntithntury.his diagram shows his rPsnttionof Mnd['sfirst law,
-l
th law of sgrgation.(From Batsln 909.)
edl 7.1
Wasthus not Darwin's thory prsnt in th hyrid plant whn th grm lls ar formd suh that gm
:rman-spaking iologists'sv- lls ar ithr of on kind or th othr, not mid. Thn if in frtilization all
nts of th spulativtahing omintions of polln lls and gg lls ar possil' populations of prog-
th sussivdvlopmnt of ny of th two kinds will yildd that will approximat to th statistialra-
.!hr
d priod of tim. On of th tios that h found in his primnts. unions r twn polln and
ortda dvlopmntal viw of gg lls of th sam kind, oth originating forms ar rovrd pur. Thus
.16l ws a spulativphilos- th old ida of on taintd, always taintd Was Wrong. Th assumption that
; of xprimnts. Mndl was a nw variant will simply dilutd y rprodution with th normal form
r Grman hyridists. His hif was also Wrong. Nw ominations of haratrsan thus prsist. This was
,ok primnts nd bsrut- dirtly rlvant to th prolm of th slmpirxgffect of outrding that
(Vrsuh ud B lb h tuge Darwin had found dunting for his thory. Mndl's onpt of grminal sg-
849), found that frtil hyrids rgation is thus sn in rtrosptas th ky that Darwin lakd.
'vrtto onr or th othr of th Also from sgrgtionflowd th possiility of omining hrditry trits
y r i d st h t e p p a dt o r m a i n in all sorts of ominations and dmonstrating that th traits hav ind-
tht Mndl rprodud many pndntly of on anothr. ndl idntifid th sour of hrditry varia-
rt. Mndl Wnt on to hoos tion with this pross of rcombitio. Trefor' h ojtd to Darwin's
rd of forms found in th wild. laim that domsti animals and ultivatd plants ar mor varial than th
..hangdonditions of
him in this dirtion, it was not orrsponding wild spis us of th fft of
ult gnus, as is oftn assrtd. lif.'' Instad,h optd for th gratropportunitis for ross-pollinations.
t oWthir multituds of spis Th aptanof ndl's work in 1900 am along with an unritial
hat h had found in his lassi adhrnto nonlnding haratrs' thus promoting skptiism toward th
rativ rol of natural sltion.If nw hratrsar produd in on non-
primnts y suggsting that lnding StP'thr is no spa for natural sltionto aumulat slight dif-
1ur)of th diffring potntials frnslittl by littl. This matrially dlayd th intgration of ndlian
gntisinto Darwinian thory.
BIBLIOGRAPHY
Y(.Lltl {'Rllv|<L}
&w'@
Batson,!. 1909. dl'sl,riiplsof Hrdit' amridg:mridg
UnivrsityPrss.
isly,L. 1959.Drtai,stur:uolutio d th Who Dislurdlt.
London:Vitor Gollanz.
rfl&
hiago:Univrsityof hiagoPrss,1985.
oly, R. 1985.origis of ndlism.2nd d. higl:Univrsityof higoPrss.
orl, V. 1'996.Gregordl:Th First Gtiist,Ofrd: Oford UnivrsityPrss.
I'
Strn,., and . Shrwood'ds.1966.Th rigin rlf Gtis: dlSour
qr dw BooA.San Franiso:.W. H. Frman.
Wiling,F.1991. istorialstudy:JohannGrgorndl, 1822_1884.llurl of
dil Gtis40: 1_2.5.
& t
www.ndlw.<lrg. Inludsth nglishtt of Mndl's186.5papron Pisumin
I nglishand Grmanand .,ssaysand ommntary,,' whih ontins,among
othrs,th follwingssays: Oly, R. 1997.ndl,Mndlismnd gntis;
YR Yq
: Paul,D. B.' and B. A. Kimmlman.1988.Mndl in Amria:Thoryand
pratie'190-1919;and Sapp,I.1990. Th ninlivsof GrgorMndl.
Britainin th rly yars of th I -R.O.
.sntationof Mndl's first law, i
72 iber
ologial lationsl
Nlihnr,harls D. (b. 1918) u n d r s t a n da n d i n t
Chrls Mihlrer, ntomologist at th Univrsit1'of Kanss, is th world's ount of th nu.l
lading prt ()n th lassifition,morphology, natural history' and volu- nr as ..irquit' g
tion of s,spially th divrs slitzrryand priritivly soirrlbs whos polmis...[utli
iology olltrsts with tlrt tlf th familiar horrylr(pis tlliierl). tc_ might s;l dadly.''
nr has usd his road kr-rowldgof ths insts tO tst idas aout th Mihnr's ordia
volution of inst soitisand to hlp modrniz th sinof taxonlmy. man of sin,sp
trr-rdhis marr studnts lrav trvll tnsivh. in th Ur-ritdStats, h has ondLltdfil
Afria, Asia' urop, nd Latin Ami to tlllt nd osv s irr thir ntomologists in mu
natural haitats.As a rsult of this work th sh:rvlm modl tatln yrs Mihnr Ws
for omprativ studis of volution. Sins,on of onl1
ilrnr's rsrh is a prmir xrpl lf how volutionr1,iology uss lristory whn h wa.
ullparativ studis of lrvior and mophology ttl undrsttrdth origin of Soity for th Studl
nw trits nd tlr divrsifiation lf spis.Baust>fihrrr'sw<lrkand his pothsis tstingand
nOuragmntof young rsrhrs,thousands of spis lf s n road knowldg of
idntifidwith rtinty.This satisfisth first rquirt-nnt of ltlrpaativ-
sr: aility to idntify tlr organisn-rs. In additiolr, ihnr's work trats th BIBLIOGRAP
phylognti rlationships of diffrnt groups of s' summarizs knowldg H u ] l ' D . L . 1 9 8 8 'S l
of thir iology, nd plains how to rr and osrv thm in th laratory. M i h n r , . D . l 9 6 '
ilrnr'sm<rrthan 400 r.rlitions, inluding two maior bl<lks on s, 12:1.51-172.
|/. lnL'J|)
a a vrital nylopdi t>f inforr-ntirn that has n ploitd by Nlih-
200.h ts
nr and othrs to tra th volution of suh phnomna as parsitism and th '!7st-rhrd,
M. J. :
transition from solitry to soial irr group-livinginsts.His taonomi insight I n M . . [ .R n , d
has nrd him ldr in th tlrorv anl prati of s),stmatis,tlr str'rdyof Smithsniirn Inst
lilssifiationardrlationshipsamong groups of organisls.At th ag of 81,
Mihnr pulishd his magrrum op.he Bs lf th World (2000), a 91-
lvlillr, StanlyI
pag tom that trats 1,200 gnra arrd sugnriland mo than 16,000
spis irr a r"rnifidlassifitiorrd surmrizsmulr of th urrlrt informa- Stanl,vL. Nlillr sp
tio on this implrtant group' insts tllt ar ssrrtiltl ologil and human vrsity of alifrnia
ommunitis as pollintors of flowring plnts, inluding many rops. rakthrough xpr
ihnr's intrstin s gan whn as oy lr gan to Ollt insts sarh of pioti
nar lris hom irr southrn aliforni' l-rtlurag1 y lris moth, an al- gthr lvirh lan1'
tur naturalist, and by P. H. Timrlk, an authority n taonmy, h stps and hmial
pulishd his first ssay on s at th g of 16. FIis ontagious nthusiasm knlrn,ldgrnnt of his
for s has inspird a larg numr of studnts and ollagusirnd has put of-lif fild, ln \97 |
t Univsit of Kansirs On th map as a entr for rsilhand sil-t- to th Natilnal Aat
tifi ladrship. Mihnr also wrrt importent ssays on th taonom of his rol in th r:rti
uttrflis,although ths ar now lss known than his work on s,and h of t,if (ISSOL), th
Was at th ntr of ontrovrsy during datsovr numril tironomy in illr Awrl to
th l950s nr'] l960s. Alog with Rort Sokl, 'hos vilvs wr sotll_ In l9-52-1953,..v
w h a t r o r r a d i a l , M i h l l r a d v o a t d t h u s o f o j t i v ,q u a n t i t t i v and Nol lauratF
dat in lassifitions (meril tolm) as an antidot tO som of th to simulat th ond
ovrly intuitiv rrd lss sirrtifipr<ldursf tlr tim. But h also - primnts h was
l i v d t h a t q t l a n t i t a t i v l n t h o d s . . t l l t t s tn o t l l o w d t o s P a r a t t h s s - on th trlrgnof
. . A l k s a n d rI . o p r :
tmtist from the organisnls thmslvsfor only v.knowing tlrir hvior,
illr 7
s a mitur of gassmimiking onsquntly, illr's rsarh from th 1990s until his dth was dvotd
r and Watr rprsnting th to th srh for th onstituntsof a pr-RNA world that wr asirto syn-
:tri dishargs that stood for thsizand that ould hav volvd into th onstituntsof th RNA world.
rk,illr analyzd th aqu. Svralsuh andidatswr suggstdy Millr nd his group.
p r o d u t so f t h i n r r a t i n g t - It is important to not that from its inption illr's rsarh on th origin
lnthsisof organi ompounds, of lif was tightly linkd to th widr ontxt of th sarh for lif on othr
protins.Signifintly,th r- plants.Aftr th founding of th National Aronautis and Spa Administra-
all numr of iologilly rl- tion (NASA) in 1958, th young Millr Was among th main ontriutors to
4h yilds in th xprimnt out th stalishmnt of th fild of oiology, latr to b alld astoiology.
gani sustans.Glyin and Svralof Millr's latr primnts lnt rdnto th hypothsis that th po-
ins,wr th major produts. tntial fr prioti hmistry xists byond arth, prhaps undr orrditions
:hrsin th 1970s, it was found prvalnt on on of Jupitr's moons' uropa' and possily on othr Jovian
lativquntitiswr otaind moons.
arth Was anlyzd.Sin mt- In ordr to disredit th study of th mrgn of lif, rtionists oftn
t h s o | rp | a n t st,h i r o r g n i rly on th mpirial datsamong origin-of-lif sintistsovr spifipr-
similar to th Millr ratlons bioti snarios and on th many thortil and mpirial diffiultis still
fad y rsarhrs.Millr, who strongly dnound rationists for pro-
ronstratdth synthsis und moting ignoran out sin' nvr doubtd that a hmil solution to
nuli aids, for ampl, th th prlrn f th mrgnof lif will vntually found.
nof a primordial hydrogn-
|LIoGRAPHY
sis has n qustiond' and
anis wr importd to arth Fry' I. 2008.illr' StnlyLloyd. In N. Kortg'd.,Nez Ditiorof Sintifi
Bbgrph,vl. 5, 154-158.Dtroit:harlsSrinr's Sons.
artils.illr nvrthlss-
illr, s. L. 1953.A produtionof amino aidsundrpossilprimitivrth
on arth might hav istd. -l.F'
onditions.Sin1'17:528-529.
r sudd in dmonstrating,
' aids nd nuli aid ass in
1953..atmosphr.''
imiry
ption lf th origin of lif' a-
mo-
l f o r m l l s ts l f - r p l i t i n g Mimiry ours whn th faturs of ln spisrsml ths rf anothr
ost fundamntalqustion that and, through that rsmlan,onfr som advantag on th mimi. Mim-
ntimatrial tht ould hav iry is a striking phnomnon in natur; its study has plyd an impotant
primitiv living systm. Millr rol in volutionary iology from th rlist argumnts aout ntural si-
r fild that a watrshd in th tion, and svralhallngingpuzzls about th volution of ompl rnimiry
lworld in whih RNA mol- systmsrmain to solvd.
rs atalysts.Th ida that RNA i m i r y i s r a m p a n t i n n a t u r ( s W i k l r 1 9 6 8 ; R a n d a l l 2 0 0 5 ) . I n i t s
ry arly in th volution of lif simplst form' mimiry ours whn a partiular fatur of on organism r-
-sliorgl, Franis rik, and smls a vry diffrnt organism. Th most familiar xampls ar ass of
r ws th sol playr in th vry aggrssivmimiry' whr a prdator lus its pry with a prt of its wn body
l s a f t rt h s u p r i s i n gd i s o v r y that rsmlsth food of th pry. Th flshy rd proturan on th tongu
i'NA molulsthat in addition of an alligator snapping turtl mimis a lodworm and lurs loodworm.
; arrying out th funtion that ating fish into th turtl's mouth. Th lurs of anglrfish and atfish ar othr
y by protins. wll-known ampls. In ths ass' th mimti fatur onfrs a lar n.
vvr'Millr, as wll as his ol- fit to its ownr nd th volutin of th fatur rats no disadvants for
am onvind that lif ould th organism ing imitatd.
alizdthat th prbioti synth- Th mor ompliatd flrm of mimiry ours whn on organism rsm-
: itslf was highly improal. ls a similar on' as in on uttrfly rsmlinganothr' ut 'ith th twist
76 imir
tht th rsmlanoffrs a nfit to th mimi Llt potntial disadvan- fnding food itm. It
tag to th nrodl (th organism ing imitatd)' This happns whn modl of a paltal spi
and mimi hav vry diffrnt l.laratristisor ologial rols and th su- nss through a rdu
ss of th mimi thratns th modl. For ampl' on oral rfs in th Pa- lowr risk of prdati
ifi, small fish alld ..lanr wrasss'' find food y piking rrrstaan mimi and modl zrr
toparasits from th sals of largr fish, whih visit spifi spots on th Dspit th simpli
rf whr th wrasssour' Thr is anothr spisof small fish, a lnny, rl irpotnt qLlst
that mimis th shap and oloation of th lanr Wrass ut, athr than sian mimis must r
pik parasits from th larg fish, taks small its from thir tail fins. Th mor mimis tha m
blnny's mimiry of th wrss allows it to los in on its targt whn it might rathr than th rvr
<lthrwis atn itslf y th largr rf fish. But on might pt th an- darrs of modl anl
noyn of a rf 6sh at having a pi rmovd frorn its tail will pr<rvoklss as th modl fo s
tolran of th modl' th lanr wrasS' and impril th wrass'sailiry to pipvin swallowtail
gain th asy mal. To undrstarrdth prsistnof this mimiry systm' W Sond, if Batsia
must undrstand th riproal ffts of modl nd mimi on ah othr's gns that ontrol p
fitnss and th onditions undr whih thos ffts ar aland. strutions for th pa
Th most hllnging ompl mimiry systmsar th hundrds of ass th alphabtialntr
in whih dngous or toxi ninrals ar imitatd y harmlss or platal that thos gnsr i:
ons. In ah s th harmlss or palatal spisgains its advantag through the fat that tlrr is
th aility of othr animals to larn th assoiation twna visul or olfa- hritan of mimti
tory u with th dangous nimal and thn assoiat tht u with th gns of larg fft
hrmlss mimi arrd lav it alon. gns wr oviousl
Th most wll_studidass ar thos involving palatal uttrflisthat trastd with th Dar
rsmlunpalat:.rl, toi oIrs.Hnry Bats (1862) rought this phnom- was th umulativ il
non to th ttnti>n of iologists whn h dsridhow rtain Amazonian disput was larifid
buttrflis in th family Pirida or striking rsmlan to unrlatd ut- onrtd inhritan
trflis in th family lioniida. Bats had notid that th hlioniid ut- pattrns and th ff
trflisflw languidly yt wr rarly attakd y irds, whil th pirids flw or-rtrolling gns (s
m<lrfurtivly and lusivly,as if viding xposur to irds. Th hliorriids nw rsarh lrersrv
ar' in fat, toi nd unplatal, and th pirids ar nontoi and palat- t a l . 2 0 0 8 ) ,t h v o l
al. Bats dsrid this systm as a dramati dmonstration of tlr powr linkd ompl tlrat i
of natural seltion' pointing out th advantaggaind y an individual of a Th dsription of
p:rlatalspis whos featurs rsmld thos of a noxious spis that whn diffrnt popr.r
would avoidd y prdators. looking modls in di
Inspird y Bats's osrvations and suggstions' volutior-raryiologists ig pattrn within th
from harls Darwin onward plungd into studying th natur and volution is starting to rvltl
of this typ of minriry. With gratr srutiny am th appriation that mim- Third, th study o
iry is a ompliatd phnomnon; as a rsult, th study of mimiry am to unpalataility and t
oupy a ntral position in volutionary biology as a nrodl systm for undr- that a prdatr wor.
standing omplitd forms of natural sltionthat ould nrold ompl pat- must di so that th
tns from simpl gntiomponl1ts. sam apparan en
omplx mimti systms an lassifid into thr typs. In Batsian sprad via th lass
mimiry a harmlss or palatal spis rsmlsa dangrous or unpalat- gstd a kin-sltion
abl, toi on. It is not diffiult to undrstand how th prdator larns to mzrny toxi uttrfli
avoid a toxi uttrfly;a prdator tht ats on omsill nd vomits, aftr and otlrrs itd thr
whih tast avsion lads th prdator to avoid Drv that rsrnlth of. ity and warning olo
icr 7.7
Limiut a potntial disadvan- fnding food itm. It is also not diffiult t s Bats'Sargumnt that a mutnt
d).This happns whn modl of a pltal spistht rsmlsiln unplatzrlon will hav highr 6t-
r ologial rols and th su_ nss through a rdud risk of prdatin. If a ttr rsmlan<rnfrsa
rmpl, on ral rfs in th Pa- lowr risk of prdation, natural sltion will rfin tht rsmlanuntil
d food y piking rustaan mimi and modl ar indistinguishal'
'hih visit spifi spots on th Dspit th sirnpliity of th asi argumnts,Batsian rrrimiryri1isssv-
.spisof small fish, a lnny, eral important qustions for whih w hv only partial answrs.First, Bat-
:lanrwrass ut' rathr than sian mi-lismust rmain rr thn thir modls lst prdators nOuntr
ll its fm thir tail fins. Th mor nrimis thn modls and assliatth shd pattr wih palatailit1'
s in on its targt whn it might rathr than th rvrs.Th pris riationship ptd among tlr aun-
l. But on might pt th an- dansof modl and mimi is unlar, spiallywhn a singl spissrvs
:d from its tail will provok lss as th modl fo svral mimti spis in th sm lotion (.g., th
d impril th wass'saility to pipvin swallowtail of th southasrnUnitd Statsand its mimis)'
:nof this mimiry systm' w Sond, if Batsian mimis must rsml thir modls losly, thn th
'dl and nrimi on ah othe's gnsthat ontrol pattrn mllst inhritd togthr so that th gnti in-
fftsar aland. strutions for th pattrn ar not silttrddr-rringsuirl rprodr-rti<ln (s
;tmsar th hundrds of ass th alphabtialntry ..volutionof s'' in this volum).Rsarhhas shown
tatd y harmlss r paltal that thos gnsar inhritd as a lok, a pross failitatdin buttrflisy
:isgains its advantag through th ft that tl-rris no rossing-ovr in fmzrls.Th initial studis of in-
ation btwna visul or olfa- hritan of mimti pattrns suggstdthat thy Wr ontrolld y singl
n assoiatthat u with th gns of larg fft, a finding usd y som sintists to argu that suh
gns wr oviousl,vth important raw matril for volutin. Tis on.
llving palatal uttrflisthat trastd with th Darwinian viw that th important rw matrial for volution
s (1862)roughthis phnom- was th umulativ impat of many gnswith small individual ffts.This
; r i dh o w r r a i nA m z o n i n dispur was lrifid only wlrn dtild studis of inhritan rvald th
g sr-rrlan to unrlatd ut_ onrtd inhritan of th svral gns that ontrolld th asi mimi
notid that th hlioniid ut- pattrns and th ffts of othr gns that modifid th xprssion of ths
[ y ids,whil th pirids flw ontrolling gns (s Nijhout 200 for a tial disussion). Although
posurto irds. h hiiorriids nw rsarh has rvidth loation of ths gns in sotn spis (Clrk
pirids a nontoi and palat- t al. 2008)' th volutionary pathway that ror"rght thos gnsinto losly
ti dmonstrationof th powr linkd ompl tlrt is inhritd as a lok rmins to dsrid.
rggaind by an individul of a Th dsription of suh Irvolutionary pathway is vn nror ompliatd
:hosof a noxious spis that whn diffrntpopulations f th sam mimti spisinrittvry diffrnt-
looking modls in diffrnt gographi loations. This is an spiallystrik-
;stions'vlutionary iologists ing pattrn within th hlioniid uttrflis,and nw rsarhinto this Pattrn
udyirrgth natur and volution is starting to rvi thsp:rthways(Krnforst nd Gilrt 2008).
Jamth ppriationthat mim- Third, th study of Batsian mimiry ld quikly to th qustion of hw
t, th study of mimiry am to unpalataility and toiity ould volv, along with a spifi visual signal
)gy as modl systm for undr- tht pdator would rmrr.]f mr'ttnttlrt is unpalatal nd toi
ln that ould mold omplx pat- must di so that th prdator n larn to avoid othr individuals with th
sam apparan and initial tast, it is unlr how th mutant gn an
'd into thr typs. In Batsian sprad vi th lssi Darrvinian sltion amon]indivi.luals.Fishr sug-
lmlsa dangrous or unpalat_ gstd kin-sltinargllmnt asd on th supposdlygrgariousrraturof
rnd how th prdator larns to many toi uttrflis,ut rnt rsarh on phylogntis (Bltran et a|.2007
n omsill and vomits, aftr and othrs itd thin) has indiatd that grgariousnssvolvd aftr toi-
void pry that rsml th of- ity and wrning ()loration in rany of ths spis, nd so th kin-sltion
78 imir
xprts would agr on is ..th most rnt omlon anestor of a turtl, ir Th lpidosallrs ar
snak, a |izard, and roodil, and ll spisdrivd from that anstor.''If Thy inlud two grol
that dfinitiotris usd' the living rpdls inlud lizds, snaks, amplrisanians lizardlik spis rstr
(a group of limlss,lizdlik arrimals),rhynhoplralians(ttutars)'ro- th -7,000 spis of
odilians, bids, and turtls. It also inluds a grat varity of tint linags, tivly alld th squarn
inluding th marin ihthyosaurs and plsiosaurs, th flying ptrosaurs, and tropial rgions, altho
th vr-popr-rlar ornithislrian (ird-hippd)and s.rtlrishian (lizrd-hippd) masss pt polar ;
dinosaurs (stlr alphatial ntry ..)it-tosaus: Th n-rodlsystrnflr volu- uiquitous and prsist
tion'' in this volum). patd volutionry l
Phylogntially th living rptils ar oftn split into thr primary linags. plt limlssnsshas
Th lpidosaursinlud th lizards, snaks,amphisbnians,and ttlatarasand group. Th most fam
.With
r th numrilly dltninirntgroup of nonavin ptils.Arhosaurs inlud narly 3,000 livi
roodiliarrsand irds, s wll as a vast divrsity of xtint spis,inludig of limblss squamats
th traditional dinosaurs. Finally, th tstudins (sorntimsalso known s th th skinks. othr imp
hlonians)inlud th turtls nd tlrtoiss, small group of mlrphologially of omplx vn()l sys
uniqu vrtbrts.Amlng rptile iologists on of tlr ky rnt rsarh f_ omplity and pris
forts has fousd on lrtlw ths majo groups ar rlatd to ah othr, lrd ifid skull and tth all
wht thsrlationshipstll us out th history of divrsifiationof ttrapd dirtly inttl pry with
vrtrats. maxillary on; whr
Turtls ar oftn onsidrdth m<lstprimitiv grop of rptilson th . mourh.
sis of thir shard olr]itionof th lak of an opning (or aps) in th hk Finally, th living a
rgion of th skull (th anapsid lndition).ammals and thir rlativshav hr) and th 22 livin
singl opning (and ar lassifids synapsids)'and all othr rptils hav a
pir of opnings in th hk rgion (diapsids). Orr th sis of thir .
trrnl distintiv morphology and tlr notion tht th anapsid ondition is
th simpIstirnd thrforth most primitiv' turtls hav oftn n onsid-
rd th first linag to split volutionarily from th othr rptils.owvr,
rnt molular nd moplrologial rsrhindiatsthat turtls,whil r-
tainly iz arrd spializd' may wll form of modifid dipsid that
has sond:rrilylost th opnings hratristiof th group. atly whr
turtls fall within th diapsids rmains ontrovrsil;som analyss favor
losr affinity to th lpid<rsaurs, whil othrs favor .rn arhosaurin affinity.
Although th rlationship of turtls t() ()thrttrpods is ntrovrsial,r-
ognizing a tllrtl is not. Both th slrll (a fusd, modifid ri ag) and th
plamnt of the shouldr girdl insid th ri ag (as pposd to outsid
th ris in all othr ttrapods) ar uniqu faturs in th history of vrt_
rt volution. For th pst 220 million vars turtls lrv maintaind ths
uniqu fturs and hav n a onsistnt fatur of lif on arth. Th liv-
ing turtls now ontin aout 325 spisthat ar widly distriutd aross
th tmprat and tropial rgions of th world. Th living turtls ar oftn
dividd into twl major gror.rps. The plurodirs,or sid-nkdttlrtls(aout Th pin snak,itop
irritsth pin fors
75 spis)'hav a horizontlly rtrta|nk joint, whil tli ryptodirs
Nlv Jrsy.Pir-rsnak
(aout 250 spis)hav vrtially ori'ntdnk joints that nl thm to
witlr gophr tortois
rtrat thir had straight bak into th shll. Plurodirsdominat th south- rptils'with narly3'(
rn ontinnts (Afria, Sotlr Amria, nd Australiil), whil ryptodirs r ftursinluding'irr
found throughout ursiir and North Amria. injtingvnom intlpt
lder Reptils 741
ommol] anesto of a turtl, a Th lpidosaurs r th most divrs and nunrous of th living rptils'
' drivd frorn tht anstor.'' If Thy inltrd two !.roups: th morplrologially primitiv tuatras (a pair of
lizards,sltaks'mphisbanians lizardlik spis rstritdto svral lffshoreislands in Nw Zaland) and
ophalians(tlrtuataras),ro- th -7,000 spis of lizards, snaks, and thi rlativs (whih ar olle-
lratvarity of tint linags, tivly alld th squamatrptils).Squamatsr most divrs in th world's
aus' th flying ptrosaurs, and tropial rgions, although thy our on all ontinntal and most islnd land
nd saurishin(lizard-hippd) masss xpt polr and trm high-lvation rgions. On of th most
rs: Th modl systl for volu- uiquitous and psistntvolutionary thms in squamat rptils is th r-
patd volutionary loss of oth th digits and lims; y on stimat om-
splitinto thr;lrim,vlirrags. plt limlssnsslras volvd minimum of 62 indepndnt tims in th
lphisirnizrns,irnd tutras and group. Th most fmous xampl is th srraks' or srpnts (s figur)'
ian rptils.Arhosurs inlud !7ith narly ,000 living spis'thy ar y far th most sussfullinag
sity of xtint spis'inluding of limlss squamats)ut many othr ampls alound,partiularly anong
:s (somtimsirlso known as th th skinks. Othr important innovations in rh squamats ar th volr.ltion
small group of morphologially of ompl vnom systms,whih rh thir gratstlvl of morphologial
n of th ky nt rsarh f- omplxity and prision in th viprs and rattlsnaks.r th highly mod-
; ar rltd to 'ah othr, and ifid skull and ttlr :rllow injtion of prisl n-rodulatdamounts of vnt>rr-t
tlf ttrapod
lry of divrsifiatior-r dirtly into pr witlr a fang lik a hypodrmi ndlthat is attahdto tlr
maillary on; lvlrn not in us, t fngs an foldd ak into th
tiv gror.rpf rptilson th - mourh.
t opning (or aps) in th hk Finally, th living rhosaurs onsist of th ids (not onsidrd furthr
ammalsnd thir rlativshav hr) and t 22 living spis of alligators' roodils, and thir rltivs.
ds),and all otlrrptils hav a
;ids).on th asis of thir x-
n that th anpsid ondition is
, turtlshv oftn bn onsid-
om th othr rptils' Howvr,
indiatstht trrrtls,whil r-
r forrrrof m<ldifiddipsid tlrt
;ti of th grollp. xatly whr
ovrsial;slnranlyss favor a
s favor an hosaurianaffinity.
)r ttrapodsis ontrovrsial,r-
.sd,modifid ri ag) and th
ri ag (as lpposd to outsid
atursin th histtlry of vrt-
trs turtlshav rnintind ths
fattlrof lif on arth. Th liv-
rat ar 'idlvdistriutd aross
rld. Th livirrgturtls ar oftn
Th pin snak,Pitulphismllurzs, a rtrof th olurid famil,
rs,r sid-nkdturtls (aout
inhaitsth pin flrstsof th southrnUnitd Statsnd th Pin Barrnstlf
:rkjoint' whil th ryptodirs
Nw Jrsy.Pin snaksurrow in sandy soil nd ar o{tnfound in assoiltin
nk joints that nal thm to with gophrtortoisolonis.Snaksar th mst sussfullinagof limlss
Plurodirsdorlinat th south- rptils'with narly 3,000 spis,and hiit a markalrangof daptiv
Australia),whil rvptodirs ar fatursinluding',in rllany spis(ut nt pin srrks),
a omplx systmfo
. injtingvnom ito prv irnd ttkrs.
742 Mods of Spiation
Uniformly prsutdfor thir skins rrd rnat,y |970 aI|roodilian spis portn of hromosorr
on arth wr onsidrd thratnd or ndangrd y th Intrnational -99% of losly rlatd;
Uniln for th onsrvation of Natur (www.iulr'org), an intrntional tions, fusions' or othr r
organiztion that r-tronitorsiodivrsity dlins.Howvr, |995 onsr- a r r a n 8 m n t sa r i s i s h i g
vation fforts around th world had rdudthis y aout half, and as of this nwly arisn rarrangm
writirrg many spiSappar to b rovringto halthy population sizs.R- ish frtility whn thy :rr
flting thir rltivly ls phylgnti rltionship to irds, ro<rdilians ularitis in miosis. Thl
ar oth th most voal and provid th gratstlvls of parntal ar of any arrangmnts,thir F l h'
.!7hit's
group of nonavian rpti1s. Alas' viw of
On of th lssi qustions in rptilian volutionry biology is ..Why did in th nt l5 yasT . h.
th dinosaurs go tint?'' Although it is tru that mny linagsof dinosaurs onditions Wr nss.1
did go xtint roughly 6.5 millin yars ago at th nd of th rtaotlsp- ing spition)whil ls<
iod (and that tintion was tiggrd y a mtorit ollision with th ariss (th individul wh
:rrth),it is important to rmmr that phylogntially,irds atually fall erozygot). mpiril w
within th dinosaus and ar therfora kind of dinosaur. Thus in a vry ral somtims not as strit:
sns th answr to this qustion is that th dinosaurs did not go tint. frnt ways in whih ra
That pigon that yotr saw in th park today was a prftlygood dinosarrr;so rombination) rthr th
was th lrikn that you at for dinnr. His mphsis on th im1
planation for how tlryl
|BLIoGRAPHY
vr' thr r ption
's?ashington'
rnst' . H., and R. \(. Barbour. 1989. Trtlsof th World' D: a dirt fft of a transl(
SmithsnianInstitr-rtion Prss. .Whit
takld svr.rl
Grn.H. w. 1997. Sks:Th uolutirlof str i Ntur.Brkly:
gading spisdiffrn
Ur-rivrsitytlf al iforrriPrss.
L,. S. Y., T. W. Rdr,J. B. Slowiski,nd R. Lawsrrn.2004.Rsolvingrptil rnst Mar's influntiirlvi
rlationships: olularand morphologialmarkrs.In J. raraftand . J. ation, during whih a lar1
Donoghu,eds.,ssmbligth Tr of Life' 45|467. Nw York: Oxford ltiv prssursaftr olt
UnivrsityPrss. of sympatri spiationin
Pinka, . R., and L. J. Vitt. 2003, Lizrds:'V/idolllsto th uolutio of Diursit.
studid today in this ont
Brkly: Univrsit,v of liforniPrss. ..tho
Pough'F. ., R. . Andrws,J. . Cadl,. L. rump' A. . Savitsky,ad K. D. ours), he itiizd
Wells.2004.rpetolog'3rd d.Uppr SaddlRivr,NJ: ParsonPrnti disrimination and othr
all. -t{.8.s. ation. Instad, h suggs
in spiation. Although h
(2004) also not th ovr
Nlods of Specition (ichal J. D. Whit)
natiolr whrl th rltivi
A ommon misonption among studntsof sinis that thoris nrust hyrid prolms i s;.l
.\rhit's
orrt to produ progrss in thi disiplins.For ampl, studntsmisun- txt is rihl
drstand why thy larn th thory of th inhritan of aquird haratrs disussions of mpiria
.!7hit
of Jan-Baptist Lamrk, rmmring him only as ..tht guy who Was Wrong Was an outstndi
about hrdity''' Howvr' that is a shortsightdviw: Lmark prsntda onvind that th phn
omprhnsivthory of volution at a tim whr-rth prossswee poorly spiartion.Althoug so
undrstood. vn if his ontriutitln was impris in dtail, it positivly in- an dout that this t
flund our undrstandingof volution today. studis of spiation.So
With that in mind' it is a plasur to ommnt on ytologist ihal J. D. many ontinu to t
Whit's lassi 1978 book ods of Spitio.This book Was th most om- ..And' at th prs
plis:
prhrrsivtratmnt of spiation until th apparanof oyn and orr's o u t l r h o d so f s p i
2004 tt. h most famous asptof Whit's ook is his mphasison th im- mor justifilto gnr
ods rfSpiation 74.]
., 1970 all roodilian spis portan of homosoml rarrangmntsin spiation. shit notd tht
rdangrdy th Intrnational -99Y. of losly rlatd nir-nalsdiff by hromosomal invsions.trnsloa.
w . i u n ' o r g ) 'a n i n t r n a t i o n a l tions, fusions' or othr rarang-lnts,and that tlr rat at whih ths r-
ins.Howvr, I995 onsr- arrangntsaris is high y volutionary standards (1/500 individuals has a
this y bout half, and as of this nwly arisn rarrngmnt)'H also notd that most rarrnglntsdimin-
to halthypopulation sizs. R- ish frtility whn thy ar htrozygous zrushtrozygosity indus irg-
iationshipto birds, roodilians ularities in miosis. Thus whn divrging populations om to diffr in
'stlvlsof parntal are of any
arangn.lnts' thir F1 hyrids ar stril,and spitionhas tlurrd.
Alas' \/hit'sviw of rchil hromoslml spcitioz fll into disfavor
,olutionary iology is ...Why did in th nxt 15 yars. Thortial argumnts showd that trmly stringnt
tht many linagsof dinosaurs onditiorrsWr nssaryfor th frtility rdr.rtiontO strong (hnaus-
at th nd of th rtaous p_ ing spiatiln)whil also not liminating th nw arrangmntwhn it first
a mtoritollision with th ariss (th individual who first ars this mutation will nssarily ht-
'logntilly,irds atually fall rozygot). mpirial wok showd that rarrangtnnthtrozygotesr
l of dirrosaur.Thus in a vry ral somtirnsnot as stril as initially livd.Instad,studisnow suggstdif-
d i n o s u sd i d n o t g o x t i n . frntways in whrh rarre1ngmnts ontriut to spiation (.g.,fftson
.s7hit.
asa prftlygood dinosaur; so romination)rthr tharr tlr dirt frtiliry,dutiorrsdisLrssdy
His mphasis on th importn of rarrangmntsWas orrt' ut his -
plantiorrfor how thy ausd spiation is not widly aptdtoday. ow-
vr' thr r ptions: on lgrrtstudy of Shrom1,syast showd
D:
f th Wrld..!ashington' a dirt fft of transloation on hyrid fitnss(Dlnri t al. 2003).
Whit takld svral otl.lrontntious issus. H rviwd gnti data r-
Isterfu Ntr.Brkl:
garding spisdiffrnsarrd onludd tht tlrrWas no vidrrsupporting
R. Lawson.2004. Rsolvingrptil rnst ayr's influntil viw of a ..gntirvolution'' that ours during spi-
marks. In J. rraftand M. J. ation, during whih a larg fration of th gr-romhangs us of nw s-
,, 45|467. Nw Yrk: Oford ltiv prssursaftr oloniztion. tVhit also strongly favord intrprttions
of symp:rtrispiation in many inst groups, suh as Rhagoltis fruit flis' still
lotusto thuoltiorlf Diuersit.
studid today in this ontxt. ontrary to mally othr authors of his tim (and
runrp'A. H. Savitsky'ar-rdK. D. ours), h ritiizd ..thologiai spiatiorrisrs''whl strongiy dvot sual
JdlRivr,NJ: PlrsonPrnti disrimination and othr haviors as ing of paramount importan to spi-
-F1.8.S. ation. Instad,h suggstdthat hyrid prolms may mot ommon arly
in spiirtion.Although his viw is not widly aptd toda,v' oyn zrnd Orr
(2004) also not th ovrmphasisplad y som authors on sul disrimi-
' Whit)
nation whn th rlativ importan of ffts from suh disrimination vrsrrs
lf sinis th:rt thoris must hybrid proinrs i spirion has rrot rrdfinitiv[ydmrrstatd.
s .F o r x a m p | s. t u d n t sm i s u n - Whit's txt is a rihly dtaild rviw of th litratur on spiation.Th
rhritanof quird haratrs disussions of nrpirial studis of spition ar tlrorough and loqunt.
..thatguy who was wrong
only s !hit was an otstanding ytologist, ut lik mrny iologists, h am
;htdviw: Lamark prsntd a onvind that th phnomna h studid wr of paramount importan in
,whn th prossswr poorly spiation.Although som of his intrprtationsar unpopular today, nO on
rprisin dtil' it positivly in- an dout that this tt Was inf]untial irl motivting svral outstnding
ay. studisof spiation.Som of his ontrvrsial viws may yt vindiatd;
mnt on ytologist i\4ihal J. D. many ontinu to ativ aras of study. This quot from his tt still ap-
o.This book was th most om. piis: ..And, at th prsI1ttim' it wlld sm that swpirrggnraliztins
apparanof oyn and Orr's aout mthods of spiation should avoidd as fr as possil (it may
's
ook is his mphasison th im_ mor justifialto gnra|lzetn 2 -Yarstim)'' (Whit 1978, 24).
744 Mond to an
Th ltrntivtO trri
BIBLloGRAPY Was to find a wr,vttl s
framwork for suh a st
oyn, J. A.' nd . A. Orr. 2004. Spitioz. Sundrlnd,A: Sinaur Assoiats.
Dlnri, D., I. olson, S. Grammnoudi, I. N. Rorts, . J. Louis' and S. G. Olivr. lution in th 1930s and
200. nginring volutilnto study spiationin y:rsts.Ntr 422: 68*72, wr thmslvs prlgr
Ortiz-Brrintos' D.' J. Riland, J. H.y, and M. A. F. No<rr.2002. Romintion t w n p r o g r s s i o n i s nl
nd th lir,rgnof hyridizing spis.Gti 1 l6, ns. 2-3: 167_|78. frnrrvork fo nrrpro
Whit, Iv{.J. D. l978. lds of Spitit;.Sn Franjsl:W. H. Frtlra.
-.'.}J.
partly austh li
Th ook,s valu is r
o m i n s a s w p i n gh i
a o u t i n d i v i d u a ls i n t
Nlond to NI: The onpt of Progress also snsitizradrst,
in uolutionIr Biolog (Mihal Rus)
|BLloGRAPHY
volutinry idas ampopul:rrin N7strniviliz:ltionjust whn onprs
1996.ol
Rus' l\4iha|.
of lruman progrss wr flourishing. The Industrial Rvolution and th gowth Biollg.Cmbidg,,
of sin itslf ld duetd popl to pt limitlss progrss in human
knowldge and in human soity. Many ommntatos hav suggstd that
idologial lifs aout human progrss Wr a motivting fator in lifs .lvlorgan,Thomas I
aout volution nd spially in lifs aout th progrssivnssof volrt-
originlly traind as irn
tion. ilral Rus st out t irrvstigatwhthr prslrl lifs of sintists
th prinipal arhitts
aout progrssivnss in human soityand in volutilnwr orrlatd.
p r i o d 1 9 1 0 - 1 9 3 . 5H. r
ond to n is th rsr"rltof this study. Rus onluds that sintists'
ology or dii (193j
lifs aout th progrssivnssof volution wr indd orrlatd with
dmonstrating tht M
thir lifsaout soial progrss.This provids a hallngto th o1tivity
rod-shapd bodis founr
of th sintifilif in volutionary progrss.Judgmnts of progrssivnss
normous influn nlt'
annot dmonstratdon str:rightforwardmpirial grounds; thy ar nor.
in mryology and sp
mativ ssssmntsof what oul1ts as ..forward'' hirng. ould th mt.
his group rsolvd th qt
phorial irrfrnfrtlm soil progrss to v<tlutionryprogrss b takn
work of Ggor d
sriously by sintists?o nsrvr this qrrstirnRtls asks thr qustions
hypothtil onstuts1
aor'rt lrg numr of lristorial and ontmporaly volutionists.Fist, do
hrdity or rprsntd
a sintist'sidas aout iologial progrss orrspond to what on would
transmittd as disrtb
pt if th sintistwr inflr'rndy ids aout soial progrss?Sond,
p o r t n t d t s i n r i
ws that sintistonsious of th rlation twn soial progrss and io-
itd valu austh l
logial progrss?Third, to whilt xtnt did th iologial viws outstrip th
T. H. organ ws [-
vidn?h largr th gap twnvidenand thor, th mor likly it
r h n d o f t h U . S .( . i v
is that th sintistwas arrid i1rossthat gap y his or lrr soial iss.
h i s s s yo n h r i d i z . r t
Rus's answrs ar intriguing. First, almost ll nintnth-nturyvolu-
f a m i l y t h t i n l u d dh i s
tionists and most twntitl.l-ntuyvolutionists Wr progrssionistswith
gnral and ladr of }
rspt both to soity and to volution. But sond, thy sm to hav r-
Sott Ky (1779_184)
ognizd that this ratd a ias. Th ognitionis rvaldin diffrns-
th Stat Univrsity of
twn thir tratmnt of voltrtionary topis and rhir nonvolutionary
rrd Johns Hopkins U
rsarh.volution Was a t()pi ior puli ltursnd popular writings, br'rt
Willim Kith Brtl<lks(
it was usully not a topi fo lrrd<lrsintifi rs:rrh.Thomas nry
sis on th volutiony
Huly, Darwin's strongst sllppotr, was prft xampl of this tn-
C r u s t a n d A r h n i
dny. H wrot dozns of popular rtils on volution ut largly onfind
onludd that th gou
his sirrtifipuliations to PaPrs on omparativ morphology.
org 745
th two groups must hav divrgd from a ommon anstor in th distnt o.o YELLow' sPo.
o.' !iL I.
p a s t . A f t r a s m m r( 1 8 9 1 ) r r dt h r r y a ( 1 8 9 4 _ 1 8 9 5 )a t t l r N a p l s m a - ro wl]le oslN' IlRRY.
I.O BNOB}tIL
rin laoratory (Stazione Zoologi) working losly with th mryologist
Hans Dris'(1'867-1'941), Morgan gav up th largly dsriptivand sp-
i! ]FID.
onrndimportant iologial
ll' th mhanisnrof mryoni
1nti) rltionships.H ollab-
imnts on th dvlopmnt of [-'""*- .' PlN' l.{'.
marin invrtratsthat hv a
tmnt of ing al to ana|yze
)sssdirtly y manipulating
o g a nt h i s w a s r h d i t i o n i n
.{?
[""":":- LA.
1"":-',-:",
h founddth Division of Biol_
Lmmrsat th Marin Biologial [';;;;:
, whih h and othr iologists
his Bryn awr piod N4organ .l.,o vsIoIAL
l m r y o l o g y .u s i n g o r g a n i s m s
.8. oNY.soot.
It was only aftr ariving at o. Lo,! oBltlD.
B^R.
lrt lr m familiar with th '.o
.u*',. ..r UYD.
<for whih h and his laoratory
l0) am most famous, sta-
".
lf rdity,r,irsinitiatd and ar- LALe
: thoisof hrdity.
:hnism<lInturel sltionrv .eo.o sP.
(90|,in whih h argud tht "r.. oRULA'
throughoutth living world. B- homologous hromosoms duing rdution division in gamt formation),
;od Vris's (1848-1935) muta- th signifianof th hromosoms was still unlar. Finally, as an mryol-
aosin ()n stp by larg-sal ogist orgn hd a strong vrsion to both Mndlian and hromosom th-
frntspis from its parnts. oris aus thy smd lik rvrsions to th long-rjtd thory f
und primntalvidn in d prformation,whih had laimd that th pross of mryoni dvlopmnt
: (othr lmrki), was was mrly th unfolding of miniatur mryoni parts into adult haratrs.
his ontmporaisnot only - Rfrring ornplx and dynami iologial prosssak to invisil parti-
nian thory outlind arlir, ut ls in th grm lls, organ laimd, might sm at first glan to x-
tl approah to volution. D planatory ut in fat ultimatly plaind nothing.
; and spisof eotbr and Irlnially Morgan gan to hang his rind on all thr thoris twn
at will. Although d Vris's mu_ th spring of \9| and th wintr of 1'911. Th major imptus am first
was found to b th rsult of a from his own studisof Drosophila. organ had gun dingth fruit fly
o othr (and thus did not in his olumia laortory around 1908 to s if h ould produ muta-
lr tim' lik Morgan' as a vial tions of d Vrisian typ in animals. H had gottn nowhr whn on day h
osrvd among his ulturs of normally rd-yd flis on that had whit
)rgan Was also skptial of th ys.It was a mal, and h rd it to a rd-ydfmal;th offspring wr all
rsultsof hyridization nd th rd.yd.But whn h rd ths offspring with ah othr, h otaind a
h p h n o m n a s s d t m i n - ratio of roughly on whit-yd fly for vry thr rd-yd flis. This was
k numrous othr partiulat basially th ptd ndlian ratio for a foss twn two parnts ht-
ntury (thos of Darwin, rnst rozygous for a rssivtrait (in this as whit ws said to ressivto
to nam a fw), was ouhd in rd, whih was dominant to Whit). Mor important, howvr, rgan
; and was thrfor ompltly notd that ll th whit-ydf]iswr mals. In his first pulishd papr on
stnof ths supposd fators y-olor inhritn (1910), rgn assumd that th fator for whit y
to Mndlian thry Wr thr- must somhow sgrgatwith th fator for malnssin th formation of ga-
1' s Was a hrditry trait, th mts. But this planation Was not wholly satisfatory sin thr Was no
l : l s r t i of o u n d i n m o s t a n i - apparntiologial asis for assuming suh sltivsgrgationunlss,as h
that Mndlians tratd phno- soon am to s' th fators for dtrmining y olor rsidd on th as-
disrt,sharply diffrntiatd sory hromosoms (in partiular th ). Sin fmals had two hromo-
:,animal oat olor) rangd ovr soms' vn if on of thm had th nw mutant fator flr whit, it would
M n d l i a n ss m dt o i n v n t a s maskd y th rd fator on th homologous' or prtnr hromosom. F{ow-
,lain any givn hrditary ondi- vr,in mals, with only on hromosom' th mutant fator would not
l d n o t s u f f i r x p l a i n a p r r i - m a s k da n d t h u s w o u l d s h o w u p a s t h w h i t - y dp h n o t y p . B y 1 9 1 1 M o r .
d (asin assof pistasis'whr gan Ws onvind that Mndl's thory ould givn a onrt matrial
anothr). also qustiond th asis if it wr assumd that fators, or what !ilhlrn Johannsn had in
lparation of th two fators for 1909 lald8s' oclpid a spifiphysial rgion, or lous, of th hro.
at ah ndd up in a diffrnt mosom.
Subsquntly th ida of gns as arrngd linarly along th hromo-
until 1910 Morgan laimd that som ws givn furthr support y osrvations On what \/illiam Batson
ad anythingto do with hrdity' (\86|-1926) and thrs had idntifid as linkag, th inhritn of two or
W i l s o n ( 1 s 5 6 _ 1 9 8 )a n d f o m r mor traits togthr. Although Batson himslf did not intrprt this ph-
: th so-alldassoryhromo- nomnon in trms f th hromosom thory, othrs, inluding organ and
lr rsponsilfor dtrmination his studnts,did. Indd,Morgan usd th ida of linkag to dvis a way of
;hhromosomsWr larly vis- atually mapping a gn'sptlsitilnon th hromosom. on oasion, Mor-
dsriptiv work had n don gan notd, linkags appard tl rokn and th gns romind, as if
h pross of distriuting th th two homologous hromsoms on whih a group of gns wr llatd
: i o s i s ( t h p o s s f s p a r a t i n g had atually hangdparts:
7.'0 org
omos()m airrshin, J ' 199 1. Thomas Hunt rgn. haptr 8 of Trsformig Trditios
i mric Biologl, 1890_1L)1.'.Bltimor:Johns Hopkins Univrsity Prss.
:hOmosm lvlorgn, T. H. l90. ulltitl d dpttil. Nw Ylrk; N1milln.
191().S linritd inhritani Droslphil.Si 2: \2-122.
1916. ritiqu lf th Thulr o|-ullutiltt.Printot-t,NJ: intn
Univrsity Prss.
rgan, . H.' A. H. sturtvnt,. .|.N,lullr,nd . B. Bridgs. t9 | 5. T
hism of dli Hrdit. Nw York: Holt.
Shin' I., ar-rdS. !(lI. 1977. hlms Hut lrg, ilr lf Gtis,
[,ingt()n:Univrsity f Kntuk Pss. -C.,..
Blgianyt()logistl. A. Jarrssns
)9 papr whirt h a||cd cbis-
lologlusmmrs of a hromo- ullr, HrmannJosph(|s90-1967)
l to tlrirstrsqr"rnt raking
!hn H. J. ullr was aout l0 yars o[d, h wnt with his fath tO s n
H. Sturtvant ( 1 8 9 1 _ 7 9 7 1 )r a -
hiit on volution at th Amrian usum of Natural History. H r-
rlintilnould osvd irr
rnmbrd ing imprssd y th small siz of fossil horss nd how thir
rt of an,vtW()]nsln th hro-
siz and natomy hangd ov rnillions of yars. Although Nlull is st
t lastth rlativpositiorrsof
known as a gntiistand won his Nol Priz (in |946) for work proving
m h o l n o s r l m T. h i s m a p p i n g .
tht _rays indu gn mutations, lr onsidd himslf an volurionisin
laoratorygroup-inluding, in
his profssilnaland prsoIrallif. ullr was third-gnrtionAmrian and
( 1 8 8 9 _ 1 9 8 )a n d H . J . u l l r
attndd olumia Univrsity ln sholarship. Thr h tlok ourss with
:arhprogram tht was tndd
two ]atAmrin iologists' dmurrd Behr wilson and Thoms Hunt
s (orn, th lnous, arrd th rat) .silson
Morgan. was a ytologist whos writings hlpd onvin a youngr
g th validit of th hromos()m
gnrtion of iologists who t<lok his lsss that th ll was dply in-
d t h N o l P r i z i n 1 9 3 3 .
volved in hrdity, devlopmnt, an.lvolutio. <lrgan' skptial at fist,
;ntisnd tlr hromosom tlr. .Wilson's
vntually provd thris y using th tiny fruit fly' Drosophil
i-th dif{rntwn rd and
mlnogstr.In his snilyar Mullr mt two soplromoswith whom h
Ld wings-lnight atl,v th
foundd a iollgy lu. Alfrd Hnry Sturtvant and alvin Blakman
ln ould t. In 19|6, for a sris
Bridgs rrjoydsharirrgwith ullr thir idas On th nw 6ld lf gerrtis
iqe o| th Thor of ullution\,
that was rating suh a sti among thir tahrs.Ths fiv indlviduals wr
had osr,lin Droslphil were
th ky figurs in puting togthr th fild of lassial gntisand making
ir h hd argud thirt smll indi-
f u i r f l i su n i v r s l l l l yk n o w n m o n g s i n t i s t s .
rys tn{to swilmpd, tlr dis-
ullr rivd his PhD for working out som omplitis that aros
ouid arrmulatdindfinitl
whn orgn nd sturtvnt disovrd rossing ov. N{ullr lvok Id
Ltionof a nr'vspis.In his latr
him to a study of what h alld gn-haatrrlations. It took him aout
rmplifidtsvirvs,larly lrv-
l0 yirrs to pulish ths rsults austh pimnts Wr tdious :lnd
arwin's thlrv.It was for othrs,
tndd ovr mny .nrtions'For ullr it was ruial dmonstration
to us th nw glltisto flru-
of how Darwinian natural sltionworkd through th sltionof modifir
hat arnto knwn as th vo_
gnesthat ould ith intnsif1'or dinrinish hif gn'snrajor ffton n
ogan systm or trait' Mullr usd gns for wing shap that wr highly
vrial,and h isolatd and mappd th modifir gnsthat usd th vri-
prolmof ntural st.ltitl.
.|ottll ations. ullr, lik D:rrwin, rognizd that volution is a gradual proess
and tht nW organ systmsdo rrot ris sr.rddnlyut a uilt up gradually.
nd is Siltt,.Prill:tlr-r,
NJ: ullr workd firs at Ri Univsity in Tas nd thn t th Urrivsity
of Ts in Austin. Whil at ths institutions h wokd on th prolms of
ln and nlltural sltin visitd.In
mutation irnd tlr grr. stitd th trr muttio to hang in th
r 51,thesis: Prsptit'so th
l' N,IA:Hrval Univrsit1.Prss.
individual gn nd agud that th gn Was th asis f lif aus all
752 ullr
]produtsof gns,as shown y mans to a radition dos of aout 150 rontgns.Mullr sugg,std that vn.
h norml gns wr alterd y tually soity would hav to prti som fom of voluntry sltivrding
pol-ttan()usmutation rts at a to rdu this rrrr'rtational
load' or humanity would suffr disproportion.rt
finding nW mutation was lik amount of tim and ost to path up its infirmitis.
ullr lrd workd out th tools
BIBLIOGRAPHY
:ruit-flystoks in whih h ould
of thswr indud ln a nor- arlson,. A. 1983' Cs' Rditil,d Slit:h Lif d Work of H. .[.
uller.Ithaa,NY: ornllUnivrsitPss.
ng daughtould rd, and
r.rllr,H. J' 1968. Studisi Getis: SltdPprsof H'.|, ullr.
hromlsonrsirr hr immtur r- Blomingtorr: IrrdianUnivrsit1,
Prss.[Mullrsltdr.ts fronrmost of
:ls that killd hlf hr sons. Th his pprs,nd this lokontainsa lmplt iliogrph1'
lf his works.]
t hrmosomullr dsignd. -',.
l supprssorof rossing ovr, ;
ap lld Bar ys, B) allowd
'trol and -rayd dsndantsfor
rs. H flund ths in aundan
:gns).lros doss r 10 tirns
all hrnsposdto it. It is not
1y yars latr Mullr showd that
t gnmutations. Brokn hro.
.h or, if ths lls ar sprm' to
[lintokshowd th sam yl
L orn (mt-riz)nd alld this th
diatl'rlgnizdthat this yl
of radition siknss, spially
ds of atlmi-omvitims irr Hi_
; Nol Prize at a tim whn th
quikly am an advot for
and th military appliations of
BlBLIoGRAPY
lf natur'fronr a study of how soil
ow fmal buttrflis sarh for a Bartral,W', and L-.Harpr. |998. Th Trulslf WiIIim.lrtrI:Nturlist's
ditio.Athr.rs:Univrsityof GorgiaPrss.
s. Th prtitionrsof natural his-
. sintistslld thmslvsgol- olldolr,B., nd . O. \ilsn. |994. .|orto th Ants: Storof Sitifi
pllrtitl.
arridg'MA: BlkllpPrssf HrvdLJllivrsity Prss.
id th naturl lvorld wr alld Wilson,, o, 1994.Nturlist.Washingttlll' D: IslandPss. -].,
)rtisin irrasfom th <lr.rrs
of
\/ithhis foundatiln now givn, Shppard gos ()n to disuss a numr ()f faitlr nd authority,
importnt issus in no-ontologial volutionary studis.Partiulr attntion on th ross filr our sil
is paid to polymorphism-th wys in whih natural sltion an ilt th t h r a t h t t r l t t t l i .
sam tim hold diffrnt forms in a population. Muh of this naturally draws xists nssrilyfom
on Shppard's own work, partiularly th studis lf srrails thet h had don ural thellogiallaim.
arlir in th dad' with fllow Darwinian A. J. ain, in th Oford oun- In th hristin rlrg
trysid. Shppard had spnt a yar in th loratory of ThodlsiusDlzhan- llogy. Howvr, sp
sky in Amria, and it is prhps aus of this that h is not ntirly always had n imptlrt
dismissiv of gnti drift as a ausal fator in volution, :rlthoughth strong thology, and famouslr
imprssion givn is that it is natural sltionthat ally ounts in dtrmin- istn of God, s wt
ing th gnti omposition of populations. By th 1975 dition Shppard is stnd his natur. Prott
vn lss kn on drift, although h dos admit tht it might nlattr in prltin susiiln ut hv gn
voltion. Anglinhurh prti
Othr topis that gt major disussion a th voluti<lnof dlminan, thority of th tholis
prottiv olortion, and mimiry. Hr w suly s somthing of a traditionally givn natu
nationalisti ias, for ths wr all topis of major intrstto Ronald Fishr volutionary thory'
inis Geticl Theor r>fNturl Sltitl(190) and wr mrrtiondlit. in sevrl rspts.on
tl in th lading Works of Amrian volutionists. Dozhansk,s Getis mirals. ny natur
d the rigi of Spis(third dition, 1951 ) has vry littl on n-rimiry,and mirals-r ki ngs lf
vn that is gorr y th tim of th d fatl fourth dition, Getis of th of th deity. \/atouJ
ullutior Pross (1970). l,wontin, a arxist onvrt' has somwhat so. L{rwvr,ny sin
ontmptuously rmarkd that th nglish intrst in uttrflis,th hif supposs tht th worl<
sour of information aout mimiry, rfltsth dilttantish,uppr-lassna. thory supp()rtsth ui
tur of nglish volutionary illogy (sLwontin |974). With good rson' In th fa of this o
A. J. ain (of vry huml origins) rtd vhmntly against this. But for h a v n t a k n .o n t h
thos intrstdin th ultural fators that influnsin,if th mphasis twn th ordr of na
is put only on partilrlar prolms, Shppard'svry attrativlywrittn littl prod aording ttl I
ook is highly suggstiv.It is vry muh writtn y irn nglishman who volutionry thory do
lovs his ountrysid and th dnizns thrlf. t h y w i l | r .l l p o i n
BIBLIOGRAPHY argud that mir:rls
a matt of maning. J
Dzhansky'T. l951. Gctisd thc rigi of Spels.3rd d.Nw York:
physial ris frlm th d
olumiaUnivrsityPrss.
Fishr' R. A. 1930. h Gtilblrf Nturl Sltil.Oxford: ()ford still livd. vllutionry
UnivrsityPrss. Anothr pla whr
Kuhn, T. |962. Tb Struturof SitifiRt,olutitls.
higo:Univrsitof om into ontat is th
hiagoPrss. tn of God is th isr
Lwontin,R. . 1974,Th }titssisrlf ullutillrhg'Nw York:
and Aushwitz. No Il-
olumiUnivrsityPrss.
Rs,' 1958.Rviwof NturlSltiod Hcredit.rcdit'12:522_52. happn. Darwinism s
Shpprd,P. . 1958.NtttrlSeltiotllld rcdit.l-ondot.t: uthinson. posits a r-riquitousstru
-i4.R. wys throughout th lrv
o n r s p o n s m i g h t
t l f v i l , n a m I y ' h tC r l
Natural thlogy that God ould hav r
Gld didd rrot tl lt h
Thology dals with our knowldg of th natur and istnof God. R- that popl would nv
vald thology is th ara that trats of <lurundrstandinglf God thrtlugh thn fir without pain.
Ntrl Thcolog 7.57
d glsn to disuss numr of faith an{authoritv, this authority th hurh lr th Bil..That Jsus did
) n ys t u ( , | i l\).a r t i r r l ar t t n t i o n on th ross for otlr sins is a rvldthologial laim. Naturl thllogv is
lih ntural sltin atl at th th azlthat trats lf lur trndrstandingof Go{ througlr rason. hat God
ion.uh of this naturally draws xistsnssarilyfrlm his vry dfinititlrr(th lnologialargumnt) is a nat-
tudisof snails that h had ]on u r r I t h o l < l g i ll i m .
r A. J. in, in tlr oflrd oun- In tlr hristiarr rligion rraturalthology taks a ak sirtto rv:lldth-
)oratoryof ThodlsiusDozhan- ology. Howvr, spially givn th Grk influn on hristianit1,,it hs
of this tht h is not ntirly always lrad an importnt rol. atholis partiularly m:rk muh of r.ratural
in volrttion,although th strong theologv,nd fnlously Sint Tlrlms Aquins ] llv a sris of proofs of th
ln tht llyourrtsin dtrmin- xistnof God, as wll as disussiorlsof how w might rasonaly undr-
By th l975 ditiorr Shppard is stand his natur. Prltstantshav rgardd rason in ligion with mor
i t t h t i r m i g h t m i l t t ri n p r o t i n suspiion ut hav gnrlly allowd som pla for nturl tholog,v.Th
Anglirrnhurlr partiulrlv, sking middl r,vytwnth xtrm iru-
ar th volution f dominn, thority lf th atllolis and th trm ilialism of man1,Protstatrts'hs
w suly s somthing of a traditionrllygivn natural thlllgya signifiantpla.
lf major intrstto Ronld Fislrr voltrtionar1,thly,spi[IyDrrvitrisnl, irrrpingstln tru.tltlrolog1,
( | 9 )a n d w r m n t i o n d l i t . in svrlrspts.on ovious pint of tnsion ()urs()vrth xistnof
utionists.Dlzhansky,sGetics mirals. Many natural thologians hv rgtld that th isttl of
i 1 ) h a s v r , vl i t t l o r r m i m i r . n d mirls-rakings of th lrvs of natu-points t th ti.,.intvI-ltIon
to fouth ditiln,Gtis lf tbe of th lity.\Wtrould not hav rrurnd into win lrd God not don
arist onvrt' has somwhat so. Howvr, any sintifith<lry,in partiular n volutionary thtlry, p-
r intrstin uttflis,th hif suppossthat th rvlrld wllks aordirrgto lw, nd th sllss of sr'rha
ltsth dilttantislr,uppr.l:lssna- thor suports th uiquit,voI Iw irrrdhn tlr rrtlnxisrnof rrrirIs.
lwontin |974).With good rson, In th fa tlf tlris rjtion'th livr hs two optiolls' oth of whih
1 vhmntlyagainst this. But for hav -ntkn. on tlr l hand, it lrn argud that thr is a diftrn
influnsin,if th mplrasis twntlr lrdr of natur rrd th >rl of g. Nomlly th rvorld dos
rd's vry attrativlywrittrr littl prod aordirlg to law, lrut somtims God fls th nd to rtrvn.
t wittn y an nglishman who vlutionary thory dos not prov th:rt mirls i.rrimpssil, only that
of. thy will rar, point aptdy tlr livr' on th otlrr hnd, it an
ergud that rilsdo not nssrrily imply a akirrgof law. It is mor
mtt of maning. Jsus's rsurrtion' for instn, did not involv a
d.Nw Yok:
tlf Spis,3rd
physial ris fronl th dad tlt th flir.rg in th hrts of his disiplstl-rath
trl Slctl.oxitld: ()tord still livd. voir-rionary tl-r<lry
has norhing ro s:ryln this.
Anotlrr pla whr volutionary thory and natul thology sm to
'lutios. <lf
hiago:LJnivrsit1, om into ()ntatis th prolm of vil. A lssi r]umntagainst th is-
tntlf Gtld is th istnof physial and mol vils, srrhs arthquks
,lltillt.tr hgt'. Nw Ylrk:
and Atrshwitz. N ll-prvrful,all-good god luld hv llowd ths to
happn. Dawirrism smingly onfirs th istn of physial vil, for it
I rdit.Hrdit 12: 522_52.
r dit1,.[,tlndon
: posits triquitorrsstruggl fl xistl-l,invlvir-rgpain nd suffring al.
"..'.,'''.'.n:n,. o. ways tlrrlughor'rttlr livirrg r,vlrld.
on splt-ts right om fror an qually lassi ountr to th prolm
of vil, nrly, that God ann()tdo th impossil. Prhaps th only world
tlrat Gld ould hv rrd Ws on'tlrt inluds pain rrd sr-rffing'If
God didd ntlt tl lt urning aus pain, thn h would hv hd o llow
naturand xistnof Gol. R- that popl would nvr lan to far fir. It is ttr to hav pain with fir
ur undrstandingof God though than fir ,ithrlut pin. Prhps th t>r-rl1, way in whi God ould hav
7.'8 Ntrl hrlklg
ratd th living world was though a mhanism lik sltion,and hn It should mntio
h nnot lmd for its ad onsquns.(Kithr 2007 raks this t l Ii g n t D s i g l lT h t l r i s r
ritiism, that Darwinism xaratsth prolm of vil, and Rus 2001 lik natur of th living
rgus tht it may not quit suh a prllm for th hristian.) tht hn w must mai
Th majtlr pla whr volutionary thory, Drwinism in partiular, Although mmrs of th
oms into touh with natural thology is th qustilnof dsign (Rus 2003). idntifid with th h
From Plto on, it was rognizd that th living world is not random ut of thm. As might 1
shows omplity, an intgratd nd funtioning omplity, for xampl' sion of ..rationismlit.
th hand nd th y and th nos. How ould this ' givn that normally on should rrvr invlk
lind laws lad to randomnss? only thrlughth intrvntion of a dsignr, ar just not ndd in v
Orlsth rply. This rgumnt was pikd up y hristins, Saint Thomas is rquird. (S Dms
patiularly, and mad th asis of th most onvining of th argumntsfor th lading ontrovrsia
t h i s t n o f G o d : t h l i v i n g w o r l d i s a s i f d s i g n d ;t h i s i s a u s i t i s In onlusion, it must
dsignd,and th dsignris God. rethr tump taditin
Philosophrs' most partiularly David Hum, showd that in major r- just historil intrst.
spts' vn if thr is a dsignr,it anntlt muh lik th hristian god. th dsignlik natur of
Apart from anything ls, why stop at on dsignr?Howvr, Hum had to plaining. orovr, lik
admit that it dos sm s though thr is somtingat work. Using a vrsion hoil languag of d
of what is known as th argumnt to th st planati<ln'Hum agrdthat Ad1l ttil d Nturl
lind law dos not ld to dsignlik phnomna, and givn that th organi is produd as th funt
world is dsignlik,thr nrust slmethingor somon hind it all. With to srv that funtion, l
this pssiility, susquntnaturl thollgins,notaly Arhdaon William onsilus dsign. Th dt
Paley t th ginning of th nintnthntury' wnt on arguing for th - flr a rtin glll or fu
istnof God on th sis of th dsignlik natur of th living world. v o l v d w s f a s h i o n d y
Darwin's mhanism of natural sltion offrs an altrnativ to th god bautiful and historill
hypothsis.It is agrdthat th living world is dsignlik,ut it is argud that h y was aov all t}
this is ausof natural sltinrathr than Gld. No longr is th st - so1thingthat had ttl h
planation th hristian god. Drwin hs shown that th world is indd th y o u l d n o t h a v o m
produt of lind law, ut lind law of a spial kind, rramly,that whih pro- its, thos long-gon
dus daptiv funtioning by a pross of omptition and atrition. m i l l i o n y r sa g o a n d l a s
Not tlrat Darwin dos not hr disprtlv th istn of Gld. In fat, h flis (s figurs on pag
himslf was a livr in a dity whn h wrot th rigi of Species' tht it is ompl, with l
vn though latr in lif (mainly ausof th prolm of vil) h baman on ag 761).Why is th
agnosti. Rathr, Darwin shows that th dsignlik ntur of th world dos on th prolm rlizd
not nssitata god. In th words of today's popular sinwritr Rihard hristiaan Huygns had
Dawkins, ..Drwin mad it possil to an intlltually fulfilld athist'' if you Want to avoid h
(1986.6\. aus th diffrnt om
Dos this now man that th livrshould hav no thologial intrstin i n d i s ) .N a t u r a t t h t
th living world? any of today's hristins dny this strongly. vn though s o n a n d L v i - S t t i1 9 7 5
on an no longr hav a ntural thology-a proof of God-on an hav a ritis of univrsal D
thology of natur. On livs in God n faith (rvaldthology), ut and StphnJay Gould (
thn on's appriation of God's powr and majsty is {lshed<ruty tln's wronghadd sllution;
nountr with th living worid. Th uty of ros' th harm of a mr. Natural thology l
haffinh, and th urnof a fun shlw tht th ator was grat and prisly what should
loving dspit all th hatrd irnd vil in th world. a now-outmodd thlog
Nturl Tholog 759
rnism lik sltion,and hen It should mntiond tht thr is t<lda,v an ative group' th so-alldIn-
s . ( K i t h r2 0 0 7 m a k s t h i s tlligntDsign Thrlists,rvho argu thirt sltionannot plain th dsign-
' o l r ror f v i l , a n d R u s 2 0 0 1 lik natur of tlr living world, tlr:rtit shtlws ..irrdtri[omplxity''' nd
for thhristian.) that hn W 1Llstmak planator rfrnto dsigning intllign.
cry, Dawinism in partiula' Although mmrs of this group dny that this intllignmust nssarily
qustionof dsign (Rus 2003). idntifidwith th hristian god, in 1atthis is th dirtion takn y most
ving wold is not random ut of thm. As might lr ptd,th volrrtionaryommunity gardsthis vr-
'ning onrplity' for ampl, sion of ..rationismlit'' With som dis]ailr,arguing on gnrillgrounds that
rld this ' gil,r.rthat normlly on slrould tlvrirrvokmirals in sinnd on spifigounds that thy
r th intvntionof a dsignr' ar just not ndd in volutionar}.stlldies.Ntural sltionn do all tht
rp y hristians, Saint Thonrs is rquird. (S Dmski and Rus 2-4for a olltion, ditd y two of
:onvinir-rg of th argumnts for th lading ontrvrsialists,prsntingoth sids to th problm.)
f d s i g n dt:h i s i s e u s i t i s In onlusion, it must strssdthat, vn though natural sltion dos
rathr trump traditional natual thology, th onntiln is of mor than
t m , s h o r v dt h a t i n m j < l - j u s t h i s t o r i a li n s t D
. a r w i n i a n s a g r f u l l y w i t h n a t u r a l t h o l o g i a n st h t
l muh lik th hristin g<ld. th dsignlik nttlof th lir,ing wlrld is tb cruial thing tht nds _
sigrrr?Howvr, Hum hd to plaining. lrlvr, lik natural th<lllgians,Darrvinins us th mt:l-
r t h i n gl t ' o r k . U s i n g v r s i o n phorial Ianguagf dsign unrsrvdly. Gorg Willirrrs, in his sminal
planation,um agrd tlrat dpttio d Nturl Sletio,writs: ..!hnvrI livthat an fft
t n . n t Ig i v n t h t t h o r g n i is produd as th funtion lf an adptation prftd y rratural sltion
or somOnhind it all. \/ith to srv that funti<ln,I will us trms appropriat to humn atifi and
ls, nrltalyAhdaon Willinr ltlsiousdsign. T dsignatiorrlf son-rtl-ring s th mns or mhanism
lry' Wllt on arguing for th x- for a rtilin glal o furltiorl or pllros will implv thzrtrh mahirrr in-
ratur <lfth living world. vlvd ws fashiond y sltitlnflr h goal ttriutlt<lir'' (\966, 9).A
ffrs altrnativ to th god autiful and historially rdolnt mpl is furnishd r' th triloit y.
; dsignlik,ut it is argud that Th y was aov all th organ on whih th natural thllogianssizd as
n Gld. No longr is th st x- somthing that h]to hav a dsignr-Paly (1802) mad muh of it. Th
wn rhat th wlrld is indd th y ould not hv om lut y pur han' Darwinians agr.Th trilo_
l k i n d , n a r l y t, h a t w h i h p r o - ites, thos long-gon n-lrineinvrtrats(thy ppard mor than 500
)nlptirionnd attritiln. million yars ]oand lastd ovr 2'i0 rnillion yars) lr:rdompound ys lik
:h-ist of God. In ft, h flis (sfigurs on pag 760).|f on uts through a lns vrtilly, on finds
,rot th rigi lf Speies, that it is ompl' with two parts' spaatdy a urvd line (sth figur
rprolm of vil) h am an on pg 761),Wh is this? \What dsign prinipl dos it sv?Rsarhrs
ignlikenalturof th world dos on th problm ralizd that irr th svntnthntury, Rn6 Dsartsand
; populr sinwitr Rihd Christian Huygns lrd solvd th prolm. It is th kind of lns you uild
n intlltullyftrlfilIdathist'' if you want to voil hromati ar:rtion(n indistint fous oming -
aus th diffrrrtOmponnt olors lf whit light lrrvdiffnt rfrativ
d h a r ' n o t h o I o g i iln t r s i n indis).Natur t th sintistsy hundrds of millions of yars (lark-
dny this strongly.vn though s o n a n d L v i - S t t i\ 9 7 5 \ |
.Vfntworth
r poof tlf God-on an hav a ritis of univrsl Darwinism, lik D,Ary Thompson (1917)
:r faith (valdthology), ut and StphnJay Gould (1997) gnrallv do not Want to laim that this is a
majstvis f]shdout y ()n's r,vronghadd solutitln;ut th Warn that on swall<rwdos tlot mak a sum-
lty tlf :r r()s' th harnr <lf tnr. Natrtral tholog1,lds ou to pt r'riqrritous adaptation, :rnd this is
l tlrtth rator WaS grar nd prislywht slrould provn not assumdas part oi tlr unsnaggagof
orid. a now-outmodlthologialmtaphor.
760 Ntrl Tholog
l-_-:=:_\
i------j
-/:\-
lLIoGRAPHY
lrkson' . N. K.' and R.
Dsartsand Huvgn
Dwkins,R. 1986.h Blt
1995. Riur()u
Dn.rski,\/.A., and M. R
amridg:ambrid
Gould, s' I. 1997. Th Dar
n o . 1 0 : 4 - 7 .
Kithr, P, 2007. Liuillg tu
Nw York: oxford Ur
.!7.
P a l y , [ 1 8 0 2 ]1 8 1 9 .N a
Rivington.
Rus, . 2001. Dr
d R I i g 'i, ' a m h r
23.Drid
Triloit e (Rdops strbrgi, Dvlttitrtriltlit frlrl l}tlhmia). MA: Harvrd Univ
Nturl Theolog 76
Upr lns
unrt
Intralnsa
owl
i l i l l s l . t i r r l n i t l t l - tB o h m i ) . *l
lp:Rn Dsrts(left) and hristian Hr.rygns(right) shr.trrg
how to avoii
hromati arrtin. Blttotll:ross-stions.o1
triloItys. 1.3,,-,lrkson
n d L v i . S e t t |i 9 7 5 . \
lBLloGRAPl{Y
amridg Univrsity
Lmridg:
Nandrtals
',!.Printon,NJ: Printorr
-.R. Pr<lntlundNa'-az-dr-tahland rrlwsplld Nandrtill, th nm first ap-
pard as Nndrthal whn th fa-rlus flssilr,vasdisovrdin th summr
of 1856. Nandr is th Grk rndition of Numnn (nw man). Th Grk
)r) form was adoptd th family of svntnth.ntury Gmn hymn om-
posr,Joahim Nut-nann.Thal was Gman for ..vlly'' fo th splling
mportantook that addrsss ll
lrangof 1901, w,ht.t it am Tal. Nirrrdrtalthn was N',vnan'svally
;y. It is on of th fw xampls nar Diissllorf, now in tlr indr-rstrilrll-rartof tlr rvstrn art tlf Grmanl,.
ntion of iologists(most notly
and th sklton rvas found y marl quarry workrs rvlro w laring ot
th tautology prolm, units Of
a av alld th F-ldhofrgrotto. Tlr workmn thouglrt that th sklton
. in volutionarythory' th vo-
might hav longd to a prhistori tvar, so thy wr not vry arful
l issusonrning sintifi -
in thir tratmnt'nd th mor fragil nd smallr parts of th fa nd tth
n disussonly a fw highlights.
Wr not rovrl.Susquntlyothr prsntativsof th plpulation to
at So maks in this ook is
whih th original Nirndrtal individul blongd wr fllunl in Blgium,
lris simplampl w lnsiir
lWhn th toy is roetia, Fratt,Italy, and Spirr,s rr",ll as frthr irstit-tlq nd Isral.In
; whit ads. ft, anothr s;litnnhad n disor,rdelow th llorth fa of th Rok
small hols and ar thrby s_
. r b o t h s m a l l a n d g r n .B u t of Giraltar in 1848, ut it rmaind r.rnstudid in th olltiorrsof th Royal
ollg lf Surgorrsin London until th first dadof th twntith ntury.
rll; th fat that grn ads ar
Thos individuals hav n alld Nr-rdrtals vr sin.
tin o/ srrrallgrn ads, ut
Nandrtal skltl matrial ws flutr1 in th Blgin provin of Namur
a t t o m m u n t l f S p , v( p r o n o u n dS p )i r r 1 8 8 6 i n s t a t i f i da r h a o -
r prolmof th units of sl_
logial olltt with stln tools :rnd nimal ons. Tlr tools Wr rOg-
dpttk d Ntrl Seltio
nizd to of th sm kind s thos found at th sourlrrvstrnFrnh sit
rd group_lvladaptation in th
of L Moustir, lug in 186. In honlr lf tht typ sit, tlr tools assoiatd
: onsistsof fast individuals who
with th Nandrtals ar lways rfrrd to as Moustrin. Th us of ra-
rings happn.Th first is tht in-
diotiv lmnts to stalish asoIut dating of prhisroi matrial was
lnd is th:rtth hrd survivs and
,rd is mrly a y-podut of th not possil until irftr World Wr II, although it was a|izd that th
o t l s t r i a n w a s l l ] tr n t h U p p r P l o l i t h i t r a d i t i t l n sa s s l i t dw i t h
:re ls sletilof fast,hrds. This
m o d r n - a p p r i n gh u m a n s s u h 2 r s( i r l _ M a g n o n ,f o t t r - ridn 1 8 6 8 . M u h r -
;ltiottfor fast individu:rl dr.
lir than th Nrrdrtalswas th ml-rrrialfound y a Drh army physiiarr
: group sltionismhings rrpon
in lava in 1891 nd 1892 that h allrdPiththrlptts(w nw all it
groups' This disussion Suggsts
Homo) rtus.All tlf this matrial Ws mtiulously studid and put into n
mpirialqustion'whih is not
volutionary prsptivduring th first dad lf th twlltith ntury y
th Strassug irnlrtlt-l-tist Gustav Sl-rrvl(1844_1916). H did not dl
ll as th rst of Sor'swork, is
witlr tlr atltirl tlllrnisnrsrvhr,lr:rrrg \1.aspodtl1r-rtprsntd.r
al onptsin suh a way as to
.thr a sustantivand dis_ straightflrwalpitur of ontirruity and hang tlrroLrghtim. This w:rs
just aout th lst tim that th hurrrilnfossil rord was rrtdin an volu-
hiologial sins,ut tlros
tionaryprsptiv.
riologistsgo aout thir work.
In th sr.rmrrof ] 908 two rothrs who had rntly n ordaind as
prists wr purstting rhallogial :rvation nar th viilag of La
hapll-au-Silltsill orrz in sorttlrrvstrnFrn. Or-rAugrrst 3 th
narvsry'Sitt5:46_47.
,rTherlri Phiklsryhil unlvr] tltrrp[t end wll-prsrvdNndrt:rltrtlri:rl.l-rytllrrrnuni-
atd thir disovry to A6 rrri Bruii, a profsslr t th ollg l
, I t i , ,Pl t,i n lN
n J. : P r i n r o n Fran in Paris. Ab6 Bruil was alrady wll known in :rrhallogy,and h
-Z..
rommnddth:rtth othrs out thir sklton into th hnds of his frind
764 |']drtls
a r l l i n B o u l ( 1 8 6 | - 1 9 4 2 I , a p a l r o n t o l o g i s tw h o w a s p r o f s s o ra t t h
Mus6um d'Histoir Naturll in Pris. Boul's monograph dsriing La
hapll-u-Saintsal ()ut in thr instllmntsin 191 1, 7912, and 191
and st th ton for th study of human ..volution''for th rmaindr of th
':::!:r,
:
twntith ntury. Thr was nothing vn faintly volutionary aout it. Gus-
tav Shwal wr()t ln of th rviws f Bor"rl'smonograph, noting th
anatomial rrors Boul had mad in his ronstrution ut, uriously
nough, pting th patntly ntivllutilnaryast of th intrprtation.
Th vry nt yar World .JrI rok out' and on of th asualtisof that
onflit Was th volutioner orinttion in th Grman tratmntof th hu-
man fossil rord.
Bfor th wr lth th Grman and th nglish litrtur tratd Nan-
drtals as n arlir stag in human volution that had om transformd
through tim into nttlmially mrdrn flrm. Undr Boul's influn th
Nndrtls wr privd in ariatur and rgardd as hving n r-
p l d y a n i n v a s i o n o f m o d r n - a p p a r i n gh u m a n s w h o a m f r o m . . t h
ast.'' Thr is no arhaologial or anatomial support for this intrprta-
tion, :rlthough it is th viw now hld y most studnts of humn ..volu-
tion.'' Various stimatsof Nandrtal statur show mals ranging from 764
t o 1 7 5 n t i m t r s( 5 , 5 , 't o . 5 . 9 , ' ) a n d f m a l sr a n g i r r gf r < l m1 5 2 t o 1 5 8 n -
t i m t r s( 5 ' 0 ' ' t o 5 ' l ' ' ) . M a l o d y w i g h t h a s n s t i m a t dt o r u n f r o m 9 0
t o 1 0 0 k i l o g r a m s ( 1 9 8 t o 2 2 0 p o u n d s ) . r n i a l a p a i t yw a s l a r g rt h a n t h This drawing showsho
modrn avrag ut prftly in proportiorr to thir gratr ody wight. th modrn stty a r
larly th Nandrtals wr haratrizdy a dgrof skltalrOustnss arhitturwhil th
.With unhangd.Th Nnd
and musularity wll yond th mor rnt human avrag. intll.
lf approimatly70,0
tual and linguisti apailitis proaly at modrn lvls' th Nndrtls
Palolithi ml from tl
qulify as an arhai form of Homo strpis.
yars ago' nd rn
Th onvrsion of Nandrtal into modrn human form was aomplishd
y rdution in thos manifstatitlnsof roustnssas a rsult of thnolog-
ial innovations that allowd thm tl gin susistnwith lss pnditur |LIoGRAPHY
of ffort. Analysis of tra lmntsshows that thir intak of animal protin
Brir, . L. 1995. I]il
was approimatly th sam s that whih hartrizsth wolf. Th us of
in th mrgnof
traps and snars' howvr, gratly rdud th anrount of ffort ndd to 7t1-72t.
ring food hom. Most mutations rdu th trait that thy ontrll, and 1 9 9 5 .T h e S t g
whn sltion that maintains r<lustnssis rlad, its manifstationswill P r n t i H | i .
show a rdtltion. Th oking lf food will rdu th intnsity of sltion 2.ultio
Prss.
tht maintains tooth siz. During th last glaiation Nandrtals wr using
Morin, . 2004. Lat Pl
arth ovns to thaw food that froze, and a rdutiln in tloth siz an volutionary aproa
shlwn fom 130,000 yars ago in a straight lin of 7o evr2,000 yars un. lf ihign, Ann A
til th nd of th Plistlnartlund 10,000 yrs ago. Trir.rkaus,., nd P. Shi
Prliminary mtDNA lmparislnsshow that th Nandrtals diffr mor clkid. Nr,vYol
from th ndition in living humilns thn th avrg diffrn twn
plpulations of th living. Th distintion twn Nandrtals and living
asatoshi (b
humans, howvr, is not as grat as tlr mtDNA diffrnstwn plpu-
ltions of himpanzs Or vn th diffrnswithin populations of him- N4astoshi Ni has
panZs. sity sin 1990.H o
Nel 765
th volutionaryrlaionships
l t n n t t l n lk1l l t l w n e s s y s t m a t i s .
nialand nroluladt, rninly
s; ltroph(lrsisis thniqu
gl. ln l972 Ni dviopd a
)twn popultitlns from l-
standrdmthod in th stdyof
Ohta, Tomoko (b. 1933)
lr loslyrlatdspis,and th Tomoko oht has n a lading figur in dvloping a thortial founda.
vn as Ni's gntilistn.Th tion for undrstandingmolular volution. In hr work sh has omind a
rsn stimi1tof th numr of dp undrstanding of pattrns of molular variation within and among
and thr-rsth dgr of prltin- spiswith rtiv mathmatil modling to approh som of th most
l n s f r t ' t l tr rh i h h t w o p r r l t i n s fundamntal qustions about molular volution.
tndrstnd. Th mthod also hs A studnt of Knihi Kojima at North aolina Stat Univrsity, ohta r-
hlds.In l979 Ni dviopd turnd to Japan aftr omplting hr dotoral dg io t967 and gan a fa-
l sustitutionspr sit twn ..otoo
mous ollaoration with Motoo Kimura (s th alphtial ntry
lata.This was anothr milston Kimura'' in this volum). Thir arly ssaysdvlopd modls for th dynamis
pd svralllntlnth()dsfor of nutral allls (i..,allls that do not onfr diffrnsin fitnsson thir ar-
s pr sit t,ntwo DNA s- rirs).In |971 seand Kimura pulishd vo landmark ssaysthat showd how
r anJN. Saitou dvlopd th th nutral thory ould onnt mutation rats,lvls of protin polymorphism
volution;rry(phylogneti)trs. in natural popultions, and long-trm rats of amino aid sustitutions.Ths
hods rn phylogntironstru- ssaysprsntda omplt pitur of th nutral thory and dmonstrtd its
ly n phyllgntistudy uss this potntial powr as an planation for smingly disparat phnomna. This
t ladri th study of mrlular work providd a thortial foundation for svral lins of sarh'from using
s mthodshav hlpd larify th protin variation to stimat popultion paramtrs suh s th fftivpopu-
and gns.In additi<ln,h hs lation siz (a msur of how many individuals atually ontriut to th nxt
, of molularvlutitln,suh as gnration)to th us of molular variation in phylognti studis.
sd y Mtoo Kirnurir and asd In ploring th nutral thory furthr, Ohta was among th first to point
nathth for o{ ntur:rl sl- out that natural populations harord mor rar llls than nutral thory
s rvll as to our undrstanding of prditd. To aount for this pattrn' sh postulatd tht som molular
s. For his lntriutitlns, Ni was variants would slightly dltriousand othrs modratly d|trious.Hr
: si n 1 9 9 7 . hypothsis ld to diffrnt statistil distriutions of fitnss ffts that wr
alignd ttrwith osrvddata on polymorphism and rats of volution.
ohta also did pionring work on volution in multign familis lik th
lr F'ullut itt. ar bridg: hmogloins. Ths familis ar sts of tandmly rptd opis of gns and
ar major parts of th gnom. ohta rognizdth importan of ths fam-
lns.mri NturIist \06:. ilis in volution and dvlopd most f th arly thory that prditd how
gntidrift would promot divrgnin th DNA squnsof ths gns
. . lf o r l u d 1 i n gg' . n t iv r r i : l l l l il t t
among opis within a singl individual, among individuals in a population,
gs ,lf th Ntill dem of
and among populations. In 1984 Tomoko Ohta was ltd a forign mm-
lg mthod:A n, Inthd for r of th Amrian Aadmy of Arts nd Sins.In 2002 sh was ltda
\irltlgl d F't,llltirl4: 46-42"5. forign assoiat of th U.S. National Admy of Sinsand ws rog.
-w-H.L. nizd y th mpror of .|:rpanas a Prson of ultural rit.
767
768 On Crowth and Form
|BLloGRAPHY
Kimura, ., and T. oht. |971. TheortilAsptsof PopultioGtis.
Printon,NJ: PrintonUnivrsityPrss.
ohta, T. t98. uolutitld Vritilof ltltigmilis.Brlin:Springr-
Vrlag. -J.T.
trst' on thinks of Darwin's works and not muh ls.Th ok is and has
always bn sn as th distintly idiosynrati opus of a highly nti i.
.sntworth
ologist, D'Ary Thompson (profssor of zoology at St. Andrws
Univrsity in Sotland, 1917_1948),who is fftivlyrmmrdsolly on
th strngth of this ook. Why has it survivd?
Th ook prsnts th as for an aorrnt of iologial struturs asd
on mhanial and physial prinipls. It aspirsspiallyto mathmatial
mthodology; hmistry hardly fatursat all. Using an stonishing srisof Diodon
ampls aross th whole rang of phylogny and of salsof magnitud,it
shows how various typs of iologial ..form'' an potntilly plaind
Left: out|ine of th poru
y various typs of physial fors. Topis ovrd inlud saling, ratios o{ ( rt h glriscus mll).Nt
surfa to volum, growth urvs' hromosom pattrns' ll strutur and any knowldg of th ma
ll division (xplaind in trms of surfatnsion),multillula arrays (asin rpliathis findingsnd
honyoms), shlls, horns, flowr pattrns (ana|zd in trms of Fionai fom th fish form on th
sris),zra striping, and shaping of ggs and on strutur (plaindas 1948,1064.1
adaptations to imposd mhanial strsss).Th losing haptr is undout-
dly th st known and th stion most dirtly rlatd to volution. It
provd to b ..wrong'' (
dals with Thompson's ..transform'' mthod-a dpition of diffrns in
sis was saltationist.Th
morphology of rlatd spis (.g.,fish) or whol strutllrs'suh as skulls,
s o n ' s r l ' m i s s i o nw a s -
in th form of oordinatd gomtri dformations (familiar nowadays in th
stil1 prvalnt-to show
guis of omputrizd morphing) (s figur).Thr natur and vlu of this
..mthod'' is disussd l i n s o f t h a l r d ys i
lswhr(Hordr 2005); it is atually no mor than a
physis.
dsriptiv dvi and is not asd on' or an indiation of, any dvlopmn-
How is it possil th
tal or volutionary mhanisms as suh. Among prsnt.day apprahs to
ago? On fator might r
volution, th losst omparisons ar morphomtris and spia1lyallom.
with som instantly stri
try' whih, through J. S. Huly' was th most dirt and pliit dvlop-
up Thompson's mssag
mnt that aros out of Thompson's ook.
Proaly mor signifia
Th survival of this arhtypially lassi book is ll th mor puzzling -
ologists wr inspird
aus it is distintly datd in oth styl-it is writtn in sholasti and ven
Goldshmidt, Julian Hu
litrary mod-and ontnt. It dats from a tim whn gntis was in its
dan, and tr Mdaw
infany, Darwinian volution Was ativly ing qustiond in favr of La-
a symol and idal of l
markism, and thr wr many gaps in undrstandingof asi ll iology.
prolms of iology. T
or surprising still, Thompson adoptd positions on thsthmsthat hav
ious swp and rang
on Gowth and Fom 769
ts of Popultio]tttis.
e Filis. Brlin:Springr-
-J.'
worth Thompson)
,' is still in print as an abrvi-
n StphnJay Gould' This
l. It is hard to think of othr
a o n r p a r a l n d u r i n g s i n -
lrimarily for thir lristorial in-
nuh ls.Th ook is and has
opus of a highly Irtii-
ssor of zoolgy at St. Andrws
.ftivlyrmmrdsolly on
?
of iologial strr"rturs asd
:sspiallyto a mathmatil
. Using arr astonislringsis of Diodon
y and of salsof magnitud, it
t'' an potntially plind
Left: out'|tneof th porupin fish (Diodo) , Rigf: outlirrof sunfish
tvrdinlud saling, ratios of (rthgorissmol). Not tht Thompson was working intuitivly, without
m pattrns' ll strutur and an1'krrorvldgof tl-rmtlrmatisinvolvd. Today' with omputrs, w n
sion),multillulararrays (as in rpliathis findigsand hav a full undrstandingof the movsneddto go
ana|zedin trms of Fionai frorn th fislr form on th lft t th fish {orm on t right. (FrlmThompson
rd on strutur (eplaind as 1948,1064.)
h losinghptr is undout-
lirtly rlatd to volution. It
provd to ..wrong'' (sHordr 2005). is position rgarding phylogn-
- a d p i t i t l no f d i f r n si n
sis ws slttionist.Th ook is in vry wy a produt of its tim. Thomp-
vhol strutrs,suh as skulls,
son's ral mission Was-as a ountr to th vauous vitalisti xplanations
rtions(familiarnow:rdaysin th
still prvalnt-to show that iologial phrromnw plibl along th
). Th natur and valu of this lins f th alrady sintifiallyhighly dvloped disiplin and mthods of
l5);it is atually no mor than
physis.
indizrtionf, any dvlopmn-
ow is it possil that this ook is as influntial now as it was 90 years
ong prsnt-dayapproahs to
ago? On ftor n-rightwll th illustrations.Th ook is rihly illustratd
omtrisand spially-rllom-
witlr som instantly striking nd mmoral piturs tht in thmslvssum
ost dit and pliit dvlop.
up Thompson's mSsag;th an ' and hirv oftn n, radily rprodud.
Praly mor signifiant is th way in whih a stady sussionf ky i-
ook is all th mor puzzling -
ologists wr inspird y th work. Aross svral gnrations (Rihard
; writtn in sholsti and vn
Goldshrrridt,JLrlian Huxly, vlyn Huthinson, Alan Turing, J. B. S. Hal-
tim whn ]ntiswas in its
dan, and Ptr dwar to StphnGould and John Bonn) th ook was
ing qustiondin fvor of L-
a symol and idal of a rigorous sintifi approah to many of th major
:rstandingof asi ll ilogy.
prolms of iolgy. Th ook appald all th mor aus of its auda-
itions or-rths thnrs that hve
ious swp and rarrg. Tlris list inluds th nms of som of th most
on th origin of Spis
many of whoin mad Thomp- ntrrralsltitlnprovids plllatilnsin all tsras and how, onvrsly,
rgs.Th ook might appar to thsarasonfirm th tuth nd plwrof sltion.This is not so muh a ir-
liologists,ut thy hav rarly ular argulnnt s ()l1tht is rirrfoing.Th argumnt is known thnially
(morphologistsand spially as a osilill of idutios. |t Was pom()tdy th nglish philosophr of
Js (biomathratiians).In an sinWillianr lVhwll, and it was from this sour tht Drwin larnd of
:s' nd ppulin rhinking- its implrtan r]d powr.
ivrsitis-this ook stnds to- Thr wr m:rjor hngs in th sussivditions of te rigin. fter
y problms and planatory Darwirr pulishd, vry fw in tlr sintifi tlmtnunity ontinud to dny
tionwith whih it has n rh truth of vtllution. Htlll,vr, Darwin's thory of natural sltion was
.s7ntworth
r ft that D'Ary sujtdtO stringntsintifianalysis,and two objtions smdvry sri-
ms of Rltiue Grotb (192) ous. Th first w:ls that Darwin had no adquatthory of hrdity-wht w
today would all a thor1.tlf gntis-and without suh a thory lt Was not
lviottsthat a mhanisrnlik sltionluld lrav a lasting fft.Th s-
ond ojtion, asd orr physiists' lultitlns aout suh things as th
lridg': tslknapPrss of hatingpowr of th sun, ws tlrat th arth was far too yung for so lisurly
a mhnism as sltion tr hav takn fft.
nnss,d.' Lndmrk \Yritigs As w now know, tlr first ljtionwas wll takn, nd it was not until
|940, 82-832. Amstrdam:
Mndl's thory of hrdit'Was rdisovrdthat this ojtion ould
ountrd.Th sond ojtilnWas not wll takn, austh hating f-
London:Nlthun.
,g Biolgi,lDUk)pmt u)ith ft of radioativ day (and th tlnsquntmuh oldr ag of th arth)
, MA: rvrd Univrsity Prss. Was n()t thn kn<lwn.Hlwvr' Drwin flt oligd to spak t oth pro-
orm' 2nd d. amidg: lms, and to ountr thm h st:rrtdto rly mor and mor on supplmn-
tary mhanisms to sltion, most partiularly stl-alld Lamarkism, th
rsion. .,-o'jfl/.'.
amrid*.' inhritenof aqr.rirdharatrs.
Unfortunatly, th nd produt of tlr various hangswas no longr an l-
gant tnddssayut lngthy and umrsonltrat. It is for this rson
that today's radrsgnrally prfr th first dition of te origi.
arwin)
of Spisb es of Ntrl BIBLIOGRAPHY
:s i th Struggl for Life, was D r w i n , . l 8 5 9 ' o t h r i g i r l fS p c i s b 1 ' sN
t lf t u r l S l t i o , l r t h
l50 opis,and it was sold out Prcsrutiltof l:t,lurdI|si th Strgglfor l'if. Londorr:Jol-rn
sold all of th opis to ook- Mury.
nmdiatlytO prpar a sond 1'959..hrigi f Spis|,htlrlsI)rt'uitt: Vrioru ?r..
,hihth last appard in 1872. Pkham,d.Philallphie: Univrsitvof Prrrrslvnia Prss'
Rihrds,R. J., nd . Rus,ls.2008. h mbridgcrlm1lil t th ,.rigi
first part Darwin prsntd his rbilg: rrilg UnivrsityPrss. -^4.R.
of Sis.''
]asonsto think it plausibl. H
;ltion,pointing out th su-
wnt on to show (in an argu-
ntogen nd hIoge (StephnJay Gould)
althus) how a struggl for
populations to a lvl gratr |n toge d P,logya dtaild :rnalysisof th rlirtionshiptwn
l s u p p o r t . n d t h n h o w ' g i v n growth of th iniividual<lrganismand its anstrYis tightly intgratdwith a
ads to diffrntial survival and historial study of rpitr.rlationand othr pattrns of orrspondn-
ltion. twn stagsin dvlopmntand thos <lfvolutiorraryhistory.
..grathain lf -
lplid his mhanism aross th Th anal<lgytwn individul dvlopmnt and th
lt' iogographi distriutions, ing'' ws fmiliar ttl risttltl :rnd othr lassial authors. Gold shows how
logy, nd othrs-showing how this id w:rs rshpdir-rth ()nttof nintnth-nturyGrtnan idalism
772 ontogny nd Phylgny
Rprodutiv
oparin, Alksandr
Somti dvlopnrnt mturirtion Shift in attrn Alksandr l. oparin, a l
lrtio fluntiaI pionrs of th l
N o h r r g i n t i m i n g Rpitulcttirl
( lration) in th Sovit Union in 19
No hngin timing lrrttil Pdomorphosis ( 1 9 6 ) ' ) p a i n p r s n t
(y truntiol-r)= P gsis a r t h . O p r i n . s i d si n i
Rtrdtio No harrg in timing Pdllmorphrlsis mtal in stlishigrn
(y rtadatit>n)= N otlt- Ralizing th nrml
No hangin timing Rt'trdtil Rpitultil fild of iohmistry, o
(v polongation)
of th origin of lif fror
pri 77