Evolution: The First Four Billion Years

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.ё n a ls с i е n с еt,h i s d i d n o t o с с u r s i m LthЬy mеn who had no aims Ьut thе ly еvеry onе of thе nеw profеssional tе an еvolutiоnist bесausе hе was at'as morе than just a sсiеntifiс thеory.

somеkind of dirесtiсlnand mеаning. Bесausеall thеsе sеminal еvolutiоnists wеrе pеoplе whо grеw up irr thе еarlytwеntiеthсеntr.lry,whеn rnuсlr uldhavе Ьееn a surprisеif this Ьad not Ьееn tЬе lnаin motivating fасtсlr. But' likе Darwin saw that if thеy wеrе to havе profеshimsеlf,this nеrv Ьrееd of еvolr"rtionists sionalstatusfor thеir асtivitiеs-soп1еthingthеy аrdеntly dеsirеd as full-timе sсiеntists-thеnthеy would nееd ttl purify thеir work Of its ехtrasсiеntifiсaspесts.Thеy nееdеdto dеal witlr dсlсrrinеsof prсlgrеss:rnd with еХtгapolаtеd

34

Тhе Historу of Еuolutionаrу Thought

STUDIЕS oi\ IltliADIATЕD POPULAT]ONS D no So P Н I I, А M l] I' А N oG А S1,]tr]l*

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By BRUCE WALLAс]] Bio|фc,ot I,a|пralorу,CoIdSqriщ llа'rbor, N.Y. (With Fivе Teхt.figures) (Recеiod 28 Jdу |95б) rштвoоuстloш It iв well kпorтn thаt the тridеspгead use of ionizing гвdiations, beсsuse of tlrеiг gеnetiс effeсts, posoв в pгoЬlem futurе genегations. The*е гadiвtions induсe gene .""g""diog mutations. TЬe vast majority of mutвtions haie dеletегiousеffесtson individuals свгrying them. Undег the pгеssuгe of сontinued mutatiоn, thеse deleteгious mutationв will aосumulatе in populвtionв. Thereforе, an iтгadiated population will, on thе avеrage, be lцrmеd-havе its .fitnesв' reduced'-by a сontinual u*р.o." to irгadiation. The pгеsеnt aгtiсle вummarizев obsrгvдtions made on iгrаdiatвd po|oйi1"". oi lиo. so1lhilamе|юnчавtюr.Somе of thе.mвterial presentedteгe haв Ьееn p"ьri'ьa pгeviously (Wallaсе, 1950, lg5l; wl]tryе & King, l95i, 1952). This summвгy,^ho,n".'',u., will intro. duсe lrew mдtоrial in вddition to еxtйding the oгiginai obsеrvatiЬns. Мдтввlдr, AND DtEтЕoDs Thе erpет'imеntnlnop]nlion.s. Tire eхpетimental population в of D' mеIюn,чoslerarе kept in luсite and sсгееnсageв.The огiginal fliев *eгe oЪйinеd fгom an oгчoniR stгвin kept' lry mass tгвnsfеr for mвny yearв. Fouгtеen letЬal. and sеmi.letЬвl-fгee sесond сhгomosomes wеге extrвсted fгom thig.stгвin thгough the usе of a вeriевof matings identiсal to tJrовedesсribed later (Fig. 2.)'.Flies сaгrying thеsе sесond сhгomosomes and miхtures of orеgon.R and .markеd stoоk' сhromosomeвotheг than thе ..""od ;;;; йlarental fliеs оf thс populвtions. Rгief dевсriptions of thе рpulationв aге given in Tаblе 1. Thе left-hand сolumn gives thе identifying numbег foг eaсh populвtion. Thе весondсolumn indiсateв tircoгrgin of thо .Stoоkв'indiоatеs populations whoвеoгiginal flies сaтгiedluth'l.г."u рopulation. *uсond сhгomosomes of oregon-R. deгivation. Tlrтeе more rеоent populations aгe subpopulations оf populationв 5 вnd 6; the deвignation in the tablе gives tie^pвгent,r p"p"ьt,l"" and the gеnегation duгing whiеh eggs wеIе removеd to вtaгt tЬе nеw The tЬird р"p'i*;;;. сolumn indiсвtеs the numlrег.ofвdulЬ in thе population .Ь,gu' .uiu.* to popotu. ".g"*, tions of aliout ]0,000 indir'iduals,.small'fo рpulвtionsЪ"qo"oТly *]iti гu'u. tь"n 1000 ildividuals. Thе last thгee сolumns of TaБlе l givе thЁ typJ ot."*po*uге, the rsomе timе, one of thе most sеriоus rivals гo thе ,.primordial soup'' hеtеrotrophiс thеory was dеrivеd frоm thе idеаs of $7;iсhtеrshziusеr (1988).Aссorсling to his ..pionееrmеtaЬoliс thеory,'' lifе Ьеgan with thе appеaranсеоf аn aLltoсatalytiсtwo-dimеnsiotr:rlсhеmоlithorrophiс mеtaЬсlliс systеm that took plaсе on thе surfaсеof thе highly insoluЬlе minеral pvritе in thе r,iсiniry of hydrothеrmal vеnts.Rеpliсation followеd thе appеaranсеof this nonorgan. ismаl iron sulfidе*Ьasеdtwo-dimеnsistof thеsе Ьioсhеnriсаl сompounds in thе сarmеtеoritеsof 4..5Ьillion yе:lrsago (Еlrrеnfrеundеt al. 2001). So it Ьonaсеous bесomеspiausiЬlе, Ьut ntlt provеl1' that similar synthеsis took plaсе on thе primitivееarth.}-orall thе unсеrtaintiеssurrоunding thе еmеrgеnсеof lifе' it apреаrsto us th21tthе forrrrаtilrifеrous. rеptilеs in thе Pеrmiаn, dinosauгs in thе Triassiс,аnd maпrtnaisand Ьirds in thе Jurzrssiс.In thе 150 yеars sitrсе 1859,and dеspitеthе input of millions of hours of sеarсhing,palеontologists havеsimply adjustеdthе rrrid-Viсtorianpiсturе: thе origin of agnаthan fishеs has Ьееrrpushеd Ьасk from thе Silurian to thе СаmЬrian, thе origin of amphiЬiansfrom thе еarly СarЬonifеrous to thе latеst Dеvonian, thе origin of rеptilеsfronr tlrе еarlу Реrmian to thе rrrid-СarЬonifеrous, arrd thе origin of mammals from thе mid.Jurassiс to tlrе latе Triassiс. Thе origin of Ьirds .!7еrе hаs rеmаinеd unсhangеd in thе latеst Juгаssiс. thе fossil rесord hopе. r-rеwfinds should providе mаjor surprisеs from timе lеsslyr.rrrrеprеsеtltativе' to timе. Thе notiсln of a gotrd fossil rесoгd was сonfirmеd in quantitativе analysеs Ьy Мaхwеll and Bеnton (|990) сlf thе vеrtеЬratеrесord arrd Sеpkoski (199З) оf thе milrinе аnimal rесord. Thе first authors lookеd at thе aссumulation of knowlеdgеfrtlm ] .990 to 1987, thе sесond аt сhаngеs from 1982 to |992. In Ьothсasеsthе sum totals of divеrsity through timе inсrеasеd,еssеntiаllydouЬling in 100 yеars, Ьut thе inсrеаsеswеrе randomly distriЬutеd with rеspесt to til1-lе,and stl thе ovеrаll pаttеrns of divеrsifiсations and ехtinсtions rеmainеd unаffесtеd. In a furthеr study Bеnton and Storrs (1994) tеstеd rvhеthеrrrеw fossil disсovеriеstеnd to fill prеdiсtеd gaps or сrеatе nеW gaps' Thе,vfound thе formеr: in 25 yеaгs of study, nеw fossil finds haсl improvеd thе сomplеtеnеssof thе tеtrapod fossil rесord Ьy somе 5% Ьу plugging gаps morе oftегtthаn сrеating llеw saps. Тhеrе havе bееn two rnajor сhallеngеsto this somеwhat сomplасеnt viеw: sampiingеffесtsand mоlесular phylogеniеs.In both саsеs thе сritiсs aссеpt that tlrе fossil rес \ : :Е Е

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molisadvantagеoшs. Thе strеngthof sе'еs,and it agrееswith thе еstimatеd 19941. Vе lrаvе alrеady disсussеd sеlесtion [fесtsof transpositions on thе host. g ovеr bеtwееnеlеmеnts irr diffеrent h i s с а u s е sс h r < l m о s o n tiеn v е r s i o l r s , (B. Сharlеsworthеt al. 1994). Somе

rеarrangеmеnts havе fеw or no harmful еffесts and саn sprеad in populations in сlrrobесausеof gеnеtiс drifl, thеrеЬy сorrrriЬuting to еvolutionary сh:rr-rgе mosomеstruсturе. But many rеarrangеmеnts disrupt gеnе funсtions аnd аrе highlydеlеtеrious:nrany humаn gеnеtiсdisеаsеsаrе сausеdЬy сhromosomе rеаrrangеmеnts(gепсlmiс disеclsеs),inсluding mаny саnсегs (Dеiningеr еt аl. 2003).If this is аn importalrt disаdvurntаgе tо having high TЕ aЬundanсе, it prеdiсtsthаt TЕs sЬould aссltmulatе mainly in gеntrmiс геgions rvith low гесomЬinаrtionrаtеs' sinсе tlrеy arе tlrеп lеss likеly tсl саusе harnrful rеarrangеmеntsаnd will thеrеforе not Ье rеmovеd Ьy sеlесtion against thе produсts o{ есtopiсеxсhangеs'As mеntiсltlеdеarliег, this is oftеn truе' For ехanrplе, a largе pгоportion of tlrе Y сhromсlsоmеs of many spесiеs сonsists of ТЕ-dеrivеd sе_ quеnсеs(B. Сharlеsworth еt a|. |994|. Howеvеr, thеrе arе othеr possiЬlе еxplanatiоnsfor this pattеrn' srrсhas thе геduсеd еffiсiеnсy of sеlесtiоrrin rеgitlns of thе gеrromеwitlr rеduсеd rесoпrЬinаtion(sееlаtеr disсr"rssiorr). TЕs miry somеtimеsinduсе favoraЬlеmutations or сhromosomе rеarrangеmеnts'and sеlесtionwill thеrl sprеad thеm throughout thе pсlpulation.Son-rе ТЕ-dеrivedsеquеnсеsаppе21rto rеgulаtеgеnе асtivity (Dеiningегеt аl. 200З) or сonfеr othеr Ьеnеfitson thеir hosts (Brookfiеld 2003). As wе arguеd for satеllitеsеquеnсеs'howеvеr, a prеsеnt-dayЬеnеfitdoеs not impl,v that sеlесtion prсlmotеdtlrе initial sprеаd of tlrеsеsеqtlеnсеs.lr is possiЬlе thаt rеgulа_ tory or еr,еnсodin8 funсtions havе Ьееn aсquiгеd by suсh TЕs long aftеr tlrеy wеrеinsеrtеdinto thеir Prеsеnt loсations and fiхеd in thе population. MAssIVЕ тЕ AссUМULAт|oN AND PoPULAт|oN sIzЕ DIFFЕRЕNсЕs Although thе ratе of inсrеаsе of TЕs Ьy trаnsposition in spесiеs suсh as D. melаnogаsle/sееmslow (onе nеw еlеmеnt in thе haploid gеnomе еlеmеnt гoughlvеvеry 10 gеnегations)'сopl,numbеr lvill inсгеasеrapidl,vovеr еvolutionarytimеsсalеsif no oppсlsingfoгсе opеrаtеs:for еxаlmplе,with thе ratе of transpositiсlnеstimatеd еarliеr, a tеnfold inсrеasе woLlld takе only 2з,000 Drosopbilа gеnеrirtiotls,lеss tlran 2,000 yеars. Lаrgе diffеrеnсеsin TЕ аЬLrrrdanсеsamong rеlаtеd spесiеs,аs in thе maizе and riсе ехamplеs dеsсriЬеd еarliеr,aге thus rеadily ехpliсaЬlе if sеlесtion prеssuгеs against thеm аrе sоmеtimesrеlaхеd. Arrothеr ехamplе is proviсlеd Ьу Drosсlphilа mirаndа, itl whiсh art autos()nlеЬесаmе attaсhеd to thе Y сhromosomе аЬout 1 million yrars ago; this hаs sinсе Ьееn transmittеd only from fаthеr to son, сausing this gеnomе геgion to stop rесomЬining ЬесаusеrессrmЬirrаtiondoеs not oссur in Drr,,sophilа ma|еs.Тhе rеgion has sinсе thеn aссumulatеd a high dеnsityof TЕ insеrtionsinto nonсoding sеquеnсеs(Baсhtrog 2003)' Gеnomеs of somеspесiеs(е.g.,lrumаns arrd mаizе) lrаvеa vеry higЬ ovеrаIlTЕ aЬundarrсе (Тablе2), and at many сhromosomal sitеs, TЕs may bе prеsеllt at high fгеquеnсiеs(or еvеn in all individuals) (Ostеrtag and Kazazian 2001)' сontrasting lvirh thе low insегtiсlnfrеquеnсiеsin Drosсsphilа. \fhy arе thеrе suсh largе diffеrеnсеsЬеtwееn spесiеs?Pеrhaps thе пruсh largеr аnrounts of intеrgеniс DNA and largеr introns in humans than in

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1 7 0 Еuolutioп of tbе Geпomе Drosophilа п1еаnthat insеrtions hаvе lеss risk of сausing harmful mutations (although this would not Ье tгuе if intеrgеniсrеgions' sеquеnсеsoftеn сontrol genе aсtivity). A morе intеrеstingpossiЬility dеpеnds on thе wеaknеss of sеlесtion against еaсh nеW TЕ insеrtion. If TЕs rarеly ехсisе from a sitе, gеnеtiс drift сan somеtimеs lеad tо a sitе Ьесoming iгrеvеrsiЬlyfiхеd for an еlеmеnt. In spесiеs with low population sizеs, suсh as humans, sеlесtion against еlеmеnts is lеss еffесtivеin opposing gеnеtiс drift' and this possiЬly allows thе prеssrrrеfor еlеmеnt Ьuildup to prеdominatе (Lynсh 2007I. Nеw insеrtions into аlгеady_оссupiеdsitеs woшld flot Ье prеvеntеdЬy sеlесtion(thеrеwould Ьe no direсt hаrmful еftесts,just a slightly grеatеr risk of есtopiс еxсhangе), and so еlеmеnts сould snowЬall ovеr timе. This may aссount for thе fixеd еlеmеnts in mаnrmalian gеnomеs' Ьut rrot for thе vеry high TЕ aЬundanсе in maize, a spесiеswhosе high DNA sеquеnсеvaгlaЬility indiсatеs a largе population sizе (Tenaillon еt al. 2002I. RесomЬination in maizе may Ье largеly сonfinеd to gеnеs' so that TЕs in intегgеniс sеquеnсеs сould not Ье rеmovеd Ьy есtopiс ехсhangе. Thus еlеmеnts would Ьuild up Ьеtwееn gеnеs' as oЬ. sеrvеd (SanМiguеl еt a|. 1.996)'Нowеvеr, thе сausаtion сould Ье thе othеr way round: TЕs may simply rесomЬiпе lеss than l.lostsеquеnсеs. тHЕ ЕVoLUтIoNARY сoNsЕoUЕNсЕs oF GЕNЕтIс RЕсoMвINAт|oN AND PoPULAтloN s|zE SеlесtionagainstTЕs is only onе Way in whiсh small population sizе and laсk of rесomЬination сan Ье importаnt for gеnomе еvolution. RесomЬination rаtеs play important гolеs in numегous еvoluгionаryproсеssеs'еspесiallythе еvolutionary advantagеs and disadvantagеsof sехual rеproduсtion. Sеvеral diffеrеnсеs among gеnomе rеgions аrе сonnесtеd with rесomЬination ratе diffеrеnсеs' аnd thе еffесts of low rесonrЬination rаtеs arе similar to thosе of small еffесtivеpopulаtion sizе (Gоrdo arrd Chaгlеsworth 2001). Sеx and rесombinаtiorr аllow sеlесtion to aсt indеpеndеntly on diffеrеnt sitеs within thе gеnomе. This inсrеasеsthе еffiсiеnсy of sеlесtion.For ехample,if favorаblе mutaсions arisе indеpеndеntly, but vеry rаrеly' at two sitеs, thе populаtion will rarеly сtrntainany individualsthat сarry douЬlе mutants'еvеn if this is thе fittеst сomЬination (unlеssthе populаtion sizе is so hugе that thе sесond mutation еvеnt happеns Ьеforе sеlесtion fiхеs the first favoraЬlе mutation). Gеnеtiс rесombination allows thе douЬlе-mutant сomЬination to Ьe formеd. Whеn thеге is littlе or no eеnеtiс rесomЬination' or whеn rеproduсtion is asеxual, sеlесtioп on onе gеnе thus impеdеs sеlесtion aсting on othеr gеnеs' making fаvoraЬlе mutations lеss likеly to sprеad and dеlеtеrious mutаtions (suсh as amino aсid сhаngеs in prоtеins) moге likеly to Ье fiхеd Ьy gеnеtiс drift, paгtiсularly whеn population sizеs arе low (Gordo and Сharlеsworth 2001). This Proсеss proЬaЬly сontriЬutеs to gеnon-lеrеduсtion in symЬiotiс Ьaсtеria that livе within inseсt сеlls (rесomЬination bеtwеen Ьасtеria of thеsе spесiеs is prеvеntеd by isolаtion);thеsеrеduсеdgеnomеsalso havе fastеramino асid sеquеnсе еvolution' as prеdiсtеd givеrrthе lowеrеd aЬility of sеlесtionto еliminatе mutations with wеakly disаdv:rntagеous еffесts(Мoran 2003).

Gеnеs irr gеnonrеr.еgion similar еvolutionаry proсеs еls of adaptation. Тhis ma сomЬirring сhrotnosomal rt (pегh:rpsаlso in pаIt саLls strikingly illustrаrе thе ( D . С h а r l е s w o r t hе t a i . 2 0 С s o m е g е n е s ' Ь е t г а r ' i n gr h е paiг, Ьut tlrе Y laсks n)()st(

thе gеnеtiс informаtion onс nonrес()mЬiningсhromoso p е d е t h е i r а Ь i l i t yr o m a i n г fastеr than tЬеir Х-linkеd d i s а d v e n г а g е o u sr . e r i e r l t si gеnеs thаn likе normal gеn s е s o f s е х . l i n k е dg е n с si n ( F г i d o l i s s o nа n d Ё l l с g г е n. gеnеs addеd to thе Y сhro сonfirm гhetthis is duе tll i mutations (Bartolсlrnёand Similar Ьut mildеrеffесt gions with diffегеntrесoml сodons that еnd irr G or С сontеnt of third сoding pos oг thе аdjaсеnt r-ronсtrdi gапs, and А' thаlitlпа, tЬerс геflесts thе aсtion of sеlес fastеr аrrd/or morе aссurаt tеin sеquеirсе(AkasЬi еt al is' horvеvеr, vеr1' small сo protеin sеquеnсеs.so that prеfегrеd ovеr AТ. In an o сomPonеnt of tlrе gеllomr whеrе rесomЬination is in1 mal сodon usagе шight Ье would Ье lеss Ьiаsсdthаn o f t h е o Ь s е r r ' е r ]r е l l t i o n s сontеnt of сoding sеquеn and for thе wеakеning of tесtеd for thе gеnеs addеd and Сharlеswсlrth 2006). Horvеvеr, thе GС с()ntе С. еlеgапs also с o r1 9 9 9 ;М a r s h a l l e t a | . 2 О 0 2 ) . t сomеs from сomparisons Ьеtwееn n thе strеngth of fеmalе mating

prеfеrеnсеsfor сoгlspесifiсs.For ехirmplе, fеmalеs dеrivеd from populations of Drosсlphilа 1lseudotlbsсt,lrasvrrrpatriсwith D . persimilis сlisplay grеatег rеluсtanсеto nlatе with hеtеrospесifiсmаlеs than do fеmalеs 1rorn aIlopatгiс populations(Noсlr 1995)' F1 hyЬrid malеs arе stегilе,Ьut hyЬrid fеmalеs arе fеrtilе,allowing fсlr gеnе ехсhangе and sеlесtionagainst hyЬrids. onе of thе Ьеst-doсumеntеdехirmplеsof rеinforсеmеntoссurs in thе Ьеnthiс and limnеtiс есOtYpеsfound in lakе populations of thе thгееspinеstiсklе_ b a с k . H o w а r d R u n d l е a n d D o l p h S с h l u t е r ( 1 9 9 8 ) d е m o n s t r а t е dЬ o t h i n laЬoratoryсrossеs аnd in F1 lrr.bridsdеtесtеd in wild populations that hyЬrids suffеr a foraging disadvаntagеrеlativе to thеir parеntal spесiеs in thе parеntalеnvirсlnmеnts.In thе laЬoratory F1 hybrids rеarеd in a сommon еnviгonmеntwith ar-rrplе aссеssto food showеd growth ratеs similar to thosе of paгеntalspесiеs.Howеvеr, in fiеld studiеs F1 h,vЬгidsrеarеd in еitlrегtypе of parеntalеnvirсlnmеnt,littoral :rnd limnеtiс zonеs of lakеs' had lowеr growth ratеsand wеrе smаller at maturity than еithеr purе parеntal spесies.HyЬrids Wеrеpoorеr at сatсhing Ьеnthiс invеrtеЬratеsand plankton in thе diffеrеnt fiеld еnvironmепtsЬесausеof intегmеdiatеgill-rakеr numЬеr and Ьody sizе. In аdditiсln, Ьеrrtlriсfеmаlеs from svmpatriс poptrlаtions wеrе rrruсh lеss likеly to mаtе rvith intеrmеdi:rtе-sizеd F1 hyЬrid mаlеs than rvеrе Ьеnthiс fеmalеs from allopatriс populatiсlns whеrе limnеtiсs did not oссur. Thеsе rеsultssupport a modеl of rеinforсеmеntof sizе-assortativеmating prеfеrеnсеs through natural sеlесtiсlnagainst intеrп-rеdiatе-sizеd hyЬrids that havе lower fitnеssin еithеr pаrеntаl environmеnt. How irrrpoгtanthas rеinfkt.Us mitoсhondrial gеnе rеgr this study showеd thirt а г r i Ь u г е dt o m t r l r i p l еo r i g , 8). Two of thе thrее dip suggеstеd Ьy thе oссurrе not in thе iiving diplс t е t r а р l l l i d sa r o \ е t h r ( ) u diffеrеnt diplсlid аnсеsto аrсlsе from diffеrеnt dip sympаtry. Bесаusеthе dl latеd from onе antlthеr,, u с r s iс tl l t , r . P о l у p l t ' i di z . r t sisсеntсharrgеin thе аdv сеll sizе, whiсh hаs Ьееn сially prodцсеd аutotrip сhаngеs in trill ratе аs a r til.ltlitу Ьеtwееn lirrеаgеs rеinftlrсеd tri Il-reltеdiffеl grаy trее frogs illustratе aгisе аnd furthег shows mеnt саlls, сan Ье сlirесt

тHЕ RoLЕ oF GЕ

Irr thеory, gеnеriс drift outward from thе еdgе whеn a small nulrlЬеr tl] а nеw popr-rizrtiсlr-r. Irrpr: foгmatiсrn сrf rrеw spесt

Thе Pаttеrп аnd Procеss of Spесiаtion rdiеdrесomЬinationalspесiation in ts of сhronrosomаl rеarrangеmеnts R е с o m Ь i n а t i o n аslp е с i a r i o no с с u r s givе risе to a nеW linеagе that is еproduсtivеly isolatеd frоm Ьoth that сhromosomal rеarrangеmеnts jor еffесtson supprеssing rесomЬithе rеarrarrgеdsеgmеnts аnd gеnеs omеs.Gеnеs unlinkеd to thе сhгorlly thosеthat arе nеutral, frееly in. Ltal spесiеs. Rеarrangеmеnts that l еffесt on hyЬrid fitnеss сould aсt to еxtеnd thеir еffесts ovеr а largеr angеmеntsproЬaЬly do play an imspесiеsЬarriеrs'Ьut not nесеssariIy rid fitnеsstraditionally еrrvisionеd. s, whiсh rеsultsfrom hyЬridization thought to havе arisеn in suсh a i g h е rf i t n е s si n n o v е l е n v i r o n m е n t s ritors do poorly. Sцсh an аdaptivе on has Ьееnsuggеstеdfor а numЬеr 1 9 9 7 ;D o w l i n g a n d S е с o r t 9 9 7 ) '

lliсation of еntirе sеts of сhromo. hat сan pгoduсеinstantanеousspееly inсonrpariЬlеwith thеir diploid эd еithег through gеnomе dupliсa,loidy)oг thrоugh thе fusion of two rtеrspесifiсhyЬridization (аllopolytrесommon in fеrns, mossеs, algaе, .ts,partiсularly angiоspеrms (Stеbhеd is not fully undеrstood. Sеvеral s, might еnаblе a nеw polyploid to h е s еi n с l u d еs е l f - f е r t i l i z a t i o nv,е g е . t h е d i p l o i d p r o g е n i t o r s p е с i е s ,o r toг spесiеs.Indееd,many polyploid propаgation' and most diffег from istribution'Inсrеasеsin ploidy altеr )pmеnt'and many othеr physiolog'havе sеlесtivеadvantаgеs in nеw origin. n in animals oссurs in thе gray trее ) еastеrnUnitеd Statеs.Thе сomplех

сontains a diplкliсl spесiеs, Hуlа сhrу51lsсеlis(2N=24) and a tеtraplоid spесiеsНуlа uеrsicolor (4N=48). Both spесiеs oссupy laгgе rеgions of al. l o p a t r y( I J . с h r l s o s с e l i s i n t h е s o u t h е r n h a l fo f t h е r a n g е ,H . u e r s i с o l o r i r r t h е northеrn half), Ьut thеy also ехist irr Sympatry in multiplе loсalitiеs whеrе thеir rаngеs ovеrlap. Diploids and tеtraploids rеproduсе sехually and arе rеproduсtivеlyisoIatеdfrom сlnе anothеr through Ьoth prеmating (mаting signal diffеrеnсеs)аnd postmating (tгiploid hyЬrids arе stеrilе) Ьarriеrs to intеrЬrееding.Malеs produсе advегtisеrтеntсalls thаt diffеr in thеir trill ratеs,rvith thе сliploid H. cЬr\,sсlsсеlЬаlrvays produсing сillls with fastеr trill ratеsthan tlrоsе of the tеtraploid H. uеrsiссllсlr(Gеrh:rrdt2005). Studiеs Ьy Сarl Gеrhardt аnd сollеaguеshavе shown that fеmalеs of Ьoth diploids and tеtraploids arе highly sеlесtivе (еsресially in sympatriс populations) whеn еvaluatingсall сharaсtеristiсsof malе advеrtisеmеntсalls, strongly disсriminatingagainst саlls with thе trill rate of thе ..wrong'' spесiеs. A rесеntstudy Ь,vAliсia Hollowаy аnd сollеаguеs(Hollowау еt al. 2006) has dеmonstratес] аn unusuаl pattеrn of pftЬe Мoпtаstrаесt аппulаris ссlпiplех.Маriпе Biollэgу |27: 705-7|1. Lаndе,R. 19ti2. Rаpid origin gеnеtiсaIlv аnсiеntfеаturеsin еrrrlrryos, dеspitе Hаесkеl.(FrсlmНаесkеl 1866.) tlrеirаЬsеnсеirl аdults,thaг strопgl}.iп.rрrеssеd

amollg taха' al1d of thе lristory of spесifiс сharirсtеrs.Тhеsе thrее prасtiсal appliсationswеrе all first proposеd during thе ninеtееnth сеntury, Ьut for somеrvhatdiffеrеrrtrеasons thеy rеmairrimportant todаy. N()t mаnу rеadеrs of tlris Ьoсlk lvilI nееd to Ье pеrsuadеd that thе proсеss of еvolution takеs plaсе. Antiсipatirrg a rathеr diffеrеnt audiеnсе, Dаrwin mаrshalеdall thе еvidеnсе hе сould fiпd to support thе сonсеpt of trаnsmutationi]mong,spесiеswhеn lrе wrotе oll the Оrigiп of Spесiеs(1859). \Whеrе .,spесifiс'' Baеr,who rеjесtеdthе notidyplans. As thе first dеvеloprnеntalrеgulator,vgеnеswеrе сhаrraсtеrizесl at а rnolесular lеvеl, a majoг suгprisе еmеrgеd:homologous dеvеlopmеntalrеgulator.v gеnеs arе shаrеd among organisп-rswith vеry diffеrеnt Ьody plarrs.Thе first сlеar instanсе was thе геаlizаtion tЬat liп-12, Notсh, аnd ЕGЛl , impoгtаnt rеgulatory gеnеs indеpеndеntly idеntifiеd in nеmatodеs, arthropods, and с h o г d a t е s ,a r е h o r n o l o g o u s( G r е е n w а l d 1 9 8 5 ; Y o с h е m е t а l ' 1 9 8 8 ) .B u t t h i s finding was quiсkly ovеrshadowеd Ьy thе disсovеry thаt vеrtеЬrаtеsсontain Ёlox gеnеs homоlogous to thosе in l1iеs,and thаt thеy arе сlustеrеd in thе g е n o m еi n t h е s a m е r е l а t i v еo r d е r ( A k a r т 1 9 s 9 ) . I } yt h е m i d - 1 9 9 0 st h е l i s t o f rеgulatory gеnеs sharеd among thе mlljtlr modеl systсn1sof dеvеloprnеntal Ьiologv ran to sеvегaldozеn (Raff 1996; Gегhart аnd Kirsсlrnеr 1997). Sur. vеys of othеr phyla rеvеalеd a rеmarkirЬly Ьroad phylogеnеtiс distriЬution, irrсluding Ьasal nrеtаzс)аngroups (suсh as сnidаrians :rnd сtеnсlphсlrеs)arrd anatomiсally ехotiс onеs (suсh as есЬinodеrms ilnd rnollr:sks).It Ьесamе сommonplaсе to spеak of a shаrеd ,.toсllkit''of аrrimal dеvеlopmеnt (Сarroll еtal.2001). All of this was сomplеtеly unеxpесtеd.Animals in diffеrеnt phyla arе so anatomiсaily distinсt that idеntifying hопlologoLlssrruсtufеsand dеtеrmin. ing phylogеnеtiс rеlationshipsrеmain сontеntious (NiеIsеn 2oo2). This phеnotypiс distinсtnеss hаs foгmеd onе сlf thе сеrrtrаl tlrеmеs of сomp.rrativс аnatomy for as long аs thе fiеld hаs ехistеd (HalI l992: Raff 1996\, Thе notion оf a Ьody plan is that еaсh phylum of ar-rimalshas а uniquе ссlmЬination of аnаtomiсal trаits, most of whiсlr аrе аrсhitесгural(suсlr аs Ьoсiy s1,пrmе. try), еmЬryologiсаl (suсh as tlrе gеomеtry of сеll division), or Ьoth (suсh as thе nunrЬеr п tion of Ziniапthropus boisеi. rtаls with rеd hair' fаir сomplехions. есtion iп Rеlаtioп to Sех. Nеw York: ld A. S. Brooks. 2ООО.Епcусlopediа York: Garlаnd PuЬlishing. w York: !0. W. Norton. . Nаturе 425:.2324. S с i е п с сЗ 1 7 z | 8 7 3 - | 9 | 2 . 'olutiсlп.Nеw York: Aсadеmiс Prеss. ars of highеr primatе phylоgеny. In уsiсаlАпthrop olo gу' 19 З 0-1 9 8 0' сеnе hominid from Dmanisi, Еаst ].Swishеr III, R. Fеrring, A. Justus, ln, G. Bosinski,O. Joris, N4.-A. dе ishvili.2000. Еarliеst Plеistoсеnе :puЬliс of Gеorgia: Taхonomy, )19-1025. A. Мorin, С. Boеsсh, B. Fruth, 'dtttIf. 1999' Mitoсhondrial :iеs оf Afriсаn hominoids. сiепcеsUSA 96z 5077-5082. rtion оf divегgеnсеtimе for major |,oklgу апd Еuсllutioп2О:4244З4. t humans. Sсiепcе ЗI5z 1'94-196. :. 2008. Thе latе Plеistoсеnе dispеrsal п с еЗ 1 9 : 1 4 9 7 - 1 5 0 2 . ;еnyof thе primаtеs аs rеflесtеd in аssifiсаtioпаnd Hиmап Еuolutioп, nd. 2007. LimЬ-чizеpгoportions in theсusаfriсаnus' Journаl of Humаn of thе primatеs' Bullеtiп of the

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Bеyor Parad Biolo

Briаn G

Тhе quеstion that and funсtion in е stood. Form has t' gаnisms; and par individuаl organis сollесtivеs, suсh a! сharaсtеristiсs of l thеy havе аdaptil Howеvеr, thеrе is (i.е.,how thе forrл сеss whеrеЬy natt diffеrеnt forms in tion сannot ехpla plaining thе difft сontribution to th Thеrе is a tеndе сan Ье gеnеratеd indеfinitе divегsit assumption that I sis of two typеs (

spесiеs morpholo1 into distinсt, not ( сonstraints on tht somе forms саn Ь aminе in dеtail сс obsеrvational, ant а m p l е . A l l t h r е еa tion of сontinuou!

п Меdiсiпе Toпrasz. |999.TЬе dеvеlopmеnt :thiсillin-rеsist aлt Stаp hу loсoссus :diсiпе340:5\7-523' ; in influепza сorrtrtll. SсieпсеЗ12: 1005.Sггаtеgiс inrеritсгions in istanсe.Proсееdiпgsсlf thе З_з1.5 8. | .S a u n d е г2s 0. 0 5 'С o l l l p a r а г i v е :еsЬеtrvееlt thеpаtlrogеnrс 1 - 218 . Diseаsе.Nеw Yоrk: oхford :althаnddisеаsе: Work in progrеss. r' andD. H. Мortorr.2003.Typе I 7 pаtiеnts. AmеriсапJourпаlof

Bеyondthе Darwinian Paradigm:Undеrstanding Biologiсal Forms Briаn Goodtlliп

\ctаСhirurgiсаBеIgiса104: lеssons' highhopеs.FDА ;/fеaturеs/2000/5 00-gеnе.html. ls of Ьiogеography of l-ruman , Cenеtics36: S21_S27' ght'С. C. BаЬЬitt.J. S. Silvеrmаn' osпr:rn, М. IЬrahim,S. A. omar, еad,J. K' Pritсhard,G. A. !Иary, o no f h t r m a n l a с t a s pе е r s i s t е n сi n е

+0.

. Е . d е М е l k е r ,a n d М . Jеr irnpеrfесtvaссination: RоуаI Soсiеtу of Lolldoп, Series B d P . А . N 4 а t s o n 1. 9 8 6 . H u п t а r l эsis.ВiоSсiепcеЗ6: З68-373. D a s z a k ,a n d B . T . Е а t o n . 2 0 0 6 . иs Disеаsеs|2: |8З4-1840. t n d е n ri г o n m е n t a rl i s k f а с t o r s i n Мediсillе l0: 570_576. l n o t Г ) а г ritn i а n l r l е d i с i n е . э Ссlеuolиtioп of Pеoplе апd }07.origins of mаjor Ьuman iаsiс' translational,zrndсliniсal :soсiаtiсln 29 4: 1З 52-1З 5 8.

Thе quеstion that I shаll ехаminе in this еssay is thе rеlation Ьеtwееn form and funсtion in еvolution' and how thе оrigin of diffеrеnt forrтs is undеrstoоd. Form hаs пvo aspесts:slrаpеor struсturе' as in tlrе morphologv of or. ganisms; and pattеrn in tinlе tlr Ьеlrirviсlr,suсh аs thе movеmеlrts of individual orgarrisms(swimming' walking, flying) or thе spatial pattеrns of сollесtivеs,suсh as floсks of Ьirds or fогaging сoloniеs of soсial insесts.Thеsе сhагtrсtеristiсs of spесiеsаrе oftеn rеfегrеdto сollесtivеly as сhаraсtеrs' and thеy havе аdirptivе, funсtiоnаl signifiс:.rnсе for tlrе survivаl of tlrе spесiеs. Howеvеr, thеrе is a distinсtion Ьеtwееn undеrstandingthе origin fPalеy's Еuidепсes of Сhristiапitу lonaldFlеming (1961, 231) spесu..yеarning aftеr a Ьеttеr God than еrе nurturеd Ьy an inaЬility to rес:е with lifе's daily dеtails (Brookе l Wеrе'ln onе Way or anothеr' thorhought.

c сhallеngеsof еpiс proportions to 1izеdthе Мosaiс narrativе of spесial of using ЬiЬliсal gеnеalogiеsto datе zinе dеsign from naturе Ьy dеmonl thе ordinary, humdrum proсеssеs lmаns Wеrе' in somе fundamеntal l moral sеnsiЬilitynot in thе human Ьut in thе primitivе iпrpulsе of a rеighЬor to thе Frеnсh matеrialism ough Lamarсkian-еnthusеd mеdiсal Angliсan-Tory еstaЬlishmеnt. Not 'tlе Ьеtwееn sсiеnсе and rеligion rеth-сеntuгytrеatisеsthat сhartеd' as lhn Drapеr, tЬe Historу of tbе Сon.!Иhitе' ) or' in thе words of Andrеw

prеsidеntof Сornеll Univеrsity, A Historу of thе Wаrfаrе of Sсieпсe luith Тhеologу iп Сhristeпdom (\896). Thе vеry titlеs of works likе thеsе did muсh to сonfirm thе imprеssion of a monumеntal dеath strugglе Ьеtwееn thе forсеsof еnlightеnеd sсiеnсе and Ьеnightеd rеligion. But thе сrisp сlarity оf this rесеivеd imagе suЬstitutеs monoсhromе abstraсtion for historiсal tесhniсolor. Thеrе is a riсhеr narrativе to Ье unсov,,days', еrеd.For a start' at lеаst sinсе Аugustine, many Ьad read thе сrеation of thе Gеnеsis rесord symЬoliсally rathеr than litеrally. Various harmonizing stratеgiеsto maintain сonсord Ьеtwееn Gеnеsis and gеology had long Ьееn еlaЬoratеd,and thе idеa of a lеngthy еarth history was wеll еstaЬlishеd in thе minds of еarly Viсtorian Сhristian gеntlеmеn.gеologists. Natural thеology took a variеty оf forms' somе lеss susсеptiЬlе to Darwin's antitеlеologiсal сhallеngе' and of сoursе thеrе wеrе traditions, likе сеrtain strands of high Angliсanism and striсt unsеntimеntal Сalvinism, for whiсh thе signifiсanсе of thе dеsign argumеnt was nеgligiЬlе. And thеrе wеrе thosе likе John Hеrsсhеl and William !Иhеwеll whosе rеligious outlook prеdisposеd thеm to strеss сrеation Ьy natural law rathеr than Ьy divinе intеrvеntion' a movе that opеnеdthе door to thе idеa of еvolution as God's mеthod of сrеation. Thе ways in whiсh еvolution and religion сould Ье madе to fit onе anothеr havе takеn,and сontinuе to takе, myriad forms. lfhat furthеr сompliсatеs mattеrs is that whеn сonfliсt Ьеtwееn еvolutionists arld rеligious Ьеliеvеrs did oссur, thе сampaign was oftеn wаgеd from thе sidе of sесular zеalots as muсh as from thеologiсal rеaсtionariеs. Takе Thomas Неnry Нuхlеy, a kеy mеmЬеr of that Darwinian gingеr group, thе Х сluЬ,who dеdiсatеd thеmsеlvеs to driving out of powеr Palеyitеs and parsons alikе' No soonеr did hе hеar of thе Сatholiс St. Gеorgе Mivart's advoсaсy of еvolution than hе fеrrеtеd out a сopy of Franсisсo Suarеz's Sсholastiс thе..сomplеtе ology to insist that еvolution was in and irrесonсilaЬlе antagonism to that vigorous and сonsistеnt еnеmy of thе highеst intеllесtual, moral,and soсial lifе of mankind-thе Сatholiс Сhurсh'' (quotеd in Brookе 1991, 308). Thеn, in irritation at thе stratеgiеsof thosе whо rеad Gеnеsis ..provе that throughgеologiсal spесtaсlеs,hе dесlarеd his dеtеrmination to ..positivеly rapе,murdеr 6( arson'' wеrе еnjoinеd'' in thе old Tеstаmеnt (quotеdin Dеsmond and Moorе 1991, 472). Bеsidеs,hе was forеvеr talking ..hymns aЬoutthе ..sсiеntifiсpriеsthood,'' prеaсhing ..lay sеrmons,'' singing to сrеation''' dеnominating himsеlf a ..Bishop'' of thе nеw ессlеsiology, or..molесular ..thе dainingfriеnds to сhurсh sсiеntifiс,'' and ехpounding his tеlеologу.'' At thе samе timе advoсatеs of сonfliсt historiеs havе also had thеir own soсial agеndasin dеlivеring pugilistiс ассounts of sсiеnсе and rеligion. In thе сasеof Drapеr' who rеmarkеd that sсiеnсе was thе twin sistеr of thе Rеformation' thе real oЬjесt of opproЬrium Was thе Romаn Сatholiс Сhurсh; thе doсtrinеof papal infalliЬility еnunсiated in 1 8 70 appallеd him and lеd him to prоphеsythat it would Ьring thе Сatholiс Сhurсh into сonfliсt with hithеrto.W.hitе, friеndly govеrnmеnts. As for thе opposition that his nonsесtarian

. 1 - \I

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Еuolutioп апd Rеligiсlп Сornеll Univеrsity had attraсtеd from сlеrgymеn strеngthеnеd his геsolvеto kееp sсiеnсе at thе forefront of thе сurriсulum; Сornеll would Ье an asylum foг sсiеnсе liЬеratеd from thе striсturеs of rеligiоus dogma. It was manеuvеrs of this kind that promptеd somе historians to rесonсеptualizеthе so-сallеd warfarе Ьеtwееn sсiеnсе and rеligiоn in Viсtorian soсiеty as a soсial strugglе Ьеtwееn two сontеnding intеllесtual еlitеs-namеly, thе old-fashionеd parson and thе nеwly professionalizеdsсiеntist (Turnеr 1978|.In this sсеnario dеbatеs aЬout sсiеntifiс knowiеdgе in gеnеral and еvolution in partiсular sim. ply Ьесamе a furthеr arеna in whiсh tusslеs for сultural suprеmaсy wеre played out. Howеvеr alluring grand narrativеs may Ье, storiеs of thе еnсountеr Ьеtwееn еvolution and rеligion that tradе in systеmiс сonfliсt or, for that mattеr' сoopеration sасrifiсе historiсal intriсaсy tо linеar simpliсity. In faсt, rеligious Ьеliеversrеspоndеd in vastly diffеrеnt Ways to еvоlution, and thеir stanсеs do not follow any straightforward taxonomy of thеologiсal oriеnta. tion' dеnominational affiliation, or doсtrinаl prеoссupation. Bеsidеs,еvolu. tionary thеory Was nеvеr еnсountеrеd in a сultural vaсuum' and dеЬatеs aЬout it routinеly Ьесamе a mеans of giving voiсе to a rangе of othеr anхiеtiеs. Somеthing of thе divеrsity of thеsе еngagеmеnts mаy Ье glеanеd from thе following thumЬnail skеtсh.

Thе Viеw from Romе Сatholiс opinion on еvolution has long Ьееn dividеd. From thе еarly days thе opposition of a group of Jеsuit thinkегs Ьrought togеthеr Ьy Popе Pius IХ to сomЬat thе forсеs of modеrnity Was ехprеssеd through thе pagеs of La Сiuiltd Саttolicа' an outlеt that ехеrtеd vеry сonsidеraЬlе influеnсе among ItalianJеsuits. Thus in thе еarly 1860s thе nесеssityof maintaining thе fiхity of spесiеsagainst Lamarсkian idеas of transformism was vigorously promulgatеd by G. B. Piаnсiani in its pagеs (Brundеll2001). By сontrast' thе еfforts of thе Еnglish сomparativе anatomist St. Gеorge Jaсkson Мivart' a Сatholiс сonvеrt' to Сhгistiаnizе еvolution through a Lamarсkian rеading of transmu. tation in his 1871 volumе Оп the Geпеsis of Species was praisеd in thе Catholiс prеss. Huxlеy might snipе that Mivart сould not Ье a sturdy sоldiеr of sсiеnсеand a loyal son of Romе, Ьut Popе Pius IХ аwardеd him thе dеgrее of doсtor of philosophу ln 1876, and the Bеlgian Ьishops prеssеdhim to takе up a сhair аt thе Univеrsity of Louvain in 1884, although hе latеr fеll into disfavor on aссount of сеrtain thеologiсal writings. Howеvеr voсifеrous thе opponеnts of еvolution undoubtеdly wеrе, thе offiсial сustodians of Catholiс dogma did not takе any aсtion against Catholiс еvolutionists until thе 1890s whеn attitudеs Ьеgan to hardеn as thе aging Popе Lеo ХIII, hithеrto a forсе for modеration, lost ground to thе inсreasing influеnсе of Roman Jеsuits and thе maсhinations of thе СiuiltЙ Саttoliса сotеriе. So whilе Мivart's еfforts еsсapеd at lеast for a timе thе сеnsurе of thе Vatiсan' othеr Сatholiс еvolutionists did not farе so favoraЬly' notaЬly thе

Frеnсh Dominiсan priе o r g а п i q И е s( 1 8 8 7 ) ,a n d rolе in thе dеvеlopmеnt |ished Еuolution аnd Dt Both thеsе writеrs su1 Ьеit within сеrtаinlimi foundations, was dirесt thеologiсal еnеmiеs of t] might Ье thе produсt Saсrеd Сongrеgation ol . . е v o l u t i o nt h е o r yi tЬаt human Ьody'' (quotеd Ьook was сеnsцrеdwas spееdily rеtraсtеd in tht find ways of rеvisiting t of thе Ьook, Ьut it too : It was muсh thе sam rесognition for his Sс Саtb oliс Sсieпtists (789

sсепt of Мш1 as thе сaU аnd Dogmа was to dеп ..to admirе' muсh in it xx). Likеwisе groundе tiс ехеgеsisto justify a intеrnational suссеss,a of a fuming Сiuiltй Саt lеnt with rесklеss assс sumptions. Thе Сongr vеry largеly on thе ma lutionism was part of z tion of tехtuаl сritiсi ессlеsiastiсal affairs. L voraЬlе rеviеw hе rесе pains, prеSеntlyсamе l Various faсtors play i51ц''-1hg tеndеnсy or indеpеndеnсе and to : suсh forсе. This indrс сеntury Сatholiсs wеr with whiсh immigran сеrns rotatеd аround t arly trеnds, notably t Ьiology wеrе thus ofl to еstablishеd ways o authority and who hа Thosе who found еvс

Еt,ollttioп апd Rеligioп gymеn strеngthеnеd his геsolvе to lum; Сornеll would Ье an asylum .еligiousdogma. It was manеuvеrs Lsto rесonсеptualizеthе so-сallеd сtoriаn soсiеtyas a soсial strugglе _namеly, thе old-fashionеd parst (Тurnеr 1978). In this sсеnario al аnd еvolution in pzrrtiсularsimslеs for сultural suprепraсy wеrе ' Ье, storiеs of thе еnсountеr Ьеs,vstеmiссonfliсt or, for that matilсy to linеar simpliсity. In faсt, ]геntWаys to еvolution' and thеir taхonomy of thеologiсаl oriеntatаl prеoссupation.Bеsidеs' еvolua сultural vaсuum' and dеЬatеs rg voiсе to a rangе of othеr anxingagеmеntsmay Ье glеarrеd from

n dividеd.From thе еarly days thе lught togеthеr Ьy Popе Pius IХ to 'rеssеd through thе pagеs of Lа :ry сonsidеraЬlеinfluеnсе among nесеssityof maintaining thе fiхity sformismwas vigorously promultеll 2001). By сontrast' thе еfforts еorgеJaсkson Mivart, a Сatholiс r Lamarсkian rеading of transmu_ is of Spесiеs was praisеd in thе vaгt сould not Ье a stuгdy soldiеr е Pius IХ awardеd him thе dеgrее :lgiarrЬishopsprеssеd him to takе 1884, although 1rеlаtсr fеll into rritings. llution undouЬtеdly wеrе' thе oftakе any aсtion against Сatholiс еs Ьegan to hardеn as thе аging i o n . I o s tg r o u n d t o t h е i n с г е а s i n g rtiorrsof tЬe Сiuiltа Саttolicct сoеastfor a timе thе сеnsuге of thе ot farе so favoraЬly, notaЬly thе

Frеnсh Dominiсan priеst M.-D. Lеroy, author of L'ёuolutioп des espёсеs orgапiques(1887), and thе Amеriсan Jоhn Zahm, who playеd ir signifiсant rolе in thе dеvеlopmеnt of thе Univеrsity сlf Notrе Damе in Indiana and pub|isЬedЕ,uolutioпапd Dogmа in 1896. Both thеsеwritеrs suppoгtеd thе thес;logiсalaссеptaЬility of еvolution' alЬеit within сеrtain limits' Lеroy's apologia, еrесtеd foursquarе on Mivart's foundations,was dirесtеd against both аthеistiс rеnditiorrsof еvolution and thеologiсalеnеmiеsof thе thеory. But Ьесausеhе hintеd that thе human Ьody might Ье thе produсt of еvolutionary forсеs, thе сardinal prеfесt of thе SaсrеdCongrеgation of thе Indеx of ProhiЬitеd Books issuеd thе judgmеnt ..еvolution thеory that is tеmеrarious and anti-Сhristian whеn аppliеd to thе human Ьody'' (quotеd in Brundеll 2001' 88). Although thе finding that his Ьook was сеnsurеdWas сonvеyеd to him privatеly' Lеroy was dеvastаtеdand spееdilyrеtraсtеdin thе pagеs of Le Мondс in Мarсh 1895. Latеr hе triеd to find ways of rеvisitingthе wholе issuе Ьy prоduсing a nеw' сorrесtеd vеrsion of thе Ьook, Ьut it too fеll foul of thе аuthoritiеs. It was muсh thе samе with Zahm, who had alrеady aсhiеvеd widеsprеad rесognition for his Sottпd апd Миsiс (1892) and Саtholiс Sciеnсe апd Саtholiс Scieпtists(1893). Although hе had еarliеr сondеmnеd Darwin's Desсеntof Мап as thе сausе of soсial inеquity, his purposе in writing Еuolutioп апd Dogmа was to dепlonstratеits thеologiсal aссеptaЬiliry.Indееd hе found ..to admirе' muсh muсh in it that is еnnoЬling and inspiring,' (Zahп-l,|896, xх). Likеwisе groundеd in Мivart's sуstеm' Zahm engagеd in dеtаilеd patristiс еxеgеsisto justify a tеlеologiсal rеndеring of еvolution. Thе Ьook was an intеrnationalsuссеss'and an Italian translation rapidly appеarеd,Ьut rеadеrs of a fuming СiuiltЙ Саttoliса wеrе assurеd that it was all a tissuе of liеs' rеdolеnt with rесklеss assеrtions' and hopеlеssly сompromisеd Ьy duЬious assumptions.Thе Сongrеgation of the Indех dеnounсed tt in 7897, foсusing vеry largеly on tЬе mаttеr of human origins. To traditionalists' Zalrm's еvolutionismwas part of a paсkаgе of radiсаl proposals tlrat inсludеd thе adoption of tеxtual сritiсism and thе ir-rсrеasingsеgrеgation of асаdеmiс and ессlеsiastiсalaffairs. Likе Lеroy, ZaЬm withdrеw his еfforts dеspitе thе favoraЬlе rеviеw hе rесеivеd from thе Еnglish Ьishop John Hеdlеy, who' for his pains, prеsеntly сamе undеr thе whiplasЬ of СiuiltЙ Саttoliса tonguеs. .,AmеriсanVarious faсtors playеd thеir parts in thеsе intriguеs. A fеar of ism''-thе tеndеnсy on thе part of Amеriсan Catholiсs to display intеllесtual indеpеndеnсеаnd to adapt Сatholiсism to arr Amеriсan сOntеYt-was onе suсh fоrсе. This indiсаtеs that Darwinian сontrovеrsiеsanlong turn-of-thесеnturyСatholiсs wеrе oftеn lеss aЬout sсiеnсеthan aЬout issuеs of idеntity with whiсh immigrant сommunitiеs wеrе grappling. Anothеr suitе of сonсеrnsrotatеd around thе routinе сonflation of Dаrwinism and modеrn sсholarly trеnds, notaЬly thе nеw ЬiЬliсal сгitiсism. Rеaсtions to еvolutionary Ьiology Wеrе thus oftеn all of a piесе with rеsponsеs to widеr сhallеngеs to еstablishеdways of thinking. Сritiсаl too Was thе mattеr of intеllесtual аuthorityand who hаd thе right to intегpгеtthе traditionЪ rесеivеd сanon. Thosе who found еvоlution in thе writings of Aquinas аnd Augustinе wеrе

.].'.]

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Еt,olиtioп апd Religioп arrogantly failing to pay duе dеfеrеnсе to thе history of nеosсholastiс сommеntary. Latеr, fеаrs of Darwinian-inspirеd еugеniс poliсiеs playеd their rolе in shaping Сatholiс еvaluations of еvolution (ApplеЬy 1999\' Dеspitе thеsе pеrsistеnt anхiеtiеs, Huxlеy's еfforts to prosсriЬе Мivart's Сatholiс Darwinism' and thе diffiсultiеs that Lеroy andZaЬm ехpеriеnсеd at thе hands of thе Сongrеgation of thе Indех, a suссеssionof Саtholiс sсholars havе сontinuеd to dеfеnd thе thеologiсal propriеty of Darwinian biology. During thе final dесаdеs of tlrе ninеtееnthсеntury, thе Сatholiс astronomеr Gеorgе Searlе dесlarеd that thе thеory of еvolutiorrеnjoyеd a strong faсtual Ьаsis and that human еvolution would not jеopardizе Сatholiс thеology. At thе samе timе thе Harvard anatomist Thomаs Dwight insistеd that thеrе was no сontradiсtion bеtwееn еvolutiorr and tеlеology. Thе thеologiаn John Gmеinеr addеd his sllpport in 1884 whеn, in Мoderп Sсiепtific Viец,s апd Сhristiап Doctriltеs Сompаred, hе wеnt so faг аs to сlaim that Augustinе had aсtually proпlotеd a vеrsion of еvolutionary thеory and suggеstеdthat Darwin сould arguaЬly Ье сonsidеrеd his disсiplе. Pеrhaps thе most сonsidеrеd Сatholiс statеmеnt оn thе suЬjесt in tlrе еarly dесadеs of thе twеntiеth сеntury appеarеd in Frеnсh tn 1927 (and in Еnglish transiation Ьy Еrnеst Меssеngег thе following yеaг) Ьy thе dirесtoг of thе Gеologiсal Institutе at thе Univеrsity of Louvain, Сanon Hеnri dе Dorlodot, who dеfеndеd not just thе lеgitimaсy Ьut thе intrinsiс doсtrinal plausiЬility of Darwinism. Not only Wеrе thеrе no sсriptural argumеnts against thе thеory' hе insistеd in Dаrtuiпism апd Саtholiс Thought, Ьut thе ..tеaсhingof thе Fаtlrеrsof thе Сhurсh is vеry fаvouraЬlе to thе thеory of AЬsolutе Еvolution'' (Dorlodot |922,4)' Сеntral to Dorlodot's diagnosiswas his dеvеlopmеntof thе сonсеpt of what hе сallеd Сhristian naturalism. Сonсеivеd as an antidotе to athеistiс and matеrialistiс еvolution' as wеll as to thе intеrvеntionist stanсе оf figurеs likе Gеorgеs Сuviеr and Alсidе d'oгЬigny, this сonсеpt еmphasized thе immanеnt workings of thе сrеator through nаtural proсеssеs.So еmphatiс was Dorlodot in this judglrrеntthat hе pronounсеd it lеgitimatе for Catholiсs to go еvеn furthеr than Darwin had donе whеn hе attriЬutеd thе initial origirr of living things to аn aсt of spесial divinе intеrvеntion. Сatholiс thеology, hе maintainеd, prеdisposеd its adhеrеnts to an advanсеd systеm of transformism and еven oЬligеd thеm to ассеpt thе idеa thаt all living bеings wеrе dеrivеd from a fеw еlеmеntary organisrns' All of this was rеpoгtеdlу in kееping with tlrе tеaсhing of thе сhurсh fathеrs. St. Grеgory of Nyssa's aссount of origins, for ..aЬsolutе ехamplе, was paradеd as a thеory of еvolution'' that lеft nO room ..еvеn for spесial intеrvеntion at thе origin of lifе'' (Dorlodot 1922,79)' Givеn Меssеngег'srolе as translаtor of Dorlodot's volumе' it is nor surprising that hе himsеlf turnеd in 1931 to a dеtailеdanalysis of thе thеologiсal propriеty of еvolution that Ьuilt on Dordolot's ..Ьrilliant piесе of woгk'' (Mеssеngеr 193 1, xxiv). Stagеd in thе introduсtory еssay Ьy Сharlеs Souvay ..noisy as poisеd Ьеtwееn thе аdvoсatеs of Protеstant Мodеrnism'' аnd thе ..stuЬЬornranks'' of ProtеstаntFundamеntalism,Mеssеngеr'svolumе undеrtп 56, ', AR: Nlastеrbooks. rshlеgaсy:Thе prеsidеnt'ssuссеss .hеfеdеralЬеnсh a dесidеd GoP аy. Rсprlrls tlf thе Nаtioltаl Сепter in thе Unitеd States.Аnпuаl ..thеistlс sсholаrshiр,''irnd N,ltkпаI Сеntеr for Sсiепсе 149 зwеЬ.org/rеsourсеs/artiсlеs/6 Rеports of thе Nаtiсlпаl Сеntеr for эduсtioп..!7еstpоrt,СT: Grееnwоod i s m :С h а n g е sа n d с t ' n t i n u i l i е s . lеsign in sсiеnсесlassrooms. с п с e s1 О 4 ( S u p p l .\ ) : 8 6 6 9 - 8 6 7 6 . :al аnalysisof IntеlligеntDesign in -2.98. "itiquе of Iпtеlligепt Dеsigп Thесlrу. y rеirsоning.lntеrпаtkпаl J ou'rnаl s? In М. Young аnd T. Еdis' еds., itiquс о[ thс Nсш Сrсаtiсlпism' vеrsityPrеss. 1984.Thе Муstсrl, of l-ifе's origiп: hilosophiсalLibrary. сation of possiЬlеroutеs of | 'B i с l l с l g2у0 3 : 1 1 1 _ 1 1 6 . irесtor.DVD. 67 minutеs. Lir Нсlш Did lt Gсt Hеrе? 8y 7atсhtowеrBiblе and Traсt Soсiеty A сritiquе of Willirrn-r сlсolatеs: )rigiпsArсhivе (updatеd Aрril 23, qs/nfl/(aссеssеdJanuary 24, 200 5I. ЭеnеsisFlood: T.he Bibliсаl Rесord NJ: PrеsЬytеriаnаnd Rеformеd. саmе Hеdoпisfs.Downеrs Grovе, IL: r'antаgеsof thеft оvеr toil: TЬе dеsign э g уо п d P h i l r l s o p h l | 6 : 7 7 1 - 7 2 4 . ,епtDеsigп Fаils: А Sсiепtifiс nsrviсkN . J : R u t g е r sU n i v е r s i t y

сoURт oP|N|oNs с|тЕD Dапiеlu. Wаtеrs.U.S. Сourt of Appеals,Siхth Сirсuit.515 F.2d 485 (1975). Еdшаrdsu. AgиiIIаrd.482U.S. 578 {,1987I. Е p p е r s с luп. А r k а п s а s , 3 9 3U . s . 9 7 ( | 9 6 8 \ , Kitzmil!еru. Douеr Аrеа SсhoolDistriсt'400 F. Supp.2d 707 (М.D. Pa. 2005). МсLеапu. ArkапsаsBoаrd of Еduсаtion.529F. Supp.(Е.D.Ark. 1982) Sеlmапu. Сobb СoltlttуSсhoсllDistriсt.390F' Supp.2d (N.D. Gа. 2005).

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This Ьook, puЬlis .!7illiams, is onе o] thоugh nеo-Darwi lishеd paradigm сommittеd to thе t сious imprеssions. thе есologist Vеro Soсiаl Behаuiсlur grouр ovеr thе inс possiЬility of grou ought to alwаys pl vidual еffесtsand l In making his сu groundЬrеaking p (1964a,1964Ь)ш , (ants, Ьееs, and w known as kirr sе сhanсеs by raising produсtion. Anothеr topiс a progrеss' thеn mu .!7il Rusе 1996). progrеss' pointing ity) arе oftеn vеr tаkеs dеvеlopmеn сrеdiЬly сomplех plех is Ьеttеr.A jеt propеllеr еnginе, lattеr. Thе main thrus and genеrally suff sсorn Was rеsеrvе lесtion. С. H. Wa similation, whеге .\Х/illiams hас fliеs.

Аdаptаtion аnd Nаturаl Selection (Gеorgе C. Williams) This Ьook, puЬlishеd iл 1966 Ьy thе Amеriсan iсhtlryologist Gеoгgе С' Williams, is onе of thе grеat dеЬunking works irr thе history of sсiеnсе.Although nеo.Darwinism, or thе syrrthеtiсthеory of еvolution, was thе еstaЬlishеd paradigm in еvolutionary studiеs, Williams (who was dееply сommittеd to thе tlrеory itsеlf) fеlt that it was infесtеd Ьy a numЬеr of fallaсior.rsimprеssiсlns.Ivlost signifiсant of thеsе Was the Ьеliеf, promulgatеd Ьy .Wynnе-Еdrvards thе есologisr Vеro С. in Апimаl Dispеrsiotl in Rеlаtiotz to Soсiаl Bеhаuioцr (|962\, гhat stlrnеtimеsnaturаl sеlесtion сan favor thе group ovеr thе irrdrvidual.Although hе did not want to dеny thе outright possiЬilityof group sеlесtion,Y/illiams fеlt that on gгounds of simpliсity onе ought to always prеfеr thе hypothеsis that group еffесtsаrе сausеd Ьy individual еffесtsаnd tlrirtsеlесtionthегеforеrnust Ьеnefitfirst thе individual' In making his сasе, Wiiliarns was onе of thе first tO makе rеfеrеnсеto thе groundЬrеaking papеrs of thе tlrеn-graduatе studеnt \Х/illiam Нamilton (I964a, 1964ь), r,vhohad shown how thе stеrility of hymеnoptеran rvorkеrs (ants,Ьееs, and w.asps)сould Ье ехplainеd as a rеsult of what has Ьесomе known as kin sеlесtion. Thе workеrs аre improvillg thеiг rеproduсtivе сhanсеsЬy raising fеrtilе rеlativеsinstеad of putting еfforts into thеir owt] rеproduсtion. Anothеr topiс attaсkеd with rеlish Ьy \Х/illiamswаs that of еvolшtionary progrеss'thеn mrrсh favorеd Ьy pеoplе likе Julian Huxlеy (7942; sее аlso Rusе 1996). \Williams was withеring in his сritiquе of various notions of progrеss,pointing out that populaг сritеria of improvеmеnt (likе сomplех. itv) aге oftеn vеr1' rrrislеading.Аp:rгt from thе faсt tlrat gеnеrally lro onе takеsdеvеlopmеntinto aссount (thе livеr flukеs rhаt affесt shееp hаvе an inсrеdiЬlyсomplех lifе history), it is Ьy no mеans oЬvigrаmsапd Еuolиtion rat еvolution сan inhaЬit thе t tlrаt еvolutiсlnis thе dеfining t а n с е st l f . l i g i r e lе v o l u t i < l nm a y amatiсаllyaliеn lifе. If wе had rеts,it likеly wor"rldЬе саrborrсommon with lifе on еarth. Arproсеss itsеlf in hе еvolr'ltitlпаry iеn than lrfс on othеr planеts. mеdiunl; it is ph,vsiсal/сlrеmiсal n е w i n s г . t t r с е. 'si е v o l u r i o n a r е а n i с е v o l u t i o n( е . g . 't h е l a w s o f r 'д,hаtamounts t() anothег uni_ hе logiс of thе сomputеr. Thеy 1есomputегis сonstГuсteсl;thеy 1е opеrаtingsystеrn.Thеsе rulеs ir Ьеhavior.Thus thеy livе in a ' l r i n t o t h е t l i g i r a ln i с d i t r m g i v е s s and rvhat it doеs. r с r е a t i n 8t,l i g i t а lе v o l u t i o n i s t с > ..digital orriсh hаvе Ьееnсallеd lеm On a сOmputеr with a Daring systеrrrmanagеs a populatiorr aу that wЬеп nеrь"сIеatlrrеs arе nory inhaЬitеdЬу thе progгams. ldttсеsmutation Ьy flipping bits r е i т u r е sТ . hе гantlommutations itlr thе rеsult thertsomе individLr_ wlriсh lеаds to a natural proсеss ll rnеditrm. а l i е n i n s t a n с еt l f е v o l u t i o n , i t i s 3aniсеvolLrtionon еarth. Pеrhaps l o г g а n i s п rеs v o l v с a d a p t а t i o n st o :hеir еnvironmеnt.A digital есol. :ng fсlrсеof еvoltrtitlrr. s е l f - r е p l i с a t i ndgi g i г a l o r g a n i s m , mеmory сlf thе сomputеr with a thеn a vеry promillеnt fеaturе of t sourсе of sеlесtivе forсеs. Para;irlly еvolvе, arrd tlrеy sеt off an to an ongoing sеriеs of еvolutionron Ьесoпtеsа targеt for еxploitaесOmесommon' thеy drivе othеr rаrio would Ьеgin with thе еvolu-rof immunity Ьy thеir hosts. This

сyсlе сan rеpеat sеvеral timеs Ьut may progrеss to thе еvolution of ..hypеrpaгаsitеs''that aсtually аttaсk parasitеs,stеalingthеir еnеrgy. Somеtimеs,whеn onе kind of сrеаturе сomplеtеly dsophilаmelапo atе studеnt, Yuiсh Сrow, his studеnts gеnеtiсs,and еvolu Сrow's influеnсе tion to thе sсorеs о dirесtly trained, Сrr

Сroш

cеs,Jamеs Franklin Сrow has :еtiсal(mathеmatiсal)and ехp r i m а r у r е s е a r с ho r g a n i s r n s :ow has аlso madе important rlturalсrops. Although Сrow tiсs of DDT rеsistanсе to ехmеiosisto dеvеloping thеory l o s t p r o m i n е n tс о n t r i Ь u t i o n s n

lvania'Сrow grеW uP mainly undеrgraduatе dеgrее from row Wеnt to thе Univеrsity of study with prominеnt gеnе. rld latеr wirr thе NoЬеl Prizе philа. Мu||er, howеvеr, wаs Thomas (J. T') Pаttегson Ье. .!Иorld ritеd Statеs' еntry into at Dartmоuth Сollеgе, whеrе rg of his ссlllеaguеsаt Dаrtlarasitology, navigation, and эs of gеnеralzoology and gеi Wisсonsin,whеrе hе has rеrougЬmutаtions arе tlrе ultilr еvolutiorrto proсееd, dеlеttions.Сrow and his studеnts bilа mеlапogаstеrto mеаsure n a singlе сhromosomе еaсh t thе lеthal mutation ratе pеr r h r е еp е r m i l l i o n . М u t а t i o n s [aсt' Сrow аnd his сollеaguеs rs mutationsarisе at lеast 10 stцdiеs suggеstthat the ratе an thosе mutations that havе lst dеlеtеriousmutations arе ; thеy havе in hеtеrozygotеs omozygotеs.Bесausе muta. еrеdfrom sеlесtionand thus ln. Thеsе dеlеtеrious mutajесrеasеthе mеan fitnеss of I this rеduсtion of fitnеssthе d othеrs,Сгow invеstigatеd / a а n d h u m a n s ,f i n d i n g i t r o

Ье surprisingly lаrgе. Thе аvеrаgе human harЬors thе еquivalеnt of sеvеral lеthal nrutatiorrs.Thеsе lnutations аrе пrоstly rесеssivе,аnd thеir dеlеtеrious еffесtsarе not manifеst unlеss individuals inЬrееd Ьесausе unrеlаtеd in_ dividuals lrarЬor а diffеrеnt sеt of rесеssivе dеlеtеrious mutations. Thе rесеssivеnaturе of thеsе mutations is tlrr.rsthе Ьаsis of irrЬгееdingdеprеssiоn. Сrow сontinuеs to havе an intеrеst in mutations and thеir еffесts in hurrans. Сrow was thе mеntor and longtimе сollaЬorator of Мotoo Kimura. Kimura dеvеlоpесlthе nеutrаl thеory of molесular еvolution, whiсh arguеs that thе vast majtlrity of rnolесr:lаrvаriants arе еithеr dеlеtеriousor nеutral with rеspесtto sеlесtion.An impoгtant fеaturе of thе nеutral thеory is thаt it еаsily gеnегatеsmаny tеstirЬlеprеdiсtions that саn allow Ьiologists to dеtесt whеthеr and what forms of sеlесtionarе Opеratingat pаrtiсuiar gеnеtiс loсi. Laying thе foundation for thе nеutral thеory was work that Kimurа did with Сrow tсl dеtеrminе how muсh vаriation would Ье ехpесtеd, providеd that this variiltion was thе rеsult of a Ьalanсе Ьеtwееn nеutral mutations addirrg tо variatiоn and random gеnеtiсdгift еroding variation. |n \964, Kimura and Сrow dеvеlopеd aл importilnt mathеmatiсаl modеl in whiсh thеy assumеd that еvеry mutation that arisеs is uniquе. In this so-сallеd infirritе allеlеs пrodеl' thе ехpесtеd hеtеrozygosity (thе proportion of thе population that rvas hеtеrozygotlsаt a sitе) of rrеutr:alallеlеs r.rndеrmlltаtion/drift Ьalаnсе еquals fсlur timеs thе produсt of tlrе еffесtivеpopulation sizе аnd thе nеutral n.lutationratе. Сгow and Kimura also puЬlishеd a highly influеntial tехtЬсlok, Аll [пtrodисtioп tсl Pсфиlаtiсlп Gепetics Thеorу, whiсh hаs Ьееn usеd Ьy gеnеrationsof studеnts and praсtitionеrs of populаtion gеnеtiсssinсе its a p p е а r a n с еi n 1 9 7 0 , In addition to his work on thе гolе of mutation in еvolution, Сrow lras madе sеvеrаl othеr сontгiЬutions to еvolutionary gеnеtiсs. As a graduatе studеnt r-lndеrthе tutеlagе of J. T. Pattегson аt thе Univегsity of Tехаs аt Austin' Сrow was an еarly invеstigator of thе gеnеtiс Ьasis of rеproduсtivе isolating Ьarriеrs Ьеtwееn Drosopbilа spесiеs.Сrow was among thе first to study rhе gеnеtiс Ьasis оf hyЬrid fеmalе inviaЬility, firrding thаt this trait mappеd largеly to thе Х сhromosomе in hyЬrids Ьеtwееn D. mцllеri and D. аIdriсhi-2. Сrow has alsо had аn асtivе inгеrеstin gеnеtiссonfliсt and thе еvolution of Меndеlism. Invеstigationsof gеnеtiс systеms thаt violatе Mеndеl's law of еqual sеgгеgаtionhаvе hеlpеd our undеrstanding of why Mеndеlism is in part a fаir gamе. Thе Ьеst knowr.rand Ьеst сharaсtеrizеd of thеsе so-сallеd nrеiotiс drivе systеms,sеgrеgationdistortion on thе sесond сhromosomе of Drosoуlbilа mеlапogаstеr, was disсnstruсtioп of Dаrlt,iп'sTheorу.Chiсago:Univеrsityof Сhiсago Prеss. |999, TЬe еpistеmologyof historiсalintеrprеtation:progrеssivityand rесapitulation in Dаrwin'sthеory.In R. Сrеathand J. Мaiеnsсhеin,еds., Еpistеm()logуапd B ioklgу, 64_9О.CаmЬridgе:CambridgеUnivеrsityPrеss' Rltsе,R. 2003. Dсlпuin аnd Design:Does Еuolиtion Наue а Purposе?Сambridgе' МA: Harvard UnivеrsityPrеss. WеЬеr,B. Н., and D. J. Dеpеw. 2003. Li,uolution апd Lеаrпiпg:Tbе Bаldu,,iпEffeсt Rесonsiderеd. СаmЬridgе' МA: MIТ Prеss.

-D,L.H.

Dawkins, Riсhard (ь. I94|) Riсhard Dawkins is a British zoologist and еvolutionary thеorist Ьеst known for his сonсеpts of thе sеlfish gеnе, thе еxtеndеd phеnotypе, and thе mеmе. Born in Kеnya, Dawkins was еduсatеd at Oxford' whеrе hе has livеd and workеd sinсе 1970. F{is 1976 Thе Sеlfish Geпe is onе of thе сlassiс works of nеo-Darwinism, unifying and сlaгifying thе еmегgingсonsеnsusаmong suсh еvolutionary thеorists as !(/illiam Hamilton, Gеorgе Williams, RoЬегt Trivеrs, John Мaynard Smith, and his doсtoral supеrvisor, еthologist Niko Tinbеrgеn. This Ьook artiсulаtrs thе thеory of nаtural sеlесtion in suсh vivid and aссеssiЬlе tеrms that it is oftеn mistakеnly rеgardеd as mеrеly a ..popular'' Ьook. Dawkins's сеntral сlaim is that thе most fundamеntal' and hеnсе most еxplanatory, pеrspесtivе on all еvolutlon by natural sеlесtion is thе .,gеnеЪ-еyе viеw,,' in whiсh Ьеnеfits to spесiеs, linеagеs, groups' and еvеn individual organisms arе sееn to Ье suЬsidiаry to thе primary Ьеnеfiсiariеs of all adapta..sеlfish'' gеnеs tions' thе thеmsеlvеs. ovеr timе' еvоlution designs and Ьuilds organisms (..survivalпraсhinеs'')that Ьеnеfitсoalitions of gеnеsЬy improving thеir prospесts for rеpliсation. TЬе геaсh of gеnеs doеs not stop at thе skin of thе oгganism: soпlе gеnеs сontrol thе dеsign and сonstruсtion of an еxtеndеd phеnotype' harnеssing fеaturеs of thе еnvironmеnt and еvеn othеr spесiеs. Thus thе bеavег'sdаm and thе spidеr's wеЬ arе just as important paгts of thе phеnotypеs of thosе spесiеs as thеiг еyеs and mouths' аnd parasitеs oftеn ехploit thе Ьеhaviorаl сontrols of thе host spесiеs' hiiaсking othеr gеnеs' survival

mасhinеs. Еmphasizing t . awkirl m е n t а l i r l s i g h t sD nattlral sеlесtion сгеatеd; that inсludеd languagе ar of еvolution Ьy natural sе сation of saliеnt сulturirl fеaturеs of сulturаl еvolu prеsеntеd thе idеil of mе аЬstraсtnеss of thе funс сlаimеd to Ье making a n еvolution of human сult thusiastiсally' with mixес In 1986' Dаlvkins put Wаtсbmаker, in wlriсh h е v o l u t i o n a r у r h е с l r yv i а thе еarly triumphs of th 1995' hе was appointеd liс Undеrstanding оf Sсi talеnts' hе has subsеquе plaining aspесtsof sсiеn thе puЬliс disсussionso N4ost rесеntly, his Ьroa tеmрt to raisе сonsсiou rеligious Ьеliеf,has attrа a l е a d i n g r o l е i n w l r a ti s thе lay puЬliс arе Dawk kin sеlесtion and thе еvс вlвLIoGRAPHY

Dawkins,R' 1976' Thе Sе |982. The Ехtсnd 1986.Тhе Bliпd \ 2О04.Thе Апсcst 2ОО6,Thе Glнtiсlltsstri'1tеgiе: optimiеruпgТechnisсhеr Sуstеmе lmd Priпzipiеп dеr Biologisсh еn Еuсllutiсlп. Stuttgart: Frommann-HolzЬoog. Turing, A. 1950. Сomputing maсhinеry and intеlligеnсe.Мiпd 59 43З-460.

fеlt Ьy many palеonto (Darwin 1859, 36.1). In еvolutionary disс many еvolutionists, Ьu philosophiсally and sс lutionary progrеss har tеndеnсy to rеad into wе think wе sее in hu two indеpеndеnt сorr сlaim that lоng_tегmd trеnd has Ьееn for thе of a long-tеrm trеnd is сrеasing. But what, еx progrеss in what sеns disсoursе has еnaЬlеd Disсцssions of еvol gists, palеontologists l е y ( 1 8 9 3 ) ,H е r Ь е r t S (1'961"), Lеdyard Stеb son (1949),Stеphеn Уa|en (1973)' Gееrat (|996}, and othеrs.Ir has Ьееn rеlеgatеdto has rеmainеd сеntral vating faсtor Ьеhind thе twеntiеth сеntur) disсussion of this valr

-Е.Е.v. апdС.Е.T. PRoGRЕss IN

Еvolutionаry prоgrеss Мodеrn organisms sееm сlearly morе advanсеd than anсiеnt onеs. Сomparе an еlеphant with a modеrn Ьaсtеrium. (Thе сomparison is apt in that сеrtain modеrn bасtеria aге prоЬaЬly vеry similaг to thе first fossil orgаnisms known, frоm roсks 3..5Ьillion yеars old') Thе notion of an ordеring among organisms datеs at lеast to Aristotlе, who arrangеd living forms on a linеar sсalе Ьasеd On dеgгее of pеrfесtion, also сallеd the scаlа псlturсlеor grеat сhain of Ьеing' For morе than 2'000 yеars aftеr Aristotlе, thе сhain Was undеrstood to Ье statiс' mеaning that organisms and thеir rаnkings did not сhangе. But in thе еаrly ninеtееnthсеntury Jеan-Baptistеdе Lamarсk addеd aп еvolutionary сomponеnt' so that orgаnisms пrovеd up thе сhain as thеy еvolved. A half сеntury latеr thе Darwinian viеw сhallеngеd thе ordеring itself. Dаrwin sаw еvolution aS a proсеss of Ьranсhing and divеrgеnсеrаthеr thirn linеar asсеnt. But Darwin nеvеrthеlеssrесognizеd progrеss' wгiting famously in on thе Оrigiп of Spеcies of ..that vaguе yеt ill-dеfinеd sеntimеnt'

Thеrе arе two Ways iI vаluе. Thе tеaturесo tionary proсеss' so to standpoint thе first a] tural (геligious,soсiа Indееd, thеy arе а di ality in еvolution w( fеatцrе wе valuе, su сoursе' wе might Ьt rathеr than trеnds gе vorеd in еvolution l trеnd' and that wе vz

hеrе too thе еvaluat as original motivatiс a trеnd in fеaturеsvi Thе sесond altеrn

Еuolиtionаrу Progrеss d morphology to automatiсally work Ьy Мurata and Yamaguсhi 'nfigurеroЬots that сan assеmЬlе 'n' sеarсhing through a spасе of ng through a spaсе of сomputеr tallizеd to сrеatе thе sсiеntifiс enl d h a s p r o v е n u s е f u Ii n a v a r i е t y ving promisе for many сomplеx

,he FossilReсord.Nеw York: IЕЕЕ ArtificiаISуstеms:Aп Iпtroduсtorу -roIапd Аrtificiаl Iпtеlligепсе. Progrаmmiпgof Сomputеrs bу \: MIT Prеss. еsignand manufaсturеof robotiс ry сomputation:An ovеrvrеw. 30:593-616. ing сrossovеrto improvе GA-Ьasеd apеrprеsеntеdat thе Proсееdingsof ry Сomputation,Washington,DC, ieruпgTесhnisсherSуstеmеund gart:Frommann-HolzЬoog. lligеnсе. Мiпd 59:4зз-460.

fеltЬy many palеontologists'that organisation on thе wholе has progressеd'' ( D a r w i n1 8 5 9 , 3 6 3 ) . In еvolutionary disсoursе sinсе Darwin' progrеss has Ьееn aсknowlеdgеd Ьy many еvolutionists, Ьut thе idеa hаs also сomе to Ье sееn as trouЬlеsomе, Ьoth philosophiсally and sсiеntifiсally.It is widеly rесognizеd that intuitions on еvo.Wе lutionary progrеss havе Ьееn сontaminatеd Ьy сultural influеnсеs. havе a tеndеnсyto rеad into thе history of lifе thе soсial and tесhnologiсal progrеss wе think we sее in human affairs. Also' progrеss is now rесognizеd as having rwо indеpendеnt сomponеnts that arе frеquеntly сonflatеd, a valuе-nеutral сlaim that long-tеrm dirесtional сhangе has oссurrеd and a valuе сlaim that thе trеndhas Ьееn for thе Ьеttеr.Among сontеmporary еvolutionists thе еХistеnсе of a long-tеrm trеnd is widеly aссеptеd.Most would agrее that somеthing is inсrеasing.But what, ехaсtly? And inсrеasing in what sеnsе?Is it progrеss? If so, progrеssin what sеnsе? Thеsе issuеs hаvе not Ьееn rеsоlvеd, Ьut thе modеrn disсoursеhas еnaЬlеd us at least to formulatе thеm fairly сlеarly. Disсussions of еvolutionary progrеss сan Ье found in thе works of Ьiologists,palеontolоgists, philosophеrs' and historians, inсluding Thomas Huхlеy (1893)' HеrЬеrt Spеnсеr (1857), Julian Huxlеy (1942)' Thomаs Goudgе (1961)'Lеdyard StеЬЬins (1,969),Еrnst Мayr (1988), Gеorgе Gaylord Simpson (1949), Stеphеn Jay Gould (1988), Franсisсo Ауa\a (\974), Lеigh Van Уa|en (1973)' Gееrat Vеrmеij (1987), RoЬеrt Riсhards (1'992),Мiсhaеl Rusе (1'996|,and othеrs. Intеrеstingly' among thе sсiеntists most of the disсussion has Ьееn rеlеgatеd to popular works. Rusе arguеs that thе notion of progrеss has rеmainеd сеntral in еvolutionary studiеs, oftеn its proof thе сhiеf motivating faсtor Ьеhind thе work that pеoplе do, Ьut that in thе sесond half of thе twеntiеth сеntury thе inсrеasing profеssionalization of thе fiеld forсеd disсussion of this valuе.ladеn suЬiесt out of thе tесhniсal litеraturе.

-E'.Е,.v.апdС.Е'T. PRoGRЕss lN WHAт sЕNsЕ?

lсеd than anсiеnt onеs. Сomparе сomparison is apt in that сеrtain ar to thе fiгst fossil orgаnisms .hе notionof аn ordеring among trrangеd living forms on a linеar :allеd thе sсаlа паtLrrае or gteat rftеr Arisrotlе, thе сhain was unisms and thеir rankings did not Jеan-Baptistе dе Lamarсk addеd sms movеd up thе сhain as thеy i viеw сhallеngеd thе ordеring itЬranсhing and divеrgеnсе rathеr ; rесognizеd progrеss' writing fa.t vaguе yеt ill-dеfinеd sеntimеnt'

Thеrе arе two ways in whiсh a trеnd in somе fеaturе of organisms might havе valuе.Thе fеaturе сould Ье valцaЬlе to us, or it сould Ье valuеd Ьy thе еvolutionary proсеss' so to spеak, in thе sеnsе of Ьеing pгеsеrvеd. From a sсiеntifiс standpointthе first altеrnativеis problеmatiс. In sсiеnсеour pеrsonal or сultural (rеligious,soсial) valuеs shoцld not Ье rеlеvant or at lеast not сеntral. Indееd,thеy arе a distraсtion. For еxamplе, if thе aсtual dеfining dirесtionality in evolution wеrе inсrеasing еnеrgy usagе' an infatuation with somе fеaturеwе valuе, suсh as intеlligеnсе' might сausе us to ovеrlook it. of сoursе' wе might Ье intеrеstеd in thе еvolutionaгy sourсеs of our valuеs rathеrthan trеnds gеnеrally. For еxamplе, it might Ье that intеlligеnсеis favorеd in еvolution by natural sеlесtion, that wе arе thе produсt of that trеnd,and that wе valuе intеlligеnсеon aссount of its advantagеsfor us. But hеrе too thе еvaluativе сomponеnt of our intеrеst is Ьеsidе thе point, еХсеpt as original motivation. Thus it is hard to sее how progrеss' if it is dеfinеd as a trеnd in fеaturеsvaluaЬlе to us, сould Ье a sсiеntifiсally usеful notion. Thе sесond altеrnativе is to intеrprеt valuaЬlе to mеan that whiсh is prе.

551

552

Еuolutioпаrу Рrogress sеrvеd' or' Ьесausе Wе arе intеrеstеd in trеnds, that whiсh inсrеases ovеr timе. Progrеss thеn Ьесomеs а purеly sсientifiс tеrm, Ьut now douЬts arisе aЬout thе aptness оf thе word. It might Ьe apt if thе еvolutionary proсеss prе. sеrvеs fеaturеs that strikе us as positivе, likе intеlligеnсе, fitnеss, or сomplехity. But supposе wе disсovеr that thе main long-tеrm trеnd is an inсrеasе in a fеaturе aЬoцt whiсh wе arе amЬivalеnt' likе еnеrgy usagе' or a fеaturе that sееms downright Ьad, likе fragility (lеading to an inсrеasing proЬaЬility of ехtinсtion). It would sound odd to сall еithеr of thеsе progrеss. I think that thеsе proЬlеms are fatal, and wе should aЬandon talk of еvolutionary progrеss and instеad dеvotе our еnеrgiеsto thе study of trеnds. But thе notion of progrеss has grеat сultural momеntum and sееms likеly to pеrsist' at lеast outsidе thе tесhniсal litеraturе, for somе timе to сomе. In what follows, I aссеdе to thе folliеs of my timе and usе thе word pro?rеss, Ьut only in thе sесond sеnsе,to mеan that whiсh is prеsеrvеd.For сonvеniеnсе,I will also usе thе phrasе degrее of аduапсement, aS though thе variaЬlеs involvеd in progrеss werе еithеr known or mеasuraЬlе. (As will Ье sееn' at prеsеnt thеу arе not.)

с) Е

advanсem

A тRЕND IN WHAт sЕNsЕ? Disсussion of progrеss has Ьееn muddlеd, еvеn rесеntly' Ьy сonfusion about how trеnds in groups work. In сurrеnt trеnd thеory it is not еnough to say that somе variaЬlе inсrеasеs.To Ье сlеar' onе must spесify whiсh of sеvегal group statistiсs is inсrеаsing-mеan' mаximum, or minimum-Ьесausе trеnds in thе diffеrеnt statistiсs havе diffеrеnt intеrprеtations. A trеnd in thе mean is an inсrеasе in thе avеragе dеgrее of advanсеmеnt among all spесiеs in еxistеnсе at a givеn timе. In thе figurе, A shows a hypothеtiсal long-tеrm trеnd. Lifе Ьеgins with a singlе spесiеs' a singlе linеagе' with sоmе low lеvеl of advanсеmеnt.As timе passеs,nеw spесlеsаrisе, and sоmе spесiеsЬесomе еxtinсt' with сhange oссurring in the origin of еvеry new spесiеs. Thе mеan at anУ givеn timе sliсе is thе avеragе lеvеl of advanсеmеnt of all spесiеs in еxistеnсе at that timе. A trеnd is an inсrеasе in that avеragе' that is, movеmеnt of thе avеragе to thе right. A сlaim that a trеnd has oссurгеd, without any qualifiеr, usually mеans an inсrеasеin thе mеan. A trеnd in thе maхimum is a risе in thе dеgrее of advanсеmеnt aсhiеvеd by thе most advanсеd spесiеs at a givеn timе, in othеr words, a risе in thе highеst lеvеl of advanсеmеnt aсhiеvеd. (In thе figurе, B is thе samе as A, Ьut annotatеd to show thе trеnd in thе maхimum' thе dottеd linе on thе right.) Мaхima arе of spесial intеrеst Ьесausе humans arе gеnеrally thought to rеprеsеntthе most progrеssivе spесiеs in еxistеnсе and, if so, would rеprеsеnt thе mахimum tn a .W.е timе sliсe that inсludеd thе prеsеnt. would Ье thе last spесiеson thе right at thе vеry top of thе graph. Finally, a trеnd in thе minimum is an inсrеasе in thе dеgrее of advаnсеmеnt of thе lеast advanсеd spесiеs (thе dottеd linе on thе lеft in figurе B). Мinima are of spесial intеrеst Ьесausеapparеntly thе lowеst lеvеl of advanсеmеnt has not сhangеd. Baсtеriа havе ехistеd, and indееd havе dominatеd thе Ьiosphеге, throughout thе history of lifе. (In othеr words, it is сlеar that figurеs A and B arе not a good rеprеsеntationof thе history of lifе.)

(A) A hypothеtiсallong Ьеgins with a singlе spе vertiсal Ьlaсk linе sеgm arisе,and somе spесiеs evеry nеw spесiеs.Hеrе strongly drivеn. (B) Saп thе maximum and mini prеdominate'Ьut a nuп whiсh inсrеasеsand dес (thе vеrtiсаl dottеd linе disсussion.

Disсussion of progr trеnd' pеrhaps Ьесaus mеasurе it). Rathеr, it mесhanism. In figurе , that all сhangеs arе iп сrеasе. Figurе С is thе mеaning that sеlесtion thе еvolution of сеrtair advanсеd spесiеs pеrs minimum stays rough driuеп, or morе prесis Figurе D is diffеrеnt in Inсrеasеs and dесгеasе symmеtriсally' Ьoth u1 limit on dеgrее of adv

Еuolutionаrу Progrеss in trеnds, that whiсh inсrеаsеs ovеr sсiеntifiс tеrm, Ьut now doubts arisе ; bе apt if thе еvolutionary proсеss prее, likе iпtеlligеnсе,fitnеss, or сomplеxn a i n l o п g - t е г mt г е n d i s a n i n с г е a s еi n а ;rt' likе еnеrgy usagе' or a fеaturе that еading to an inсrеasing proЬability of l еithеrof thеsеProgrеss. , and wе should aЬandon talk of еvoluг еnеrgiеsto thе study of trеnds. But thе lomеntumаnd sееmslikеly to pеrsist, at 'r somе timе to сomе. In what follows, I э thе word progrеss, Ьut only in thе sесгvеd.Foг сonvеniеnсе,I will also usе thе lugh thе variaЬlеs involvеd in progrеss will bе sееn' at prеSеntthеy arе not.)

(l) Е

Strongly driven

Weakly driven

adVanсement ? maх

B dlеd, еvеn rесеntly' by сonfusion аЬout nt trеnd thеory it is not еnough to say lеar, oпе must spесify whiсh of sеvеral n. maximttm. оr minimum-Ьесausе diffеrеnt intеrprеtations. A trеnd in thе :grееof advanсеmеnt among all spесiеs gurе, A shows a hypothеtiсal long-term еs,a singlе linеagе,with somе low lеvеl 'spесiеsarisе' and somе spесiеs Ьесomе : origin of еvеry nеw spесiеs.Thе mеаn еvеl of:rdvаnсеnrеntof all spесiеs in ехrеasе in that avеragе,that is, movеmеnt that a trеnd has oссцrrеd' without any n thе mеan' l thе dеgrееof advanсеmеnt aсhiеvеd Ьy гimе,irr othеr words, a risе in thе highеst : figurе,B is thе samе as A, Ьut annotatеd hе dottеd linе on thе right.) Maхima arе з gеnеrallythought to rеpгеsеnt thе most f s o , w o u l d г е p r е s е ntth е m а х i m u m i n a Ъ would bе thе last spесiеson thе right at .еndin thе minimum is an inсrеasе in thе vanсеdspесiеs(thе dottеd linе on thе lеft :еrеstЬесausеapparеntly thе lowеst lеvеl |tеriahavе ехistеd, and indееd havе domhistоry of lifе. (In othеr words, it is сlеar )prеsеntationof thе history of lifе.)

(A) A hypothеtiсallong-tеrmtrend in advanсеmеnt'that is, prоgrеss.Lifе bеginswith a singlеspесiеs,with somе lоw lеvеl o{ advanсеmеni(thеsiпglе vеrtiсalblaсk linе sеgmеntat thе Ьottom lеft).As timе passеs,nеw spесiеs arise,and somе spесiеsЬесomеехtinсt,with сhangеoссuгringin thе origin оf еvеrynеW spесiеs.Hеrе all сhаngеsarе inсrеаsеs,so thе trеnd is said to Ье stronglydrivеn. (B) Samе as A, with dotted lines addеd to show thе trеndsin thе maximum and minimum. (с) A wеаklу drivеn trеnd in whiсh inсrеasеs pгеdominatе,Ьut a numЬеr of dесгеasеsаlso oссur. (D) A passivеtrеnd in whiсh inсrеаsеsаnd dесrеasеsarе еqually frеquеnr,Ьut а lowеr limit on сhаngе (thеvеrtiсaldottеd linе) forсеslineagеsto diffusеto thе гight.Sееteхt for d i s с us s i o n . Disсussion of progrеss has not Ьееn сonсеrnеd with thе ехistеnсе of a trеnd, pеrhaps Ьесausе:1trеnd is takеn for grantеd (dеspitеouг inability to mеasurеit). Rathеr, it has Ьееn aЬout thе pattеrn of сhangе, thе undеrlying mесhanism. In figurе А, natural selесtion strongly favors аdvanсemеnr' so that all сhangеs arе inсrеasеs.Thе mеan, maхimum, and minimum all inсrеasе' Figurе С is thе samе ехсеpt that thе nеt sеlесtivе advantagе is wеak, mеаning that sеlесtion Somеtimеs favors dесrеasеs.(Тhе сlassiс ехamplеs arе thе еvolution of сеrtain rеduсеd forms, likе parаsitеs.)Thе геsult is that lеss advanсеd spесiеs pеrsist or arе rеplaсed Ьy dесrеasеsfrom aЬovе' and thе minimum stays roughly thе samе. Тhе trеnds in A and С arе sаid to Ье driuеn, or morе prесisеly,A is strоngly dгivеn, whеrеas С is wеakly drivеn. Figurе D is diffеrеnt in that thеrе is no ovеrall sеlесtivе advantagе to inсrеasе. Inсrеasеsand dесrеasеsoссur еqually oftеn' so thе only геndеnсyis tо sprеad symmеtriсally, Ьoth up and down. Howevеr' thеrе is a Ьoundary, a lowеr limit оn dеgrее of advanсеmеnt,that Ьloсks thе sprеad of thе group on thе

5 54

Еuolutionаrу Рrсlgress lеft. This Ьoundary сan Ье thought of аs thе lowеst lеvеl of advanсеmеntсonsistеnt with bеing аlivе. Thе rеsult is that tirе mеаn and maximum inсrеasе whilе thе minimurn stays thе samе (as in thе wеаkly drivеn сasе).Thе trеnd is said to Ьe pаssiuе. Thе сurrеnt сontrovеrsy ovеr mесhanism Ьеgan in 1996 with а Ьook on progrеss Ьy StеphеnJay Gould in whiсh hе raisеd thе passivе mесhanism to thе lеvеl of plausiЬility. At issuе is thе rolе of sеlесtion in progrеss, that is, whеthег thе (prеsumеd)trеnd in thе history of lifе is wеakly dгivеn Ьy sеlесtion or mеrеly passivе.(A strong|ydгivеn mесhanism is rulеd out Ьесausеthе truе minimum is thought to havе rеmainеd staЬlе: Ьaсtеria pеrsist.) Gould h:rs bееn widеly nrisintеrpгеtеd as сlaiming that еvolution is random, Ьut thе iпrpliсation of thе passivе mесhanisпris not randomnеss.Within еaсh еvolving linеagе sеlесtion doеs opеratе, and organisms arе adapting. Rathеr, thе impliсаtion is that еvolving linеagеsas a group аrе govеrnеd Ьy many diffеrеnt sеlесtivе forсеs, сomplехly сonfigurеd' so that sеlесtion favors inсrеasеs and dесrеasеsеqually oftеn. Thе proсеss сan Ье undеrstood аs purеly dеtеrministiс, likе thе diffusion of a gаs. Unfoгtunatеly,as will Ье sееn,thеrе arе hardly аny data availaЬlе,so from a sсiеntifiс pеrspесtivе thе dеЬatе aЬout mесhаnism is taking plirсе in a nеar vaсuum. Still, thеory has produсеd somе insights.Foг ехamplе, it is now сlеar that in thе dеbatе ovеr mесhanism two сommon oЬsеrvations arе irrelеvant. Thе first is thе pеrsistеnсеof nonprogrеssivе spесiеs,thе staЬlе minimrrm. This oЬsеrvation is not hеlpful Ьесausе tlrе minimum is prеdiсtеd to rеmain unсhangеd in Ьoth сasеs, passivе and wеakly drivеrr.Likеwisе, thе гisе in thе maximum and thе сurrеnt ехistеnсе сlf somе highly advanсеd spесiеs,likе hun-tans,arе not hеlpful. A rising mахimum is ехpесtеd in all сasеs. A тRЕND IN WнAТ? Мany сandid:rtеsfor a long-tеrnrtrеnd havе Ьееn proposеd: (1) fitnеss;(2) сomplехity; (3) aЬility to sеnsе' сontrol' or rеspond to thе еnvironmеnt; (4) Ьody sizе; (5) intеlligеnсе;(6) vеrsatility or еvolvability' mеaning thе aЬility ttl сhangе in еvolцtion; (7) thе rеvеrsе of еvolvaЬility, аlso сallеd dеgrее of еntrеnсhmеnt;(8) еnеrgy intеnsivеnеssoг ratе ofеnеrgy usagе;and (9) dеpth and sophistiсationof mесhanismsof inhеritanсе'For thе most paгt advoсatеs of all thеsе сandidatеs havе takеn a сasе-Ьuilding аpproaсh, with thе rеsult tlrat thе litеraturе offеrs a numЬеr of сlеvеr and somеtimеs-powеrfulargumеnts in favoг of еaсh. Unfortцnаtеly, somе of thе most intеrеsting-likе fitnеss, сomplехiry, еnеrgy intеnsivеnеss, and aЬility to сontrol thе еnvironmеnt-arе diffiсult to opеrationalizе. Thus thеrе аrе hardlу any data on thе sсalе of lifе as a wholе, for any trеrrd statistiс, for аny of thеsе сandidatе variaЬlеs. For somе of thеsе variablеs, many wоuld want to сount a long.tеrm trеnd, if onе сould Ье doсumеntеd, as progrеss.onе variаЬlе is fitnеss,цndеrstood as aЬility to sцrvivе and rеproduсе. [t might sееm that Darwin,s prinсiplе of natural sеlесtion virtually guarantееs a trеnd in fitnеss. Latеr organisms

shoцld Ье moге fit thl ..in tJ Darwin's tеrrns, also knеw, natuгаl sеl on gеologiсal timеsсal duсе just сonstant сh; nеss. On thе othеr adaptations, suiting tl vorеd ovеr thosе morе offer advantagеs on ti Thеrе arе somе Ьiolo suсh as сеrtain сеllr"rl pathwаys. But wе do rigorous way, or how that is сlеar now is thz arе possiblе in prinсip Thе most widеly rес As a possiЬlе Ьasis ior сonnесtion with progr usеd in еvolutionaгy st for progrеss, supеrfiсiа suЬtly сonnoting advar nition makеs this usag onе сan say anythinga tain a tесhniсal dеfinit sегrsеsof сomplехity. l typеs, numЬеr of physi dеvеlopmеnt (sееfigur donе, and with onе еЬееn dосцmеntеd.Nor in сhangе in thеsе vаri somе еvidеnсе within hаs bееn a trеnd in nur Thе еxсеption is с< nеstеd, to somе ехtе within wholеs' ovеr Ьеrs of lеvеls havе еvс Ьaсtеria. AЬout 2 bili Ьiotiс rеlationship to protozoans suсh as l сеlls, Ьut Ьесаusе thе nеxt lеvеl up. Thеn а joinеd to produсе thс gaе), thе anсеstor of а lias to muskrats. Finа individuals joinеd to сallеd a Ьryozoirn. V

Еuolutioпаrу Progrеss ,Wеstlеvеl of advanсеmеnt сoni m е a na n d m a х i m u r n i n с r е а s е ,еaklydrivеn сasе).Thе trеnd is эеgan in 1996 wit.Ь a Ьook on аisеdthе pаssivе mесhаnism to lf sеlесtion in progrеss, that is, lf lifе is wеakly drivеn Ьy sеlес:hanismis rulеd out Ьесarrsеthе staЬlе:Ьaсtеria pеrsist.) Gould lat еvolutiоn is random, but thе .s7ithin еaсh еvolvandomnеss. risms arе adapting. Rathеr, thе lp arе govеrnеd Ьy many diffеrl that sеlесtion favors inсrеasеs Ье undеrstood as purеly dеtеrrdly any data availaЬlе, so from ranism is taking plaсе in a nеar hts. For ехamplе, it is now сlеar non oЬsеrvations arе irrеlеvant. эесiеs'thе staЬlе minimum. This l u m i s p г е d i с t е dt o r е m а i n u n lrivеn. Likеwisе, thе risе in thе righly advanсеd spесiеs' likе huкpесtеdin all сasеs.

: bееn proposеd: (1) fitnеss; (2) :еspondto thе еnvironmеnt; (4) :volvaЬility, mеaning thе aЬility olvaЬility, also сallеd dеgrее of е of еnеrgyusagе;and (9) dеpth nсе.For thе most part advoсatеs ilding approaсh, with thе rеsult and somеtimеs-powеrfulargurе of thе most intеrеsting-likе and aЬility to сontrol thе zе. Thus thеrе arе hardly any trеnd statistiс,for any of thеsе vant to сount a long-tеrm trеnd, rе variablе is fitnеss, undеrstood sееmthat Darwin's prinсiplе of :nd in fitnеss. Latеr organisms

should Ье morе fit than еarliеr onеs' on thе wholе, having Ьеatеn thеm, in Darwin's tеrms'..in thе raсе for lifе'' (Darwin 1859,363). But as Darwin also knеw, nаtural sеlесtion produсеs adaptation to loсal еnvironmеnts, and on gеologiсaltimеsсalеsthеsесhangе dramatiсally.Thus sеlесtionmight pгoduсе just сonstant сhangе with no nеt improvеmеnt' no nеt inсrеasе in fitnеss. On thе othеr hand, on long timеsсalеs organisms with gеnеral аdaptations, suiting thеm to a widеr variеty of еnvironmеnts' сould Ье favorеd ovеr thosе morе narrowly adaptеd. Arе thеrе gеnеral adaptations that offеr advantagеson timеsсalеsof hundrеds of millions or Ьillions of yеars? Thеrе arе somе Ьiologiсal mесhanisms that may havе pеrsistеd that long, suсh as сеrtain сеllular transport mесhanisms and parts of сеrtain mеtаЬoliс pathways. But wе do not know how to idеntify gеnеral adaptations in any rigorous way' or how to tеst whеthеr thеy havе bесomе morе pгеvalеnt. All that is сlеar now is that Ьoth altеrnativеs, a long-tеrm trеnd and its aЬsenсe, arе possiЬlе in prinсiplе, and thеrеforеthis is an еmpiriсal issuе. Thе most widеly rесognizеd сandidаtе for a long-tеrm trеnd is сomplеxity. As a possiblе Ьasis for progrеss it is rеally a stеalth сandidatе Ьесausе a dirесt сonnесtion with progrеss is rarеly aсknowlеdgеd. But thе way сomplеxity is usеd in еvolutionary studiеs suggеststhat it funсtions as a kind of сodе word for progrеss, supеrfiсially valuе frее and thеrеforе sсiеntifiс sounding Ьut still suЬtly сonnoting advanсеmеnt. Thе aЬsеnсе of a widеly known tесhniсal dеfinition makеs this usagе problеmatiс. NoЬody knows what сomplехity is, so onе сan say anything at all about it. Thе litеraturе of rесеnt dесadеs doеs сontain a tесhniсal dеfinition' or rathеr sеvеral dеfinitions for еaсh of thе sеvеral sеnsеsof сomplеxity. Thеsе inсludе сomplехity in thе sеnsе of numЬеr of part typеs,numbеr of physiologiсal or Ьеhavioral proсеssеs'and numЬеr of stеps in dеvеlopmеnt (sееfigurе on pagе 556). Howеvеr' littlе еmpiriсal work has Ьееn donе, and with onе еxсеption' no trеnd that spans thе history of lifе has yеt bееn doсumentеd. Nor do wе know vеry muсh aЬout thе mесhanisms involvеd in сhangе in thеsе variaЬlеs, passivе vеrsus drivеn. (On a smallеr sсalе thеrе is somе еvidеnсе within thе animals, ovеr thе past 500 million yеars' thаt thеrе has bееn a trеnd in numЬеr of part typеs and that it has Ьееn passivе.) Thе ехсеption is сomplехity in thе sеnsе of nеstеdnеss.organisms arе nеstеd, to somе еxtеnt' likе Сhinеsе Ьoхеs, with multiplе lеvеls of parts within wholеs. Ovеr thе history of lifе, оrganisms with еvеr grеatеr num' Ьеrs of lеvеls havе еvolvеd. Thе first living things 3.5 billion yеars ago wеrе Ьaсtеria. AЬout 2 Ьillion yеars ago somе Ьaсtеria joinеd togеthеr in a symbiotiс rеiationship to produсе thе first еukaryotiс сеlls, similаr to modеrn protozoans suсh as Аmoеbа' Thеy wеге supеrfiсially similar to baсtеrial сеlls, Ьut Ьесausе thеy еvolvеd as сoloniеs of Ьaсtеria, thеy oссupiеd thе nеxt lеvеl up. Thеn aЬout 600 million yеars ago сlonеs of еukaryotiс сеlls joinеd to produсе thе first multiсеllular individual (proЬaЬly a kind of algaе),thе anсеstor of all modеrn multiсеIlulаrs' from mushrooms to magnolias to muskrats. Finally' aЬout 480 million yеars ago сlonеs of multiсеllular individuals joinеd to form thе first сolonial animаl' a сoral-likе animal сallеd a Ьryozoan. Мodеrn сolonial organisms inсludе Ьryozoans, сorals'

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Еuolиtionаrу Progrеss

A сlassiс ехamplе of why dеfinitions of progressin tеrms of movеs to сomplехity arе proЬlеmatiс.Thе uppеr skеlеtonis of a lion and thе lower skеlеtonis of a whale. By any measurеmеntthе ЬaсkЬonеof thе whale is thе simpler' and yеt adaptivеly the whаlе's ЬaсkЬonе is what is nееdеdfor a martnе mammal, just аs thе lion's ЬaсkЬonе is what is nееdedfor a tеrrеstrialprеdator. Whalеs dеsсеndedfrom land tеtrapods(four-lеggеdanimаls)sO thеir bасkЬonеsmark a sеlесtion-drivеn move from сomplехityto simpliсity.(From MсShea 199|. з|5.\

thе soсial insесts, and many vеrtеЬratеs,from fish to humans. !Ие do not know if еvеn highеr lеvеls-сoloniеs of сoloniеs-havе Ьееn rеaсhеd. Thеrе arе no сlеar-сцt сasеs. Humans sеem to аssoсiatе at many lеvеls' Ьut only wеakly at thе highеr onеs. Intеrеstingly,if humans havе not tгuly rеaсhеd thе сolony-of-сoloniеs lеvеl, аnd if no othеr organism has еithеr, this raisеs a novеl possiЬility: thе only variaЬlе in whiсh wе сan doсumеnt a trеnd, сomplеxity in thе sеnsе of nеstеdnеss,in faсt doеs show a trеnd, Ьut that trеnd might havе еndеd, pеrhаps 480 million yеars аgo. Pеrhaps thе agе of progrеss is ovеr. BIBLIOGRAPHY Ayala,F. J,1974. Thе сonсеptof Ьiologiсalprogrеss. In F. J. Ayala and T. Dobzhansky,еds',Studiеsiп thе Philostlphуof Biologу, З39_354.London: \4aсmillan. Goudgе,T . ^. 1961. Thе Asсent of Lifе. Toronto:Univеrsityof Toronto Prеss. Gould' S. J. 1988.On rеpiaсingthе idеaof progrеsswith an opеrafionаlnotionof dirесtionality. In М. H. Nitесki, ed.,Еuolutiсlпаrу Progrеss,319-з38.Сhiсago: Univеrsityof Сhiсago Prеss. 1996. Full Hoиse: Thе Sprеаdсlf ЕхсеIlепсefrсlmPIаto to Dаrшiп' Nеw York: Hаrmony Books. Huxlеy'I. S.1942. Еuolиtioп:Thе Мodеrn Sуnthesis. London:Allеn аnd Unwin. Нuхiеy,T. Н. 1893.Еuolutiсlnапd Еtbiсs.London:Мaсmillan.

Mаyr. Е. 1988.Toшаrd, Еu сlluti сlпi st.СаrnЬ M с S h е a 'D . W . 1 9 9 1 .С о апd Рhilosсlphу6, n 1 9 9 8 .P o s s i Ь llеa hypothеsеs... Аппuа RiсhаrdsR . . J . | 9 9 2 .т h ( апd ldеologiсаl Rес Сhiсago Prеss. R u s е 'М . | 9 9 6 .у , , n u 7' , Сambridgе,МA: На Simpson,G. G. 1949.Th Prеss. S p е n с е rH' . l 8 5 7 . P r o g r StеЬЬins'G. L. |969, Тh North Сarolina Prеs Van Valеn,L. 1973.A nе Vеrmеij,G. J. 1987.Еuо Prеss.

Еvolutionary psyс

S o m е Ь е h а v i o r а |s с i е study of human Ьеhav In this Ьroad sеnsе,еvс q u i r y a r е d е f i n е dn o t l Ьut Ьy thе kinds of quе fеr from paradigms' W ologiеs with whiсh thе; a fiеld of inquiry, еvolu I i с а t i o no f Т h е D е s с е п iп Маn апd Aпimаls in t i с u l а t е d p a г a d i g m sЬ е d i s t i n с t t h е o r i е sа Ь o u t human Ьеhavior and m Thе first of thеsе pa thе 1970s. Thе сorе idс n a t u r a Ia n d s е х u а |s е I t many spесiеs сhoosе m. mаlе сourtship display sеlесtion will tailor m w a у . s o с i o Ь i o I o g i s ras1 рrеsеnсе in a populatiс thеory. Aссordingiy' hr nations of how huma anothеr-to ехplain hr с h i е s , f o r m a n i f е s t i n gl thе сеntrаl thеorеtiса

Еuolutioпаrу

Psусhologу

Mayr, Е. 1988. Тotuаrd а Nеtu Philosophу of Biologу: Оbseruаtioпs of ап Еuolutioпist. CаrnЬridgе,NlА: Bеlknар Prеss оf Hагvirrd Univеrsity Prеss. .Whаt МсShеа, D. !и. 199 [. Сomplехity and еvolution: еvеryЬody knows. Вiсllogу апd Pbilosoph1'6, no.3: 303_325. 1998. PossiЬlе largеst-sсalеtrеnds iгr orgаnismlrlеvolution: Еight ..livе hypotlrеsеs.''Aпп uаI Reuieш'сlf Есolog1,ап d Sуstепlаti сs 29 : 29 3_31'8. -Гhe Riсhards, R. J. 1992. Мetllliпg of Еuсllutioп: Thе Мсlrpholtlgiса! Сoпstruсtioп апd Idеologiсаl Rectlnstruсtion of Dаrшiп's Tbеorу. Сhiсago; Univеrsity of Сhiсago Prеss. Rusе, М. |996. Мoпаt! to Маn: Tbе Сoпсеpt tlf Progrеssiп Еuolutiсlпаrу Bitllogу. Саmbгidgе, МA: Hаrvаrd Univеrsity Prеss. Simpson, G. G. 1949, Thе Меапiпg of Еuolutiсlп' Nеw Havеn' СT: Yаlе Univеrsity Prеss. Spеnсеr,H. 1857. Progrеss:Irs lаw and сausе. Weslt,instеr Rеuiеtl,67:244-267. Stеbbins, G. L, 1969. Thе Bаsis of PrсlgrеssiuеЕuolutioп, Сhapеl Hill: Unir.еrsity of North Сarolina Prеss. Vаn Valеп, L. |97з. A nеw еvolutionаry lаw. Еuolutitlпаrу Thеor1'1: 1-30. Vеrmеij, G. .I' 1987. Еuсllutionапd ЕsсаIаtiспz.Prinсеton' NJ: Pгinсеton Univеrsity

in tеrmsof movеs to is of a lion and thе lowег Ьaсkbonеof thе whаlе is the е i s w h а t i s n е е d е dt o г а m a r i n е reеdеdfor a tеrrеstrialprеdatoг. 4gеdanimals)so thеir omplехitvto simpliсity.(From

lm fish to humans. !7е do not riеs-havе Ьееn rеaсhеd. Thеrе эсiatе at many lеvеls, Ьut only tumans havе not truly rеaсhеd o r g a n i s mh a s е i t h е r ,t h i s r a i s е s iсh wе сan doсumеnt a trеnd, сt doеs show a trеnd, Ьut that 1 yеars ago. Pеrhaps thе agе of

:ss.In F. J. Ayala and ,у of Biolсlgу, 3з9-З54. London: Univеrsityof Toronto Prеss. ss with аn opеfational notion of паrу Progrеss,319_338. Chiсago: се froп Plаto to Dаrшiп. . ' s i sL. o n d t l n :A I l е n a n d U n w i n . n;Мaсmillan'

Prеss.

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Еvolutionary psyсhology Somе Ьеhavioral sсiеntists dеfine еvolutionary psyсhology as simply thе study of human Ьеhavior and psyсhology from an еvolutionary pеrspесtivе. In this Ьгoad sеnsе,еvolutionary psyсhology is а fiеld of inquiry. Fiеlds of inquiry arе dеfinеd not Ьy speсifiс thеoriеs aЬout thе phеnomеna thеy study, but Ьy thе kinds of quеstionsthеy posе aЬout thеm. Fiеlds of inquiry thus diffеr fronr paradigms, whiсh аrе dеfinеd Ьy thе spесifiс thеoriеs and nrеthodologiеs with whiсh thеy answеr thе quеstions that dеfinе a fiеld of inquiry. As a fiеld of inquiry, еvolutionaгypsyсhology Ьеgan with Charlеs Darwin's puЬliсation of Tbe Descепt of Мап in ] 871 and Tbе Ехpressiсlll rlf the Еmotions in Мап апd Апiпшls tn 7872.It was not until a сеntury li1tегthat сlеarly artiсulatеd pаradigms Ьеgan to emеrgе in еvolutionaгy psyсhology, еaсh with distinсt thеoriеs about' and spесifiс mеthods for studying, thе еvolution.of humаn Ьеhаvior urndmеntillity. Thе first of thesе paradigms was human soсioЬiology, whiсh еmегgеd in thе 1970s. Thе сorе idеa of soсioЬiology was that Ьеhavior has еvolvеd undеr natural and sехuаl sеlесtionIustаs org:rniсform has. For ехamplе, fеmalеsof many spесiеsсhoosе matеs Ьаsеdon thе quality of malе сor-rгtship displays. If malе сourtship displays vary in quality, and that variation is hеritablе,sехual sеlесtion will tailor malе сotlгtship Ьеhavior to fеmalе prеfеrеnсе.In this way, soсioЬiologistsarguеd' a form of Ьеhavior сan Ье an ildaptation' whosе prеsеnсеin a population is to Ье ехplainеd Ьy thе prinсiplеs оf еvolutionary thеory. Aссоrdingly, humаn soсioЬiology sought to offеr еvolutionary ехplanations of how humаns arе Ьеhavioraily adаptеd to soсial lifе with onе anothеr-to еxplzrin human Ьеhavioral adaptаtions for dominanсе hiеrar. с h i е s , f o r m а n i Г е s t i n ga r t d d е a l i n g w i t h a g g r е s s i o n а, n d Г o г m a t i n g . l n d е е d , thе сеntrirl thеorеtiсal proЬlегr-rof humirn soсioЬiology Was to ехplain thе

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Еuolutiслtаrу Psусhologу еvolution of altruism, thе pеrformanсе of aсts thаt Ьеnеfitothеrs at a fitnеss сost to thе асtor. In thе fiеld of еvolutionarypsyсhology today, lrumаn soсioЬiologyhas Ьееn supеrsеdеdЬy sеvеralaltеrnativеparadigms' two of whiсh arе partiсularly notaЬlе. Onе of thеsе сonfusingly goеs Ьy thе namе ..еvсllutionаrypsyсhology.'' Тo avoid сonfusion, and to distinguislrthе fiеld оf inс1шiryfrom thе pаradigm, (сapitаlizеd). I will rеfеr to this par:rdigmas ..ЕvolutiсrnaryPs,vсhсllrlgy'' Еvolutionary Psyсhology is a maгriаgе of сognitivе psyсhology with еvolutionary Ьiology. In сontrаst with soсioЬiology, Еvolutionarv Psyсhologists arguе thаt, whеn z-rЬеhavior hаs еvolvеd undеr sеlесtion, it is not thе Ьеhavioг thаt is аn ad:rptаtion, Ьut rathеr thе psyсh 7 9 - 5 9 3 .

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Ford, Еdmund Brisсo (1901-1988) Е. B. Ford, usually сallеd Hеnгy Ьy lris сollеaЕ.uеsаnd friеnds, is Ьеst known as thе сhiеf proponеnt of thе rеsеarсhprogram known as есologiсal gеnеtiсs. Intеndеd to throw light on thе rolе of gеnеtiс divеrsity in thе adaptation of organisms to thеir еnvironmеnts, eсologiсal gеnеtiсs Ьеgins with oЬsеrva. tions of variation in nаtural populations. Brееding studiеsthеn dеtеrminеthе mannеr of inhеritanсе of thе diffеrеnt variаtions. Cоmparativе analysеs of diffеrеnt haЬitats and сorrеlations of еnvironmеntal and gепеtiс variaЬlеs lеad to hypothеsеsаЬout thе rolе of variation in еvolutionary adaptation. Finally' ехpеrimеntal manipulation of еnvironmеntal variaЬlеs rеsults in thе сonfirmation or rеjесtion of thе hypothеsеs.Ford's rеsеarсhphilosophy is sеt out in lris sеminal Ьook, Есolс-lgiсаlGenеtiсs. Ford spеnt his еrrtiresсiеntifiс сarееr at Oхford Univеrsity' Hе matriсulatеd as an undеrgraduatеin 192О, reсeivinga BA in zоology iп |924 and a BSс in \927. tle wаs appointеd Dеpartmеntal Dеmonstrator in Zoo|ogу (1927|, thеn Univеrsity Dеmonstrator (1929) and Lесturеr at Univеrsity сol. lеgе (1933). Iп 19З9, hе Ьесaпrе Rеadеr in Gеnеtiсs, thе first еxpliсit аppointmеnt in that suЬjесtin oхford. Hе еstaЬlishеdthе Gеnеtiс l-aЬoratoriеs in 1951 and rесеivеd an appointmеnt as Profеssor of Есologiсal Gеnеtiсs in

1.96З, a post hе hеll of All Souls Сollеgt сеivеd its Darwin Nz Ford сamе to thе еst in thе Lеpidоptе thеr of a populatio: Their pеrsonal оЬsе сality rесords going thе papеr puЬlishеd aЬility of thе popult lесtion, whiсh allolr hе maintainеd his i on thеsе animals ir Ьеstsеllеrfor thе sеr Ford's earliеstwс lеy of a gеnе that af. cheureuхi. Thеy sh gеnе' but also оn t} growth. Although I gеnеs' tЬe Gаmmаr Ford is pеrhaps was thе first to offс parеd for Julian Ht Ford's ЕсologiсаI С rеnсе toBеthеr in th spесiеs in suсh pro1 rесurrеnt mutatlon whiсh onе morph i anсеd polymorphis nеtiс polymorphisr among loсi сontrit ..su what hе сallеd phеnotypеs' Batеsi amplеs of supеrgеn Although not gi R. A. Fishеr in thе ] thе Sсarlеt Тigеr m Ford was aЬlе to tr yеarly сеnsusеs.Fi quеnсy wеrе too g1 сontеntion at thаt of еffесtivеpopula that thе original an Ford was also in populations. For п thе hindwings of ,V

Ford in thе influеnza sеquеnсе that rаt will сausе futltrе еpidеmiсs. ast еvеnts' thе status quo for alсtion o{ futurе еvolutionary trarinking and prеsсiеnсе that has rolесulаr еvolutiоnist.

on: Wеidеnfеldаnd Niсolson. .W. 'x, and М. Fitсh. 1999. \. Sсiепcе286: 1921-7925. .om anаlogоusprotеins. Sуstеmаtiс v:rriаblесodons.Jourпаl of эlution:Мinimum сhangе for a ):406416. .Гreпds in Gemе of thе proЬlеms. | . С o х . 1 9 9 7 . L o n g t е r m t r е n d si n t,vpеA. Proсееdiпgs сlf thе 7.-7718. :tiоrrof phylogеnеtiс treеs. Sсienсе for dеtеrmining сodon эvеd r-rrетhod е rаtеtlf fiхlrion tlf rnut.rtionsin

-r.K.l.

)a8uеsand friеnds, is Ьеst known 21In known as есologiсal gеnеtiсs. ltiс divеrsity in thе adaptation of al gеnеtiсsЬеgins with oЬsеrvaееdingstudiеsthеn dеtеrminе thе iations.Сomparativе analysеs of rоnmеntal and gеnеtiс variaЬlеs ln in еvolutionaryadaptation. Fi)nmеntal variaЬlеs rеsults in thе . Ford's rеsеarсhphilosophy is sеt o х f o r d U n i v е r s i t 1. H е m а t r i с u rg a BA in zoology iп |924 and a rеntаl Dеmonstrator in Zoo|ogу 9) and Lесturеr at Univеrsity Сolin Gеnеtiсs, thе first ехpliсit ap. aЬlishеdthе Gеnеtiс LaL.loratoriеs rofеssorof Есoloяiсal Gеnеtiсs in

196З, a post hе hеld until his rеtirепrеntin \969. }-ord was еlесrеda Fеllow of All Souls Сollеgе in 1958. Hе was a Реilow of thе Royаl Soсiеty аnd rесeivеdits Darwin Меdal tn 1954. Ford сamе to thе study of gеnеtiсs and еvolution through his lifеlong intеrеst in thе Lеpidoptеra. Bеforе attеnding Oхford, hе Ьеgan a study with his fathеr of a population of Меlitаеа (Еиphуdrаs) аuriпiа, thе Marsh Fгitillary. Thеiг pеrsonal oЬsеrvationsеmЬraсеd thе yеаrs from 1917 to 1935, and loсаlity rесords going Ьасk to 1881 mаke this an unusually lеngthy study. In thе papеr puЬlislrеdwith his fathеr in 1930, Ford attriЬutеdthе unusual variability of thе population during pеriods of aЬundanсе to thе rеlaxation of sеlесtion,whiсh allowеd suЬoptimal phеnotypеsto survivе.Throughout his lifе hе maintainеd his irrtеrеstin moths and Ьцttеrfliеs,prrЬlishingtwo volumеs on thеsе animals in Сoliins's Nеw Naturalist sеriеs. Ilts Bltttеrfljes wаs a Ьеstsеllеrfor thе sеriеs. Ford's еarliеstwork at Oхford was a сollaЬorativе study with Julian Huхlеy of a genе that affесtеd thе ratе of darkеning of tlrе еyе сolor in Gаmmаrlls chеureuхi. Thеy showеd that thе гirtе dеpеnds not only on thе Mеndеlian gеnе,[.lutаlso on thе sizе of thе еyе, thе tеmpеraturе'and thе ratе of Ьodily growth. Although not thе first to аttгiЬutе еvolr'rtionaryimportanсе to ratе gеnеs'thе Gаmmаrus studiеsprovidеd an important еarly ехamplе. Ford is pеrhaps Ьеst known fоr his woгk on gеnеtiс polymorphism. Hе Was thе first to offer a сritiсal dеfinition of thе phеnomеnon in a papеr prеpaгеd for Julian Huxlеу,s The Nеtu S1'5ngmаtiсs(1940), rеfinеd as follows in FordЪ,Есologiсаl Gепеtiсs (|964,84). ..Gеnеtiсpolymorphisrrris thе oссurrеnсе togеthеr in thе samе loсality of two or morе disсontinuous forms of a spесiеsin suсh proportions that thе rarеst of thепr сannot Ье maintainеd Ьy rесurrеnt mшtation.'' Hе distinguishеd Ьеtwееn transiеnt polymorphism, in whiсh onе morpЬ is in thе proсеss of rеplaсing anothеr, and staЬlе or Ьalаnсеd polymorphism, maintainеd by staЬilizing sеlесtion.Hе notеd thL1tgеnеtiс polymorphism oftеn irrvolvеd largе phеnotypiс еffесts, сlosе linkagе among loсi сontriЬuting to thosе еffесts, and strong sеlесtion, lеading to ..supеrgеnеs.'' what hе сallеd Hе intеrprеtеdthе ABo Ьlood grouрs' Rhеsus phеnotypеs, Batеsian mimiсry in Ьuttеrfliеs, and hеtеrostyly in plants as еxamplеs of supеrgеnеs'a position Ьornе out Ьy suЬsеquеntrеsеarсh. Although not givеn to mathеmatiсal nrodеling, Ford сollаЬorаtеd with R. A. Fishеr tn thе |947 study of а сolor polymorphismin Pапахiа dсlmiпulа, thе Sсarlеt Tigеr moth. Sinсе hеtеrozygotеs at this loсus arе distinguishaЬlе, Ford was aЬlе to traсk gеnе frеquеnсy and populаtion sizе in a long sеriеsof yеarly сеnsusеs.Fishеr,s analysis еstaЬlishеd that thе сhangеs in gеnе frе. quеnсy wеrе too gгеаt to Ье Ьrought aЬout Ьy gеr.rеtiс drift аlonе, a point of сontеntion at that timе. Although сhallеngеd Ьy Sеwall \Иright on thе Ьasis of еffесtivеpopulation sizеs,suЬsеquentwork on this сolony has еstaЬlishеd t h а t t h е o r i g i n a l a n a l y s i sW а s с o r г е с t . Ford was also intеrеstеdin thе еffесtsof polygеnеs on variation in natural populations' For many yеars hе studiеd variation in thе pattегns of spots on thе hindwings of Мапiсllа ;jиrtiпа,thе Mеadow Brown Ьuttеrfly. Tlrе pattеrns

587

.'88

Fсlrm and Funсtiсlгr wеrе геmark:rblyсonstant ovеr widе arеas of Еuropе, Ьut thеy shoц,еdgrеat сhzrngеsovеr slight distarrсеsin sor.rthwеstЕngland arrd thе Islеs of Sсillv' Spot pattеrns on thе lаrgе islands tеndеd tсl Ье similar, whilе tlrosе from smallеr islands diffеrеd rаdiсally. Ford attriЬutеd tl"rесonstanсy сlf pattеrn ovеr timе to staЬilizing sсlесtion and thе diffеrеnсе bеtwееn largе .rnd small islаnds to thе еffесt of avеrirgingovеr divеrsе haЬitаts orr thе largе islands. In addition to his gеnеtiс woгk' Ford madе сorrtriЬutionsirr sеvеral othеr fiеlds. Не puЬlishеd on thе сhеmistry of Ьuttеrfly pigllrеnts'on thе arсhaеology of southwеst Еnglаnd' and on thе сontеnts of сhurсhеs in thе Oхford distriсt. Pеrhaps Forсl'sgrеaсеstirrrрilсton thе str'rdyof vari:ltihnR. G. d was pеrhapstoo irrсlinеd to a r rС l а r k е( I 9 9 5 , 1 6 2 ) m a k е s ,,stt||Еcologiсаl Geпеюir, thе hеld, still it is thе сlеaгеst i o r l i s rv. i е w ,а n d s t i l l i t i s а j o у

саl Mеtпоirs сlf FеIloш's of thе 4. Еvolutionarystudiеs on Iеrеtlit1,1'9 47 |-488. g r l l еi n n з т t t г l lс o t l t l i т i t l n isl l а |it1,1: 11З_174' Сhарrllan аnd Наil. -onr.1tln:

-1.til.

еll) iquеlyvaluaЬlеrеsourсе in tlrе loraтionof thе most irnportarrt lrough thе еarly twеntiеth сеnlhologistsЬе livеly? Thе livеlis tlrеthеoгistsaround a сеntral h сеnturyand pеrsistsеvеn toпсtitlп сoa|d jr-lstas wеll havе -rattеrsimplу, whiсh сamе first, tavеthе forrnsthеy do? Do thе r sеrvеd?If so, funсtiorralism is t. or arе orgenismiсforms or еrаtingproсеssеs?lf thе lattеr,

thеn tlrе funсtions that Ьody рarts sеrvе arе thе Ьy.produсts' llot thе сausеs ..puге morphology.'' of fоrпr. Rrrssеll rеfеrrеd to thе prсl-f Galton, Franсis (1'822_1911)

ipеlago,whеrе Pеtеr and f Darwin'sfinсhеs.Thе эlown oif and a сratеr fоrmеd o work out rеlаtionships quеntiаlordеr in whiсh thе е еafth'splatеsslippеd around

гinationsfor tourists. Darwin's ls provеn tо havе thе longеr.

апd Аdаptаtiсlп iп thе Gаldpаgt.>s ss. Dупаmiсs of а Nаtиrаl Populаs. Сhiсagсl:Univеrsity of Сhiсago уiп'sFiпсhеs.Pгinсеton,NJ: rcs.SсiспtifiсАmeriсап, oсtoЬеr: сrоеvolutionaryrеsponsеsto ,olиtion49:241-251. е RаdiаtЙlttсlf Dапuiп's Finсhes. пbridgеUnir,еrsityPrеss. d Sсicпсе tlп thе Gаldpаgos ;les.Nеr,l,York: СolumЬia

Franсis Galton Was born nеar Birmingh:rm,Еngland. Hе originаlly intеndеd to pursuе a сarееr in mеdiсinе, Ьut hе switсhеd to СamЬridgе Univеrsity to study mathеmatiсs' influеnсеd Ьy his сOusin Сharlеs D;rгwin (1809_1882). Darwin and Galton wеrе grandsons Ьy differеnt marriagеs of Еrasmus Dаrwin (1731_1802),a physiсian' sсiеntist,poег' аnd invеntor. Сharlеs Darwin would Ье a сruсial influеnсе on Galton throughout muсh оf his сarееr. In faсt, it was Galton's rеading of oп the o rigiп of Speсiеs (1 859) that gavе him thе idеa of improving tlrе human raсе throllgh sеlесtivеЬrеесling,for whiсh hе сoinеd thе tеrm сltЗеniL.s. Sevеral yеaгs aftеr grаduating from СamЬridgе with aп ordinary dеgrее, Galton оrganizеd and finanсеd an Afriсan ехpеdition that rеsultеd in thе first Еuropеan ехploration of northеrn NamiЬia. Hе made сarеful mеasurеmеntsof latitudеs, longitudеs' and altitudеs, rеflесting his lifеlong intеrеst in thе appliсation of numеriсal and quantitativе mеthoс.lsto whatеvеr happеnеd to intеrеsr hinr. Не puЬlishеd his rеsr'rltsin tЬеlourпttl rуarе ехplorations of сasеs,suсh аs dorninanсе,sехual sеlесtiоn,аnd rnimiсry' tO supрort thе prесеdingthеorеtiсalwork' Nеvеrthеlеss,in thе fourth and fifth сhaptеrs' Fishеr ехpands his thеorеtiсal disсussion to morе gеrrеral issuеs сotlсеrning thе сausеsof gеnеtiсvariation' irrсiudingrаndom gеnеtiс drift. Thе last fivе сhaptеrs of Gепetiсаl Thеorу ехplorе natural sеlесtion in human populations' partiсularly soсial sеlесtiorrin lruman fЪrtility. Fishеr's сеntral oЬsеrvation is that thе dеvеlopmеnt of есonomiеs in humаn soсiеtiеs struсturеs thе Ьirth ratе so that it is invеrtеd with rеspесt to soсial сlass. In thе final сlraptеr,Fishеr offеrs stratеgiеsfor сountеring this еffесt.Dеspitе

602

The Gеnеtiс Basis of ЕvоlutionaryСhangе Fishеr's еspousal of this еugеniсs thеsis in this part of thе Ьook, hе intеnds thе disсussion to Ье takеn as an insеparaЬlе ехtеnsion of thе prесеding part. Tbe Gепetiсаl Thеorу of Nаturаl Seleсtioп is a point of dеparturе in еvolu. tionary thought, rеsponsiЬlе in part for thе originаtion of thеorеtiсal population gеnеtiсs and what is сommonly сallеd thе modеrn synthеtiс thеory of .!Иright's еvolution. Gепetiсаl Theorу was followеd Ьy Sеwall and J. B. S. Нaldanе's major works in 19З1 and 1,9З2,rеspесtivеly. Fishеr's viеws on thе rolе of natural sеlесtionin еvolution arе widеly aссеotеdtodаv. в|вLIoGRAPHY Fishеr,R. A. 1930 [1958].Тhе GепeticаlТhеorуof NаturаlSеlесtioп.2nd еd.Nеw York: Dovеr Publiсations.Rеlеasеdin a variorum еdition by oхford Univеrsity Рrеssin 1999, editedЬy J. H. Bеnnеtt. Haldanе,J. B. s. 19З2.The Саusеsof Еuolиtion.London:Longmans. .Wright, s. 1931.Еvolutionin Меndеlianpopulations.Gепеtiсs16:97-159.

-R.А.s.

Thе Genetiс Bаsis of Euolшtionаrу Сbапge (Riсhard Lеwоntin) In1974, Riсhard Lеwontin, a profеssor at Hаrvard Univеrsity, puЬlishеd an important assеssmеnt of thе statе of еvolutionary gеnеtiсs. Lеwontin's 1974 Ьook,The Gепetiс Bаsis of Еuolutioпаrу Сhапge, сonsidеrеd thе trеmеndous сhangеs within evolutionary gеnеtiсs wrought Ьy thе introduсtion of tесhniquеs from molесular Ьiology and thе suЬsеquеnt dеvеlopmеnt of thе nеutral thеory of molесular еvolution. Lеwontin's Ьook is notеworthy for its analysis of thе naturе of еvolutionary gеnеtiсs and thе dynamiсs of sсiеntifiс сontfovеrsy and сontrovеrsy rеsolution. Lеwontin and his сollеaguе, J. L. HuЬЬy, hаd touсhеd off a molесular revоlution in 1966 whеn thеy introduсеd thе bioсhеmiсal tесhniquе of еlесtrophorеsis as a mеans of rеsolving gеnеtiс diffеrеnсеs at thе molесular lеvеl. Lеwontin Ьеliеvеd that this nеw еlесtrophorеtiс analysis of gеnеtiс variaЬility rеsolvеd thе еarliеr disputе Ьеtwееn thе сlassiсal and balanсе positions ovеr variability and sеlесtion. Thе high lеvеls of variaЬility detесtеd with еlесtrophorеsis sееmеd to support thе Ьalanсе position and thе importanсе of Ьalanсеd polymorphisms in еvolution. The сlassiсаl position's Ьеliеf was that most mutations wеrе dеlеtеriousand most wеrе sеlесtionpurifying. Thеy strugglеd to ехplain еlесtrophorеtiсvariaЬility until Мotoo Kimura and othеr Ьiologists Ьеgan to arguе that most of thе dеtесtеd polymorphism was in faсt nеutral, nеithеr sеlесtеdfor nor against. In Тbе Geпetiс Bаsis of Еuolиtioпаrу Сhапge, Lеwontin еmphasizеd thе сontinuity Ьеtwееn thе еarliеr сlassiсalЬalanсе сontrovеrsy and thе thеn-raging nеutralist-sеlесtionist сontrovеrsy. For him, thе nеutralist position was rеally just thе сlassiсal position Ьrought in linе with nеwеr moleсular rеsults. Whilе this intеrprеtation of thе сontrovеrsy makеs a goоd сasе foг сontinuity in еvolutionary gеnеtiсS, it nеglесts thе growing сontriЬution of bioсhеmiсal rеsеarсh to thе dеvеlopmеnt of molесular еvolution.

Having arguеd that Lеwоntin advanсеs an tiсs is inеvitaЬle. The с rеsult of a dееpеr,irrе and nеutralist position aЬility; thеir fondnеss сеpting оr rеjесtingvа еmЬraсеd сhangе and Ье aсtivеly maintainеd hеld сonviсtions rеgar his еlесtrophorеtiс dat inspirеd its transformа Tbe Gепetiс Bаsis o1 duсing еvolutionary g сarеful analysis of thе . сapturеs how gеnеtiс 1 Lеwontin, informatioп Ьody of data that wou and rеfinеd. In light of rеlationship bеtwееn t] tion, Lеwontin arguеd plaсеd with modеls th: Indееd, Tbe Gеnеtic B thе rеal proЬlеms of е standing thе intеraсtiv

вlвLIoGRAPHY

Diеtriсh'М. R. 1994.Th lourпаl of thе Histo Нubby, J. L., and R. С. l hеtеrozygosity in na Drosophilа psеudoo Kimura,М. 1983.TЙeN СambridgеUnivеrs Lеwontin,R. С. 1974.Тt Сolumbia Univеrsit Lеwontin'R. С.' аndJ. L hеtеrozygсlsity in nа hеtеrozygosityin na 542595-609.

Geпetics аnd the I (ThеodosiusDoЬz

With grеat сharisma a Ьесamе a rallying poin giп of Spесies (1'937\l

Gеnеtiсsand thе origin of Spесiеs 60З part of thе Ьook' hе intеnds thе rsion of thе prесеding part. is a point of dеparturе in еvolu.iginationof thеorеtiсal popula:hе modеrn synthеtiс thеory of .tХ/right,s and J. B. S' ly Sеwall spесtivеly.Fisher's viеws on thе :ly aссеptеdtoday.

f NаturаlSеIеctioп.2ndеd. Nеw ' r u mе d i t i o nЬ y o х f o r d U n i v е r s i t y London:Longmans. о n s .G е п с t i с s1 6 29 7 _ | , , - * . o . , .

Сhаnge lаrvard Univеrsity' publishеd an )nаry gеnеtiсs.Lеwontin's 1974 шge, сolsiderеd thе trеmеndous ;ht Ьy thе intrоduсtion of tесhеquеnt dеvеlopmеnt of thе nеuin's Ьook is notеworthy for its сs and thе dynamiсs of sсiеntifiс

Having arguеd that onе сontrovеrsy has Ьееn transformеd into anothеr' Lеwontin advanсеs an ехplanаtion for why сontrovеrsy in еvolutionary gеnеtiсs is inеvitaЬlе. Thе сonnесtion and pеrsistеnсеof thеsе сontrovеrsiеs WaS a rеsцlt of a dееpеr, irrеsolvaЬlе idеologiсal сonfliсt. Advoсatеs of thе сlassiсal and nеutralist positions hеld a сonsегvativе attitudе toward сhangе and vаriaЬility; thеir fondnеss for staЬility lеd thеm to sее sеlесtion as a mеans of aссеpting or rеjесting variations. Advoсаtеs of thе Ьalanсе position' howеvеr, еmbraсеdсhangе and variaЬility; thеy favorеd thе idеa that variation сould Ье aсtivеly maintainеd in a population Ьy Ьalanсing sеlесtion.Thеsе dееply hеld сonviсtions rеgarding thе valuе of сhangе еxplainеd for Lеwontin why his еlесtrophorеtiс data did not rеsolvе thе сlassiсal-balanсе сontrovеrsy, but inspirеd its transformation. Thе Gепetiс Bаsis of Еuolutionаrу Сhаngе should not bе undеrstood as rеduсing еvolutionary gеnеtiсs to сonfliсting idеologiеs. Lеwontin prеsеnts a сarеful analysis of thе сhаllеngеs of produсing a thеory of gеnеtiс сhangе that сapturеs how gеnеtiс proсеssеs intеraсt and dеvеlop ovеr timе. Aссording to Lеwоntin, information aЬout molесular level сhangеs was finally providing a Ьody of data that would allow thеoriеs of еvolutionary gеnеtiсs to Ье tеstеd and rеfinеd. In light of nеw molесular data, Lеwontin сlaimеd that thе еntirе rеlationship Ьеtwееn thеory and faсt would havе to Ье rесonsidеrеd. In addition, Lеwontin arguеd that thеoriеs of individual gеnеs would havе to Ье rеplaсеd with modеls that сonsidеrеd gеnomеs and thеir сomplех intеraсtions. Indееd' Тhе Gепеtic Bаsis of Еuolutionаrу Сhапge еnds with thе сhargе that thе rеal proЬlеms of еvolutionary gеllеtiсs сan only Ье addrеssеd Ьy undеrstanding thе intеraсtivе сontехt of allеlеs. вIвLIoGRAP|.]Y

ad touсhеd off a molесular rеvoЬioсhеmiсal tесhniquе of еlес. Liffеrеnсеsat thе molесular lеvеl. :tiс analysis of gеnеtiс variaЬility ;siсalаnd Ьаlanсе positions ovег f variaЬility dеtесtеd with еlесposition and thе importanсе of :lassiсalposition's Ьеliеf was that rе sеlесtionpurifying. Thеy strug. rtil Motoo Kimura and othеr bi:есtеdpolymorphism was in faсt ,he Gеnetic Bаsis of Еuolutioпаrу ity Ьеtwееn thе еarliеr сlassiсalсontrovеrsy. еutra1ist-sеlесtionist Ьrought in position сlassiсal ;t thе

Diеtriсh,М. R. 1994. Thе originsof thе nеutralthеoryof molесulаrеvolution. Jourпаl of thе l{istorу of Biologу 27: 21_59. HubЬy,J. L., and R. С. Lеwontin.1'966.A molесularapproaсhto thе studyof gеniс hеtеrozygosity in naturalpоpulationsI. Thе numЬеrof allеlеsat diffеrеntloсi in Drosophilаpsеudoobscиrа. Gепеtiсs54 577_594. Kimuгa,М. 1983. Tbе Nеиtrаl Thеorу of Мolесulаr Еuollltioп. СamЬridgе: СamЬridgеUnivеrsityPrеss. Lеwontin,R. С. 1974. Тhе Gепеtiс Bаsisof ЕuolиtioпаrуСhапge.Nеw York: ColumЬiaUnivеrsityPrеss. Lеwontin,R. С., and J. L. НuЬЬy.1966.Ь molесularapproaсhto thе studyof gеniс hеtеrozygosity in naturalpopulations.II. Amount of variationand dеgrееof hеtеrozygosityin naturalpopulationsof Drosophilа psеиdoobsсurа.Gепеtics -М.R.D. 54:595-609.

intеrprеtationof thе сontroversy rtionary gеnеtiсs' it nеglесts thе :h to thе dеvеlopmеnt of molесu-

.!Иith grеat сhаrisma and a passion fоr thе subjесt' Thеodosius DoЬzhansky Ьесamе a rallying point for thе еvolutionary synthеsis. Geпеtiсs апd the origiп of Speсies (19З7) was his prinсipal сontriЬution to that movеmеnt' and

Genetiсs аnd thе Оrigiп of Speciеs (ThеоdosiusDoЬzhansky)

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Gеnеtiсsand thе Origin of Spесiеs it Ьесamе an еssеntial tеХt in еvolutionary Ьiolоgy. On onе |evel, Geпеtiсs is a splеndid summary of еvolutionary gеnеtiсs in thе 1930s' Ьoth zoologiсal and Ьotaniсal. It Ьrought togеthеr divеrsе information and сrеatеd an еasy-to.undеrstаndmodеl for еvoiution lеading to nеw spесiеs.It also prеsеntеd an unеxpесtеd lеvеl of sophistiсation and сorrfidеnсеto Ьiologists suspiсious of еvolutionary studiеs. Likе no othеr сontriЬution to the suЬject,,Gепetiсs gave a Ьurst of еnеrgy to spесiation studiеs as a resеarсh program. i\4uсh of Gепetiсs survеyеd knowlеdgе aЬout thе gеnеtiс struсturе of populations. Reсеnt dеvеlopmеnts in teсhniquе had givеn gеnеtiсistssеnsitivе tools for monitoring nеW mutations, сhangеs in gеnе frеquеnсy, and сhangеs in сhrоmosomе struсturе. DoЬzhansky prеsеntеd сonsidеrablе data on thе еxtеnt of this variation in Ьoth laЬoratory and wild populations. Hе also dеsсriЬеd еxpеrimentаl systеmsthat monitorеd сhangеsto thе gеnеtiсstгuсturе of populations ovеr timе and ovеr сhanging еnvironmеntal сonditions. In part, DoЬzhansky's survry сеlеЬratеdthе hard-еarnеdеmpiriсal sidе of population gеnеtiсs.It also aimеd to prеsеntthе suЬjесtas rigorous, with thе еvo. lutionary sсiеntistsin full ехpеrimеntalсontrol of thеir data. DoЬzhansky dеfinеs еvolution in narrow tеrms: a сhаngе in thе frеquеnсy of allеlеs within a population. This is purposivеly opеrational. ceпetiсs offеrs many еxamplеs of suсh a сhange.It also prеsentsa dеtailed disсussionоf mесhanisms driving thosе сhangеs.This disсussion,along with thе latеr disсussion of isolation and divеrgеnсе,makе for thе two thеorеtiсal spinеs in Geпetiсs. Dоbzhansky arguеd natural sеlесtion drivеs adaptаtion and is a prinсipal agеnt in shifting thе gеnеtiс struсturе of populations. Sеlесtion usеs thе raw matеrial of gеnomе variation rеsulting from random mutation as wеll as from thе many othеr gеnеtiс and сhromosomаl pгoсеssеsat Work within organisms. Although potеnt' sеlесtionwas not DoЬzhansky's only еvolutionary mесh.sИright's..shifting anism. Hе was hеаvily influеnсеd Ьy Sеwall Ьalanсе'' thе.Wright's ory of population gеnеtiсs and notion of adаptivе landsсapеs (sее \Хiright 19З7, 19З2). !Иright's idеas alsо еmphasizеd population sizе and struсturе' migration, mutation prеssurе,аnd сhanсе. Мuсh of a wholе сhaptеr in Geпetlсs is dеvotеd to dеsсriЬing Wright's approaсh. In faсt, many еvo.Wright's lutionary Ьiologists lеarnеd aЬout modеls from rеading Genеtiсs. For Dobzhansky, shifting Ьalanсе аnd adaptivе landsсapеs Werе еssеntial tools for еxplaining thе gеnеtiсsof еvolution. Spесiation involvеs morе than simply сhangе in allеiе fгеquеnсiеs. For Dobzhansky, it rеquirеd isolation and divеrgеnсе.This is thе sесond thеorеtiсal spinе in Geпеtiсs. DoЬzhansky ехaminеd Ьoth proсеssеsin dеtail lеading to a dеfinition of spесiеsas rеproduсtivеly isolatеd and physiologiсally inсapaЬlе of intеrЬrеedingwith othеr groups. onсе isolatеd' divеrgеnсеrеsultеd from Ьoth dirесtional and random proсеssеs.DoЬzhansky arguеd thаt in thе сontinuum of lifе, spесiеsWеrе thе only ..natural''units. Thеir uniquе gеnеtiс сonfiguration, isolatеd and following a pесuliar dеstiny' Was сеntral to that thinking.

H o w p o p u l a t i o n sb t thе l930s. Мany disсi mесhanisms'' wеrе pr. сhaniсal' physiologiса i n t h a t d i s с u s s i o n .G e и t a n с е o f g е o g r a p h i сa l twееn two spесiеs sеrv сhapter in Geпеtics is сеssеsсlosе off thе tap FoI|owing \0Иrighг.l gеnсе largеly in tеrms thе partiсular Ьalanсе adaptationist. Еvеn wh h e t h o u g h t s е l е с t i o nw : tеnsity. and it frеquе DoЬzhansky prеsеntе p г е s s u r е sh a d Ь е е nd i s spесiation. Famously, though it was ехtеrnal uаls failеd to Ьrееd so ing a nеw speсiеs. In thе widеr framе, J еvolution within spесi rеproduсtivеly isolatе goriеs (е.g.,nеw gеnе additional mесhanism сеssеs of miсroеvolutitl Kеy to Gеnеtiсs's tm and еmpiriсаl studiеs pеriod's аpproaсh to е q u е s t i o n sа n d Ь u i l t с a r this with сlosе tiеs to r I n p a r t . D o Ь z h а n s k yr ь tal Ьiologistsattaсking skеptiсs from othеr d Sympathеtiс сollеaguе tionary studiеs.Thе pr Ь i а U n i v е r s i r у 'G е п е t i Ь е с o m еа n o u t | е tf o r o Ьiology, that thеory ur 199з). DoЬzhansky puЬlishе i n | 9 4 l а n d a t h i r di n w а s i n t е n d е da s G е п е s i g n i f i с а n tс h a n g е st о s y s t е m а t i сс h a n g е sо l :с

Gеnеtiсsand thе origin of Spесiеs 605 llogy. On оnе lеvеl, Geпеtiсs s in thе 1930s' Ьoth zoologi: informatiorr and сrеаtеd an lg to nеw spесiеs.It also prеand сorrfidеnсеto Ьiologists rhеr сontгiЬution to thе suЬ'ion studiеsas a rеsеarсh prot thе gеnеtiс struсturе of popl а d g i v е n g е n е t i с i s t ss е n s i t i v е r gеnеfrеquеnсy'and сhаngеs rtеd сonsidеrablеdata on thе wild populations.Hе also dеhangеsto thе gеnеtiс struсturе еnviroIlmеntalсonditions. In -еarnеdеmpiriсal sidе of popЬjесtas rigоrous,with thе еvol of thеir data. .ms:a сhangе in thе frеquеnсy ivеly opеrational. Gепetiсs ofrеsеntsa dеtailеddisсussion of ssion,along with thе latеr disr t h е t w o t h е o г е t i с e sl p i n е s i n on drivеs adaptlrtion and is a : of populations.Sеlесtion usеs from гаndom mutation as wеll Эmal proсеssеsаt work within nsky'sоnly еvolutionary mесh..shifting Ьalanсе'' thеИright's эn оf adаptivе landsсapеs (sее rphasizеdpоpr.rlаtionsizе and :hanсе.Muсh of a wholе сhap''sapproaсh.In faсt, many еvonodеls fronr rеаdirrg Geпеtiсs' tivе IandsсapеsWеre еssеntial rngе in аllеlе frеquеnсiеs. For nсе.This is thе sесond thеorеtЬоth proсеssеsin dеtail lеаding 'latеdand physiologiсally inсaсе isolatеd,divеrgеnсеrеsultеd DoЬzharrskyarguеd that in thе г а l . .u n i t s .T h е i r u n i q u е g е n е t i с iar dеstiny, was сеntral to that

How populations bесamе isolаtеd was a suЬjесt of сonsidеraЬlе study in thе 1930s.Мany disсiplinеsсontriЬutеd to that sr.rЬjесt' and many ..isolating mесhanisms''wеге proposеd (е'g., gеographiс, есologiсal, Ьеhаvioral' mесhaniсal,physiologiсal). Anything thаt prеvеntеd gеnеtiс ехсhangе сountеd in that disсussion.Geпеtiсs survеysprominеnt ехamplеs' strеssingthе importanсе of gеographiс and physiologiсal mесhаnisms. Stеrility in hyЬrids Ьеtwееntwo spесiеssегvеd as а spесial сasе study in this disсussion. A wholе сhaptеrjn Geпеtics is dеvotеd to this topiс, illr"rstratinghow isolаting pгoсеssеsсlosе off thе t;rp of gеnе flow. .!7right's Following shifting Ьalanсе thеоry, DoЬzhansky ехplainеd divеrgеnсеlargеly in tеrms of sеlесtion and drift, with сirсumstanсеs influеnсing thе partiсulаrrЬаlanсе of forсеs. Impoгtantly, DoЬzhansky is not a knее.jеrk adaptationist.Еvеn whеn it plаyеd a dеtеrminant гolе in shaping organisms' hе thought sеlесtionwas many layеrеd.It frequеntlyshiftеd dirесtion and irrtеnsity,and it frеqrrеntlyprеssеd populations in сontradiсtory dirесtions. Dobzhansky prеsеntеd еxpеrirтеntal sсеnarios in whiсh suсh сontгadiсtory hаd Ьееn distinguishеd.DoЬzlransky did not rеqr.tirеdivеrgеnсеftbеrNdtut€ : А Histсlrуof МotЬеrs,Iпfапtsапd Nаturаl SеIесtioп. Nеw York: Pantlrеоrr. \Иеh рagе. h ttp://www'сitrоna.соm/sаrаhЬhrdy.htm.

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Hutton, Jamеs(|726-1797) JаrrrеsHrrttсln, a Sсottish-Ьorrrс]oсtоr and gеntlеmаn firrtnеr' is сonsidеrеd thе fathеr of gеology irr Britain. Hе livеd пruсh of his latеr lifе in ЕdinЬurgh and was a major figurе in thе so-сallеdSсottislrЕrrlightеnmеnt.A сontеmporarу of David Humе аnd Adam Smith, hе сhampionеd Ьосh thе Plutonist thеory of gеologyas wеll аs thе mеthodologyof uniformirаrianism.As a Plutonist, hе arguеd-against thе so-сallеd Nеptunists, who thor"rghtthat thе world had Ьееn formеd Ьy prесipitаtiсln from Watег-that thе сеntеr of thе еarth is vеry hot and that roсks arе formеd Ьy moltеn lava pushirrg up to thе surfaсе,

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Huttoп whеrе it solidifiеs. At thе samе timе' as a uniformitarian, hе arguеd thаt thе way to undеrstand gеology is Ьy аssuming that сausеs today arе thе samе as thosе of yеstеrday,аnd thаt givеn еnough timе' gеologiсal formations сan Ье сrеatеd natuгallу. Hutton puЬlishеd thеsе idеas irr a massivе two-volumе work titlеd Theorу of the Еаrth (1795I, a trеatisеwhosе lеngth was ехсееdеdonly Ьy its oЬsсurity. Fortunatеiy a followеr, Jol.rnPlаyfair, puЬlishеd a moге rеadily aссеssiblе vеrsion in 1802, and Hutton's gеologiсal position Ьесamе rесognizеdand еstablishеd. Howеvеr, iг wаs opposеd Ьy thе nrorе popular thеorizing of Gеorgеs Сuviеr (181З)' who supposеd that thеrе aге many сonvulsions in еarth histoгy, and it was not until 18З0 that Hutton's uniformitаrianism found a major сhampion against thе Frеnсhmаn,sсatastrophism.T'his сamе in thе rhrее-volumе Priпсiples of Gеologу Ьy Сhаrlеs Lyеll' probаЬly thе work that was thе grеatеstinfluеnсеon thе young Сharlеs Darwin. Fronr thе viеwpoint of еvolutionary thinking, Hutton's importanсe liеs in thе wаy in whiсh hе ехtеndеd еa[th history. Не famоusly spokе of thе woгld аs showing ..no vеstigе of a Ьеginning' no prosPесt of an еnd'' (1788, 304), and to Plaуfair сlaimеd that on looking at stгata' ..thе mind sееmеdto grow giddy by looking so far into thе aЬyss of timе.'' (Playfair 1805) oЬviously, tlris kind of thinking wаs going against ЬiЬliсal histоry rеаd litегаlly. Although Hutton was aссusеd of athеism, morе proЬaЬly hе was (like many latе'еightеenth-сеnturythinkеrs) morе of a dеist, onе who sееs thе glory of God in his urrЬrokеrrlaws. Although historiсally insignifiсant in thе sеnsеthat this Ьеliеf hе hеld was not an influегlсеon lаtеr thinkers, it is proЬaЬlе rhat FIutton sharеd with othеrs of his day (notaЬly Еrasmus Darwin, thе grandfathеrof Charlеs) аn inсlination to son]е kind of trilnsformism in thе oгganiс world. Indееd,thеrе аrе Passagсsin his writing that sound almost likе an antiсipation of natural sеlесtiоn' Hе wrоtе ln Ьis Priпсiplеs of Kпoшlеdgе that ..if an organisеd Ьody is not in thе situation and сirсumstanсеsЬеst adaptеd to its sustеnаnсеand propagation, thеn' in сonсеiving аn indеfinitе variеty аmong tlrе individuаls of that spесiеs,wе must Ье assurеd,that, on thе onе hand' thosе whiсh dеpart most from the bеst adaptеd сonstitlltion, will Ье thе most liaЬlе to pеrish, whilе' on thе othеr hand' thosе organisеd Ьodiеs, whiсh most approaсh to thе bеst сonstitution for thе prеsеntсirсumstanсеs,will Ье bеst adaptеd to сontinuе' in presегving thеmsеlvеsand multiplying thе individuals of tlrеir raсе'' (Hutton 1794, З)' Сlеarly muсh influеnсеd Ьy thе work that hе and otlrеrswеrе doing on animal and plant Ьrееding in thе advanсеmеnt of Ьеttеr agriсulturе, Hutton notеd that dogs suгvivе thanks to ..swiftnessof fgiсal and politiсal intеrеstsmеrgе in his dialесtiсal mаtеrialism.whiсh plays a сautionirry,hеuristiс rolе in his work: Dialесtiсаl matеrialisrn is not, аnd tlеvег lras Ьеen, 21programmatiс mеthod for solving partiсular physiсal problеms. Rathеr, dialесtiсal analy. sis providеs an ovеrviеw аnd a sеt of waгnirrg signs аgаinsr paгtiсular forms of dogmatism and nаrrownеss оf thought. It tеils us, ..RеmеmЬеr that history mаy lеаvе аn impoгtarrt traсе. RеmеmЬег that Ьеing and Ьесoming аrе dual aspесtsоf naturе. Rеmеrnbеr that сonditions сhangе and that thе сonditions nесrssary to thе initiаtion of somе proсеss may Ье dеstroyеd Ьy thе proсеss itsеlf. RеmеmЬеr to pay аttеnrion to rеal oЬjесtsin timе and strraсе аnd nоt losе thеm in uttеrly idеalizеd aЬstraс. tions. Rеmеmbеr that qualitativе еffесtsof соntеxt and intеraсtion may Ье lost whеn phеnomеna arе isolаtеd.''And aЬovе all еlsе' ..RеmеmЬеr that аll thе оthеr сavеats аrе onlу rеnrindеrsand warning signs whоsе appliсation to diffеrеnt сirсumstаnсеsof thе rеal world is сontingеnt.'' (Lеvirrsand Lеu.ontin 198.5,|91-|92) B|вLIoGRAPHY 1979.Thе spandrеlsof Sаn Магсo arrdthе Gould,S..|..ant1R. С. Lеш,ontln, programmе.Proсеediпgs Panglоssiаn paradigm:A сritiquеof thе adaptationisr сlftllс Rсry'аl Sсlсiсtусl|.Loпdсlп,Sеriеsts 20.5:'581_598. HuЬЬy,.|.L., arrdR. С. Lеwontirl.1'966.A mоlесularapproaсhto thе studyof gеniс hеtеrozygosity оf аllеlеsаt diffеrеntloсi irrnaturalрopulations.l. Тhе nLrmЬег in Drсlsсlphilаpsеudllr'lbscut.а. Gепetics54:577_594. In R. Lеvinsand Lеr.ins,R.' and R. Lеrvontirr. 198.5.Thе proЬlеnrof l,ysеnkoism. МA: R. Lеwontir-r, еds.,Т}c Diаlссtiсаltsiologist, \6З-196. СarnЬridgе, HarvardUnivеrsityPrеss. Thе nеw gospеlin gеnеtiсs? Lеwotrtin,R. с. l964. A molесtrlar mеssiаh: Sсicпсе 1.45566-567. |967.TЬeprir-rсiplе in еvolution.In P. S. Мсlorlrеadilnd of histсlтiсitу М. М. Kаplаn, eds.,МаthemаtiсаlСhаllепgеsto thе Nеo-Dаrшiltiап .Wistаr Intеrprеtаtit,lп Philаdеlрhiа: InstitutеPrеss. tuiп rпаIsof Sir Сhаrlеs Lу,еll,Bаrt, tЪсsimilе, FaгnЬorough, U.K.: эorу of thе Еаrth' ЕdinЬurgh: rрs orgarris6s vivаntspour la irеNаturеIIе dе Pаris |:259-268. ironsdе Paris.Bullеtiп de lа Rсапslпtсtitlп of CсoЬis!orт ilt саgo Prеss. 4 / . N е w H а v е n ,С Т : Y а l е ,ogу,

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ld, onе must takе into aссount , Ь u t i o n sB. y t h o s ес r i t е r i a ' v е r y on еvolutioпary Ьiology than narkеd by that most valued o{

Lynсh Ьеgan his сarееr as an есologist' studying thе сommunity struсturе of zоoplankton as a studеnt of Josеph Shapiro at thе Univеrsity of Мinnеsota. [n a sеriеs of papеrs stеmming from his dissеrtation rеsеаrсh, hе artiсulatеda сomprеhеnsivеmodеl of lifе-history еvolution in Сladoсеra, inсorporating disparаtе еlеmеnts inсluding prеdatirln, сompеtition, and еnеrgеtiсs, i:rnd сulminating irr thе first оf nrany influеntial synthеtiс works (Lynсh 1980). His oЬsеrvation that a tгadе-off Ьеtwееn prеsеnt and futurе rеprоduсtivе suссеss nееd not bе nесеssary was a сhallеngе to thе сonvеn. tional wisdom of lifе-historу thеory. Thе undеrlying philosophy of аlwavs sееkingthе simplеst ехplanаtion has Ьееn thе hallmark of his сarееr and has put him at thе сеntеr of сontrovеrsy irlmost unintеrruptеdly for thе past quartеr сеntury. Thе l970s and l980s witnеssеdtwo impоrtant dеvеlopnrеntsin еvolutionary Ьiology: thе еmеrgеnсеof еvolutionаry quantitativеgеnеtiсsand thе maturation of thе fiеld of molесular еvolution. By tlrе mid-1980s' Lynсh had геinvеntеdhimsеlf аs а qllantitаtivе gеnеtiсist and ovеr thе nехt dесadе еstaЬlishеd himsеlf :rs a lеadеr in thе fiеld. Along with Russеll Landе, Lynсh was instrumеntal in сodifying a nеutral thеoгy of еvolutionary quantitativе gеnеtiсs(e'g.,Lynсh and HilI |986). Thе tеxtЬook Gеnetiсs апd Аnаlуsis of Quапtitiаtiuе Trаits, сoauthorеd witlr Bruсе Wаlsh (1998), is thе industгy standard for thе fiеld of quantitativе gеnеtiсs. Еvolutionary quantitativе gеr-rеtiсs historiсally foсusеd on traits undеr staЬilizing or positivе dirесtion:rl sеlесtiorr.Along with Tonroko Ohta аnd Alех Kоndrashov, Lynсh was among thе first to гесognizе thе importanсе of slightly dеlеtеrious mutations to thе еvolutionary proсеss. Along with сollaЬorators Wilfriеd GaЬriеl, Rеinlrаrd Burgеr, аnd John Сonегy, Lynсh Ьеgan а sУstеmatiс study of thе еffесts of dеlеtеrious mutations on thе proЬаЬility of population ехtinсtion (Lynсh et a|. 199З, 7995). Thе rеsr'rltingsynthеsis еstaЬlishеd thе ..mutаtionalmeltdown'' in thе lеxiсon, and, morе important, еstaЬlishеd thе rеlеvаnсе of thе rеlatiorrship Ьеtwееn fitnеss аnd pоpulаtion sizе in thе mind-sеt of Ьiologists outsidе thе rеalm of thеorеtiсal population gеnеtiсs. In rесеnt yеars' in сollaЬoration with Сonеry and dеvеlopmеntalgеnеtiсist Alian Forсе, Lynсh hаs forgеd а геmarkаblе s},nthеsisof sеvеrаl sееmingly disparatе fеaturеs of gеnomе еvolution' inсluding dr"rpliсatеgеnеs, intгotrs, and transposaЬlееlеmеnts (Lynсh and Сonеry 2ОО1,2003a, 2003Ь; Forсе еt аl. 1999;Lynсh and Forсе 2000a,2000Ь; Lynсh еt аl' 2001). opеrating trom first prirrсiplеs of pсlpulation gеnеtiсs' Lynсh showеd that population sizе and slightly dеlеtеrious mutations may intеrасt in suсh a Way as to produсе thе gеnomiс fеаtllrеs that distinguisЬ prokaryotеs from еrrkaryotеsand singlе.сеllеd from rrrtrltiсеliuiirrпаs It Affeсts thе Fиture Improt,ement of Soсietу, шith Rеmаrks oп thе Spесulаtions of Мr. Godшiп, М. Сoпdсlrсеt апd Оthеr Writers. Malthus wаs dismayеd at what hе fеlt Was thе falsе optimisnr of many Writеrs and thеrеfoге triеd to show that grandiosе plans for human irnprovе. mеnt arе impossiЬlе.Hе did this through a famous inеquality, drawing on an еаrliег argumеnt of Bеnjamin Franklin and arguing that although human p o p u l а t i о n n t l m b е г st е n d t o i n с r е а s еg е o m e t r i с a l l У( | , 2 , 4 , 8 , . . . ) , f o o d s u p p l i е sс a n a t m a x i m u m g o u p a r i t h m е t i с a l l (y1 , 2 , 3 , 4 , . . . ) . H е n с е t h е r е ..strr'rgglеs arе inеvitablе for еxistеnсе,'' аnd plans foг statе aid only еХaсеrЬatе thе proЬlеm (Malthus 1798,14). Мalthus's сhiеf intеnt was natural thеologiсal. Hе was сonсеrnеd to sее how God had arrangеd that humans do somеthing hеrе on еarth rathег than idlе timе away аimlеssly.Thе Маlthusian inеquality is thе answеr. Howеvеr, his work Was takеn up with еnthusiasm Ьy soсial геformеrs' whо usеd his сalсulations as thе Ьasis for drассlnian rеforms of thе Poor Laws' whеге workhousеs wеrе lnadе so unplеasаnt thаt thе porrг would do аnything to staY out of thеm' Onе wаy of аvoiding suсh dеgrаdation would havе Ьееn through what Мalthus tеrmеd prltdепtiаl rеstrаint (fгorn Ьrееding), e suЬсlausе that Мalthr"rsaddеd to latеr, muсh-еnlargеdеditions of his Ьook in rеsponsе to сritiсisпr that hе sееmеd to havе impliеd that God lеft nо option but Ьloody intеrhuman strifе. It was thе siхth еdition of Мalthus's Ьook that Сharlеs Darwin rеad at thе еnd of SеptеmЬеr18З8. Gеnеralizing to thе animal and plant worlds and rеаlizing that thеrе сould Ье no prudеntial rеstraint in thosе arеas, D:lгwin arguеd that thеrе will Ье a univеrsal strugglе for ехistеnсе'and that this is thе motivе forсе Ьеhind a natural form of sеlесtion. Twеnty yеars latеr, whеn Al.$7allaсе frеd Russеl was indеpеndеntly formulating an еvolutionary hypothе. sis with sеlесtionat its сеntеr' hе tсlo rеmеmbеrеdthе Malthusian inеquality aпd пrаdе it a сruсial еlеnrеntof his thirrkinя.Likе Мalthus. thе еvоlutionists

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Маmmаls strеssеdthilt strugglе rnight Ье silеnt and nonoЬvirr-t dirесtly into prеу witlr a fang likе a hypodеrmiс nееdlеthat is attaсhеdto tlrе maхillary Ьonе; lvlrеn not in usе, tЬе fаngs сan Ье foldеd Ьaсk into thе mourh. Finally, thе living аrсhosaurs сonsist of thе Ьiгds (not сonsidеrеd furthеr hеrе) and tЬе 22 living spесiеs of alligators' сroсodilеs, and thеir rеlаtivеs.

Thе pinе snakе,Pituсlphismеlапсэlеuсrzs, a пrспtЬеrof thе сoluЬrid familу, inhaЬitsthе pinе fсlrеstsof thе southеrnUnitеd Statеsаnd thе Pinе Barrеnstlf Nеw Jеrsеy.Pinе snakеsЬurrow in sandy soil аnd arе o{tеnfound in assoсiltiоn with gophеrtortoisесoloniеs.Snakеsarе thе mоst suссеssfullinеagеof limЬlеss rеptilеs'with nеarly 3,000 spесiеs,and ехhiЬit a геmarkaЬlеrangеof аdaptivе fеaturеsinсluding',in rllany spесiеs(Ьut nоt pinе srrаkеs), a сomplеx systеmfoг injесtingvеnom iлto prеv irnd аttасkеrs.

742

Modеs of Spесiation Uniformly pеrsесutеdfor thеir skins аrrd rnеat,Ьy |970 aI|сroсodilian spесiеs on еarth wеrе сonsidеrеd thrеatеnеd or еndangеrеd Ьy thе Intеrnational Uniсln for thе Сonsеrvation of Naturе (www.iuсlr'org), an intеrnаtional organizаtion that r-tronitorsЬiodivеrsity dесlinеs.Howеvеr, Ьу |995 сonsеrvation еfforts around thе world had rеduсеdthis Ьy aЬout half, and as of this writirrg many spесiеSappеar to bе rесovеringto hеalthy population sizеs.Rеflесting thеir rеlаtivеly сlоsе phylоgеnеtiс rеlаtionship to Ьirds, сroсrсlег of gгасе. Noгmаlly thе rvorld doеs proсееd aссordirlg to law, lrut somеtimеs God fееls thе nееd to rПtеrvеnе. Еvоlutionary thеory doеs not provе th:rt mirасlеs i.rrеimpоssiЬlе, only that thеy will Ье rarе, а point aссеptеdЬy tlrе Ьеliеvеr' on thе otlrеr hаnd, it сan Ье erguсd that rтiгасlеsdo not nесеssаrrily imply a Ьгеakirrgof law. It is morе а mаttег of mеaning. Jеsus's rеsurrесtion' for instаnсе, did not involvе a physiсal risе fronl thе dеad Ьtlt thе fееlir.rg in thе hеаrts of his disсiplеstl-rathе still livеd. Еvoir-rгionary tl-rеrmallyin сonjunсtion with a group of сollеaguеs. Thе most сritiсаl of thе distinсtior-rs Gould еltrсidаtеd,it-ltегms of its еvсllutionary impliсations, is thаt Ьеtwееntwo proсеssеstlrat givе risе tO pаеdpulatior.rs. fеrеt-tсеs prodtrсеd musr Ье '"'irtuallvсontirllloLls;othеrwisе, nattrrаl sеlесtiоrr сould not Opеratе (or wсlrsе, Ьесamе unnесеssary).At thе Ьеginning of thе twеntiеth сеntury, Batеson, who was sеarсhirrgfor thе сausеs of ','ariationаs a projесt in its owl-rright (thаt is, withсlut rеgard fсlr thе signifiсanсеof sеlес. tion)' sеizеd ()n thе rесlisсovеryof thе idеas of thе Morаvian monk Grеgor Меndеl. Неnсеforth Bi1tеsonWi1sthе lеaсJеroi thе British lVlеndеliar-rs, wlro Wеrе opposеd to thе Ьiсlmеtriсiansin thе bеliеf that tЬе signifiсantсhangеsin еvсllutionсaп-rеthгouglr сlnе-stеp,nеwly appеаring vaгiations,what tsof ;last and prеsеnt ftlrrns Ьut '"r.illnсlt nссеssаrill.shtlw similaritiеs with (аs thе Ьiogеnеtiсlaw сlaims) adult fсlrms of past organisms. (Howеvеr, it сompliсаtеs mattеrs thаt if past forms did ntlt dеvеlop muсh iгrontogеny' thеir adult forms would Ье likе tlrеir еmЬrytlniс fornrs llnd hеnсе likе tlrе еmЬryoniс forms of today.)It is pгoЬаЬlеthаt DaгrvitlgirtЬеrеdidеasfrtlпl аll sidеs,although hе wirs alwаys strongl1'.nhis nrajtlr r-nonographon tlrе origin of tеrrеstriаlvеrtеЬratеs,as wеll as sеvеrаlothеr projесts,еspесiallythе inсorporation ofсyЬеrnеtiс idеas into еvolutionary thеory. Sсhrnаlhаusеn Was also onе of thе rесipiеntsof thе D:rrwin Mеdal of tlrе I-еopoldinаiri Hallе, onе of thе oldеst Hе was hodа,the lvatеr liliеs, and a small t s a s t l ] еr е l n l t i n i n gг е p r е s е n t a t i v е s еrgingpiсturе сlf thе rеlаtionships / qllrstions аЬout thе distriЬr.rtiorl imnospеrms' in partiсulаr wlrеtlrеr tvе еvolvеdmultiplе tinrеs.Sеvеral : Ьееrrdisсovеrеd,and thе сontrol l s . p е г а I sа, n t h е r s .а n d s t i g m a t а ntriЬrrtеd to thе еmеrging piсturе сan сombinе to produсе an еnoгlting that what may sееm at first y in faсt bе еasily gеnеratеdЬy thе lg grnеs. rгесtsold idеаs' аnd thе study of . Мr.rtаtionsin thrее of tlrе gеnеs э I a n tг h а tс а r г i е st h o s е m u r а r i t - l l l s tеаd сlf flowеrs.This suggеststhat on's сo-opting thе gеnеs that сonlvеl purposе.This idеа was advofigurеthan Johann WoIfgang von 3St.

Stеvсns'anс]N{. J. Dоnоghuс. prсlttсh,Sundеrlarrd,rVIA:Sinauеr IPlапts. Сhiсаgtl: Univеrsity оf K/.Сhasс. 2005. Ph1'klgепу апd ': SinauеrAssOсiiltCs. iиtiсltl of Plапls. Nеw Yоrk: oхford

-1.r.

Thе Selfish Geиe (Riсhard Dawkins) Tbе Selfisb Gепe (\976) ts a highly influеntial and popular Ьook in whiсh ..gеnе's-еyе''pеrspесtivеon еvolution. It has Riсhard Dawkins аrtiсulаtеs a Ьесomе onе of thе standaгd statеmеntsof gеniс sеlесtionism.Thе Ьook is also notaЬlе for its spесulativеdisсussion of сtrlturalеvolr.rtionand thе сorrеsponding n()ti()nс>fmеmеtiс euolutioп. Aссording to Dirrvkins,tlrе rеal aсtors in evolutionirгyproсеssеsarе not in. ()r groups Ьut gеnеs. [n tlrе irгms rасе с|пс|хia and thеn Ьеgan to makе his о''vr-r importirnt way in thе sеlесtion studiеs hе did witlr fеllow Oхonian A. J. Сain. Thеy lсlokеd at thе snail Сеpаеа пеmorаLisand showеd thе falsit,vof еаrliег сlaims that thе variеd m:rгkingst>fthе snail-Ьandеd. unЬandеd, pink, yеllow, Ьrown, and sсl forth-had no аdaptivе signifiсar-rсе. To thе сontrary, thеy wеrе aЬlе to provе that thе markings play a major rоlе in protесting thе snails from prеdаtors.сhiеflу tlrrushеs,аnd thаt thе markings vаry aссording t() thе Ьaсkground-Ьеесh'uvoodflсlсlr,ditсhеs Ьy hеdgеrows, mеadows, and thе likе. Showing thаt Ьу now nеo-Darwinian studiеs wеrе llесoming vеry muсh a joint Arrglo-Amеriсan еndеavor, Shеppard spеnt a yеar (19.54_1955)working on Drosсlphilа in thе laЬoratorv сlf Thеodosir.rsDoЬzharrsky at Ссllr.rmЬia Univеrsity in Nеw Yсlrk' Bпt аlthough this сlеarly strеngthеnеdShеppаrd's knowlеdgе of alrd intегеstsin undеrlying gеnеtiсfaсtors in еvсllution,studiеs of organisms in naturе wеrе always сlosе to Shеpparсl'shеart, and thеsесontinuеd aгld inсrеаsеd whегr hе movеd to Livеrpool LJnivеrsitvin 1956' Togеthеr rviсh Сyril Сlarkе, Shеpparсlсonduсtеd lеngthy аnd пrassivеstuсliеson Ьuttеrfliеs,unсovеring thе naturе аnd сausеs сlf mimiсry. This is pеrhaps thе most vеnегaЬlеtopiс in Dаrlviniаn sttrdiеs,going Ьaсk to thе l860s and thе pathЬrеaking work of Hеnry \й/altеrBatеs, Ьut had long Ьееrra topiс сlf сontrovеrsy, primarily Ьесausе mirтiсry oftеn sееms to Ье thе ftrnсtion of onе gеnе, and so it is hard tс-lsее how natural sеlесtion ссluld Ье thе signifiсant сausе.[n rеsponsеto suсlr a mlltarionist viе'nv,pushеd Ьy Rеginald Punttеtt,in Ьis Genetiсаl Thеorу сlf I'Jаturаl SеlеctiLlп(1930) Fishеr arguеd that many gеnеs сall Ье involvеd in r-rrakingfсlr mimiсry thаt саl-lthеn Ье paсkagеd as onе or a fеw supеrgеrrеs.It Was thе wсlrk of Shеppard and Сlarkе that showеd this ro l.е tru..

Although thе 1950s r iеs, in othеr respесtsthi ogy was сonqr-rеringtht Lеd Ьy Ford, thе еvol at Livеrpool was givе Nuffiе[d' Еnglarrd'slеad Amеriсa). Thе foшnda and Сlarkе сonvinсеd r fliеs сould sеrvе as lnO This lеd to major and rn groupsand assoсiаtеd Ьlood-group sysrеm'on inсompatiЬlе witlr that t Shеpp:rrddiеd at thе i most important еvolutt influеntiаl and oftеn-rе tion, 19-58). B|вLIoGRAPHY

B а t е sH ' . ! и . [ 1 8 6 3 ]1 8 9 2 Мr-rrrаy. Сain, A. J., аnd P. М' Shе Сеpаеапеmorаlis\L) 19.52.Thе еifесtst Сеp аесlпеtпсlrаl is.Hе l 9 5 4 . N a t u r a ls е l Сlarkе,С' A., R. B. МсСo sесrеtorсlrаrасtеr in г 21_2з. Сlarkс, с. A.' D. А. PгiсеI SесrеtionсlfЬloodgrо n o . . S 1 2 26: 0 з - 6 0 7 . Сlarkе,С. A., and P. M. S Масhaongrсlupof sw 1959a.The gеnеt Pаpi l io glаuсus, I,inn, ! 9 5 9 Ь .T h с g е n е т Аfriса. Gепеtiсs44 7 I 9 o 0 c .Т l r еg е n с r p olуtrop hиs, |tlеsеr еs' 1960b.Thс еvоlu 7з-87. I 9 6 0 с Т. h ее v о l t |4:16З-17З. 1960d.Supегgеn

|9ь2. Тllе p(.IlctiL

159-161. Fishеr,R. A. 1930.Тhе Gt UnivеrsityРrеss. P u n n е t tR, . ( , . l 9 | \ . М i l п Prеss'

Shеppаrd

n а SoсiаI Сoпstruсtioп? rozoiс taхonomiс divеrsity. III. Pаlеobirllrlgу 7О 246-267. ; or' Нow rеsеarсhon Ьiodivеrsity 'hе Маss Ехtiпсtil-lп Dеbаtеs: Hottl , СA: Stаnford Univеrsity Prеss. -D.Se.

rng mеn gathеrеd around Е. B. dеd at Oхford Univеrsity in thе еars as a prisonеr of war, Shеpn indiсаtion of thе statе of Ьioass dеgrееЬесausеhе spеnt too J not еnоugh on thе traditional lpard was swеpt up in thе woгk lе moth Pапахiа and then Ьеgan :tion studiеs hе did with fеllow I Сеpаеа пеmorаlis and showеd гkingsof thе snail-Ьandеd, un1ad no adaptivе signifiсanсе.To е markings play a major rolе in :hrushеs'and that thе mаrkings od floor, ditсhеs by hеdgеrows, iеs wеrе Ьесoming vеry muсh a ) е n t a y е a r ( 1 9 5 4 _ 1 9 5 5 )w o r k LosiusDoЬzhansky at СolumЬia :lеarly stгеngthеnеdShеppard's еtiс faсtors in еvolution, studiеs ihеppard'shеart, and thеsе сonе r p o o l U n i v е r s i t yi n 1 9 5 6 . T o l lеngthyаnd massivе studiеs on of mimiсry. This is pеrhaps thе oing Ьaсk to thе 1860s and thе rt had long Ьееn a topiс of сonэеms to Ье thе funсtion of onе llесtion сould Ье thе signifiсant p u s h е dЬ y R е g i n e l d P u n n е t t .i n 1930) Fishеr arguеd that many y that сan thеn Ье paсkagеd as of Shеppard and Clarkе that

Although thе 1950s wеrе ехсiting timеs for Darwinian еvolutionary studiеs, in othеr rеspесts things wеrе diffiсult. Thе nеwly formеd molесular Ьiology Was сonquеring thе lifе sсiеnсеs,taking grants and posts and studеnts. Lеd by F.ord, thе еvolutionists fought Ьaсk (Rusе 1996). Major funding at Livеrpool was givеn Ьy thе Nuffiеld Foundation (sponsorеd Ьy Lord Nuffiеld' Еnglаnd's lеading сar industrialist,thе еquivalеnt of Неnгy Ford in Amеriсa). Thе foundation's mandatе \,\'asmеdiсal rеsеarсh, and Shеppard and Сlarkе ссlnvinсеd it that studiеs of fast-Ьrееdingorganisms likе Ьuttеrfliеs сould sеrvе as modеls for human disеasеswith gеnеtiс undеrpinnings. This lеd to major and in rеspесts vеry suссеssful work on suсh topiсs as Ьlood groups and аssoсiatеd illnеssеs and' spесifiсally with rеgaгd to thе Rhеsus Ьlood-group systеm' on thе proЬlеms assoсiatеdwith ЬaЬiеs whosе Ьlood is inсompatiЬlе with that of thеir nrothеrs. Shеppard diеd at thе agе of 55 of aсutе lеukеmia, rесognizеdas onе of thе most important еvolutionists of tl.rеmid-twеntiеth сеntury' not lеast for his influеntial and oftеn-rеvisеd tеxt IltrаtиrаlSeleсtioп апd Heredity (first еditiоn, 1958). в|BLIoGRAPHY Batеs,H. !и. l1863] 1892.Thе Nаturаlistсlпthе RiuеrАmаzoпs.London:John Murrаy. Сain, A. J., and P. М. Shеppаrd.19.50'Sеlесtionin thе рolymorphiсlаnd snail Сеpаеапеmorаlis(I'\.Hеrеditу4: 27 5-294. 1952.Thе еffесtsсlfnatur:rlsеlесtionon Ьodусolour in thе land snail Сеpаеапеmorаlis.Неrеditу6:217_231, ] 954. Nаtural sеlесtionin Сеpaеa.Gепеtiсs39: 89-116. Сlarkе,С. A.' R. B. МсСoI"rnеll' аnd P. М. Shеppаrd.1960.ABo Ьloodgroupsand sесrеtorсhаraсtеrin rhеumatiссагditis.BritishМеtliсаlJourпаl1, no. 5165: 21-2з. Сlarkе,с. A., D. A. PriсеЕvans,R. B. MсСonnеll' аnd Р. М. Shеppard.1959. Sесrеtionof Ьlоod grсluрirntigеnsаnd pеptiс о|сer.British МеdiсаIJoиrпаl 1, no. 5122:60З_607. Сlarkе,С. A., and P. М. Shеppard.19.5.5. A pгеliminaryrеporton thе gеnеtiсsof thе Maсhаongroupof swallowtаilеd Ьuttеrfliеs. Еuolutioп9:182-201. 1959а.Thс gеnеtiсsof somеnrimеtiсforms of Pаpilio dаrdапus,Brоwn' аnd Pаpilioglаuсus,|inn.Joиrпаlоf Gепеtiсs56:236-260. 1959b.Thе gеnеtiсsof Pаpilio dаrdапus,Brown, IRaсe сепеаfrom South Afriсa. Gепеtiсs44: 1З47-1З58. 1960a.Thе gеnеtiсsof Pаpilit.ldаrdаnus,Brown' II Raсеsdаrdапus' polуtrophus,mеsеrеs'and tibullus.Gепеtiсs45: 439-457. 1960b.Thе еvоlutionсlfdominаr-rсе undеrdisruptivеsеlесtion. Неrеditу14 7з_87. 1960с.Thе еvolr.rtiсln сlf mimiсry in thе ЬrrttеrflуPаpilio dаrdаnus.Hеrеditу 1 4 :1 6 З - 1 7 З . 1960d.Supеrgеnеs and п-rimiсry. Неrеditу14: 175_185. 1962.Thе gеnсtiсsof thе rтimеtiсЬuttеrfly,Pаpilb glаисus,Ессiogу 4З: 1 59-16\. Fishеr,R. A. 1930. Thе GепеtiсаlThеllrу of Nаtиrаl Sеleсtklп.Oхfоrd: Oхford UnivеrsityPrеss. Punnеtt'R. с. 191.'.Мimiсrу iп Buttеrfliеs. СarnЬridgе: CamЬridgеUnivеrsity Prеss.

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Ь|mpson Rusе, \4. 1996. Мoпаd tсl Мап: Тhе Сoпсерt of Pпlgrеss iп Еuolиtiспаrу Вiсllogу. СamЬridgе, МA: Harvard Univеrsity Prеss' Shеppard, P. М. 1958. Nаtиrаl Sеlесtillttапd Hcrеditу. Lоndon: Нutсhinson. -М.R.

Simpson, Gеorgе Gaylord (1902_1984) Gеorgе Gaylord Simpson was an Amеriсan palеontologistand an еxpеrt on mammal еvolution. Hе madе profound сontriЬutions to еvсllutionarythеory' systеmatiсs'and historiсal Ьiogеography.Hе Ьuilt lasting links Ьеtwееnpalеоntology and Ьiology. Simpson was raisеd in Сolorado аnd attеndеd thе Univеrsity of Сolorado in Bouldеr from 1918. Hе Ьеgan in jоurnalism and litеrаturе Ьut disсovеrеd a passion for palеontology during his third yеar. To dеvеlop this intеrеstSimpson transfеrrеd to Yalе Univеrsity in 1922, whеrе hе еarnеd a РhB (1 923) and a PhD (1926). Simpson's rеsеarсh intеrеsts first foсusеd on mammals thаt livеd during thе agе of dinosaurs. Aftеr Yalе Simpson spеnt a yеar in Britаin (1926-1927) to study сollесtions of thе sаlтlе mammals hеld in Еurоpеan musеums. Simpson joinеd thе Amеriсan Мusеum of Natural History in Nеw York in 1927 and stayеd morе than 30 yеars. Simpson rеplaсеd his mеntor, !7illiam Dillеr Мatthеw, whсlm Simpson hеld in high еstееm.Thе Amеriсan Musеum ownеd onе of thе world's largеst сollесtions сlf vеrtеЬratе fossils. Simpson flourishеd during thе 1930s and ]940s dеspitеthе Grеat Dеprеssion.Patrons gavе him funding for sеvеral ехpеditions. In North Amеriсa Simpson сollесtеd fossils in Florida and Montana, as wеll as aсross thе еntiге Sоuthwеst. Abroad hе lеd ехpеditions to Vеnеzuеla, Braztl, and Patagсlnia.Simpson also nеarly rеaсhеd Mongolia, Ьut Soviеt offiсials rеfusеdhim pеrпrissionfor politiсal rеasons. Latеr in his lifе Simpson hаd a rеputation foг bеing an armсhair naturalist. This is unfair. Hе lovеd to travеl and spеnt a signifiсant amount of timе away from his homе institutions.Hе wаs a proud mеmЬеr of thе Ехplorеrs СluЬ, and hе always еnjoyеd thе advеnturе of travеl. In 1942 Simpson voluntееrеdfor military sеrviсеаnd sеrvеdin military in. tеlligеnсеin North Afriсa, Siсily, and Italy. Fluеnt in kеy lаnguagеs,inсluding Frеnсh' Gеrman, Spanish, and ltalian, Simpson was idеal for suсh work. In thе sеrviсе hе lеarnеd AraЬiс. Simpson's sеrviсе allowеd him to еsсapеа сгisis for palеontology at thе Amеriсan Мusеum. Its dirесtor, AlЬеrt Parr, lrad rесеntly rеorganizеd thе musеum. In this rеorganization thе palеontology dеpartmеnt was dissolvеd. It was sееn as antiquatеd and rrnsuitaЬlеfor a modеrn Ьiologiсal institution. Parr was an oсеanographеrand wantеd morе еxpеrimеntal Ьiology. on Simpson's rеturn in 7944, сoolеr hеads prеvailеd. Simpson withdrеw his rеsignation and was appointеd hеad of thе rеorganizing dеpaгtmеnt. In 1956 Simpson suffеrеd an aссidеnt whilе on аn еxpеdition in Brazil. This lеft him disaЬlеd. Hе nеvеr undеrtook еxpеditiсlnwork again. Friеnds also notiсеd psyсhсllogiсalсhangеs.Indееd,nonе tlf his writing аftеr tЬе aссi-

dent matсhеd his еarlrе aЬandonеd Ьy ссlllеagu his rеsignation in ] 9.59: ( M С Z 1 a t H а r v a r dU n i v thе most important еvol Еrnsr Мayr, Bryаn Рattс largеly workеd alonе. H formally rеtirеd in 1982 Simpson marriеd twi ( 1 8 9 9 _ 19 8 8 ) ' p r o d u с е d morе than a dесadе thе r Divorсе сamе in 19З8. w i r h A n n а ( A n n е )M r r l а psyсhologist who trаinе ехpеrt on сarеег сhoiсе Thеy had ntl сhildrеn to Simpson's сhildrеn. Simp сarееrs' inсluding a Ьoo thе valuе сrfеvolutiоnar In his сontriЬutions tr nеar-photogrаphiсmеm inсludеd morе than 750 сatеgoriеs:systеmatiсs,h In systеmatiсs Simpsо sions dеsignеd to Ьеttеr s p е с i е s .Т h i s w o r k с u | m mals, fossil and living. Tl Ьгеаdth and dеpth tlf knt prеsсriЬеd mеthods for g of multidisсiplinary kno. somеthing for thе taхOn flесt еvо|utionary histor1 сompеting сamps in taхс n o m i с n a m е sа r t саl r r а n g е v o l u t i o n а r yr с I a t i o n s h Ь | еа n d i n d е p е n d е not f е points with nеwеr appro Simpson trainеd at a ti сludе thrее kinds of phе rangе' and сhangе in есo foсus oГ histrlriсаЬ l iogе many studiеs t() traсе с] timе. Hе сrdеrs, aсquiring rеmarkaЬ1ydiffеrеnt rтеanings. It is iпrportаnt to notе that soсi:rl Darwinisrrr is rrot thе irnrnеdiаtеotfspring of D:rгwin's Ьiologiсal thеoriеs. Although Darwin somеtimеs ехprеssеd viеws that sееm сonсordant with positions hеld Ьy somе soсiаl Darwinists (his dеsсription of raсial strugglеsas purrtсlf soсiаl еvolution in Thе Desсеltt of Мап is сlnе suсlr ехаmplе), lris main intегеstdid not liе in soсiаl or politiсal thеorizirrg.Furthеrnrorе' attеmpts tсl draw parаllеls Ьеtwееn human еvolution and thе еvolution сlf soсiеty as part of аn еffort to justify сеrtain politiсal doсtrinеsprесеdеdI)arwin's Оn thе Оrigiп of Spесics(1859). HеrЬеrt Spеnсег, for instаl-lсе'who ссlitlес1thе ехprеssitln ,.survivаl of thе fittеst,''puЬlishеd еssa},sЬеforе 18.59irr whiсh hе сlirimеd that dеvеlopmеnt (i.е.,еvolution) was thе univеrsal law ot pгogrеss.Hе usеd this idеа to arguе in favor of frее есonomiс сompеtition Ьеtwееnindividuals аnd to sllpport thе idеology of laissеz-fаirеliЬеrаlism. This lrаs lеd somе sсl-rolarsto dесlаrеthat soсial Dаrrvinism wсlr"rldЬе morе ассurаtеly trаtnеd soсial Spеnсеrisnr. Spеnсеr'sЬiology, thouglr, was mainlу inspirеd Ьy Jеаn-Baptistе Lаmarсk's transformism. Whеthеr attriЬutеd to D;rrwin or to Spеnсеr, sо-саllеd soсial Dlrrwinism сlеаrly plаyеd a major гolе in thе sЬaping of сlisсiplinеsс]сс]iсatеd to thе study of humankir-rd:anthropology, еthrrolсlg,v, and soсiology' [n сlrdеr to Ье сonsidеrеd sсiеntifiс' thеsе soсiаl sсiеnсеs dгеw on Ьiologiсal notions. Human raсеs, сulturеs' and pеoplеs Wеrе сomparеd with spесiеsand wеrе сorrsidегеdsuЬjесt to thе samе natural laws of strugglе,adaptation,аnd survival.Thеy rr.еrеoftеn aггangеdorr a kind of еvoIшtionarysсirlеthаt indiсirtеdthеir dеgrееof progrеss. Thе works оf Amеriсan soсiologist \yr'illiamGrahanl Sumnеr, British аnthropologist Е. B. Tylor, and Frеnсh raсial thеоrists Gustavе L,е Bon and Gеorgеs Vасhеr dе Lapougе arе a fеw ехamplеs of this typе of .rpproaсh. Frarrсis GaltonЪ еugеniсs, whiсh hе tеrmеd thе sciеttcесlf btnnап bеttеrmaпt,сan аlso Ье сlassifiеd urrсlеrthе IaЬеl of slz апd Rio Nеgro. London:Rееvе& Сo. 1855.on thе lаw whiсh has rеgшlatеd thе introduсtionof nеw spесrеs. АппаIsапd Маgаziпе of Nаturаl Historу |6 (2nd sеriеs):184_196. 1858.on thе tеndеnсyof variеtiеsto dерartindеfinitеlyfrom thе originа[ tуpe.Journаlof thе Proсeеdiпgsof tbе LiпneаnSoсiеtу,ZсlologуЗ, no.9: 53_62. |876. Thе GеogrаphiсаlDistributioпсlfAпimаIs.2vols.London: -М.F. Масmillan& Сo.

T h е f i r s tp u Ь l i сr е р r е s е аnd Сriсk 1953.)

Watson,Jamеs (b. 1928)' and Сriсk, Franсis (Т9Т6-2004) .Watson In 1953 thе Amеriсan Jamеs and thе Еnglislrman Fгanсis Сriсk, working at Сambridgе Univеrsity, disсovеrеd thе strrrсturе of thе dеoхyribonuсlеiс aсid (DNA) molесulе, long suspесtеd to be thе сarriеr of thе gеnеtiс infoгmation within thе сеll. Thеy found that it was a pair of long molесulеs, гwinеd around еaсh othеr in a douЬlе hеlix (sееfiguге)'Vеry еxсitingly, thе DNA molесulе сonsists of four diffеrеnt kinds of smallеr moleсulеs:adеninе (A), thyminе (T), guaninе (G)' and сytosinе (C). Thе opposing DNA molе. сulеs arе linkеd togеthеr, with onе of еaсh kind of smallеr molесцlе always spесifiсally pairеd on thе othеr DNA molесulе with anothеr of thе four molесulеs (A with T, and G with С). s7atson and Сriсk immеdiatеly hypothеsizеd that thе ordеr of the smallеr molесulеsсarriеsthе infoгmation of hеrеdity,and so it provеd latеr in thе dесadе whеn thе so-сallеd gеnеtiссodе was сrасkеd. It was found that information fronr thе DNA is usеd to makе thе Ьuilding Ьloсks of thе сеll, as wеll as thе еnzymеsusеd to drivе proсеssеs.Iп 1962Watson, Сriсk, and a Univеrsity of London rеsеarсhеr, Мauriсе Wilkins' iointly rесеivеd thе NoЬеl Prizе for Меdiсinе or Physiology for thеir dеtеrmination of thе struсturе of DNA.

Initially, rеlationslrip vеry tеnsе' with thе for and thе lattеr rеgarding autoЬiographtcal Тhе L guagе). Thе systеmatis provе thаt evoIutionаг standing' and that lrеnс standing aЬout wholе о сonfеrеnсеon thе philor quitе еqual to that of \й4 thе sаmе Ьiolоgv dерaг ..tlrе most unplеаsаr-rthr s o n i s d е t a i l е di n h i s с i : and thе morе traditiona By thе 1960s, howеvс iar tесhniquеsсould thr аtiorr, that wеrе inrpfAnimalsand Plantsunder (C. Darwin),507 Domestication Vascularplants,diversificationof, 87-88,88/ Velctciraptor, 4 53, 520f Venerealdisease, 440 Veracausa,699, 9 16-917 Vermeij,Geerat,551 VertebratePaleontology(A. S. Romer), 835 Vertebrates. SeeOrigin of vertebrates Vestiges of the Natural History of C,reatien(R. Chambers\,6-7, 8f, 474, 649, 81.9,893-896,903, 9t4 Vicariance allopatricspeciation, 187, 18 8 f Vicariancebiogeography, 450 Vicq-d'Az-yr, Felix,581 Virchow, Rudolf, 626 Virus-mediated process transduction, 679 V i r a l i s m3, 0 , 3 6 3 ,4 4 6 4 4 7 , 6 4 5 , 7 7 3 Vogt,Oskar,885-886 Volcanisnr, massextinctions and, 7r8-719

s77

L)78

Index Vortriige iiber D escendenztb eorie (\Weismann), 909 Voyageof the Beagle,The (C. Darwin), 504-505,903 Vries,Hugo de,748, 850, 852, 896-897 Wdchtershduser, G., 59,62-63 'S(addington, ClonradHal, 216, 403, 5 6 1 f ,8 3 3 , 8 9 8 - 9 0 0 lfade, Michael.|ohn,900-901 'Wagner, Andreas,5 18-519 'Wagner, Giinter, 638 '!fake, David 8' 901-902 Walking.SeeBipedalism 'Wallace, AlfredRussel , 1.4,22-23,29, 5 0 6 ,5 1 1 ,6 3 9 ,8 6 4 ,8 9 5 ,9 0 3 - 9 0 6 ; Batesand,445; biogeography,44T, 448,449f; Cuvierand,502;Jenkirr a n d ,6 6 3 ;L v e l la n d ,8 1 8 - 8 1 9 ; Malthus and,703-704;religion,513, 514; superorganism concept,874 'Wallace, Bruce,33f, 34f Wallace's line,448, 449f,904 '!7alsh, Bruce,70 I Wanderingpolygyny, 242 Warfield,B. B., 3.58,363-364 "Warm little pond" term,49,60-61 'Water, origin of life and, 51-54 Vy'ater Babies,Tbe (Kingsley),650 'Watson, James,906-909 Watson,John 8., 541 'Weber, Bruce,327 Wedekind,C., 250 Wedgwood, .fosiah,9/, 504, 508 'Weischaus, Eric, 529 'Weismann, AugustFriedrichLeopold, 333-334,545, 570,674, 832, 868, 909-9rO 'Weldon, W. F. R.. 537,754,801-802, 910-913 \ W e l l sG, . P . , 6 4 6 \ W e l l sH, . G . , 6 4 6 Wells,Jonathan,377 V{eltrdthsel, Der (Haeckel),622, 626 West-Eberhar d, M. J., 247 lilhat ls Darwinism?(Hodge),357 Vlhat Is Lrlei (Schrddinger), 886 W h e w e l lW , i l l i a m ,l 5 i , 1 8 f ,3 5 ' t , 7 7l , 894,973-919 'Whitcomb, John C., lr., 359,373,428, 842 White, Andrew, 350-352 White,EllenG., 359, 842

\White,Michael D., 199,742-744 J. 'Whitman, Charles,215 'Whooping 439 c