Takingallthesefactstogether,itisobviousthattheview,whichwasentertainedbyMantellandtheprobabilityofwhichwasdemonstratedbyyourowndistinguishedanatomist,Leidy,whilemuchadditionalevidenceinthesamedirectionhasbeenfurnishedbyProfessorCope,thatsomeoftheseanimalsmayhavewalkedupontheirhindlegsasbirdsdo,acquiresgreatweight。
Infact,therecanbenoreasonabledoubtthatoneofthesmallerformsoftheOrnithoscelida,Compsognathus,thealmostentireskeletonofwhichhasbeendiscoveredintheSolenhofenslates,wasabipedalanimal。Thepartsofthisskeletonaresomewhattwistedoutoftheirnaturalrelations,buttheaccompanyingfiguregivesajustviewofthegeneralformofCompsognathusandoftheproportionsofitslimbs;which,insomerespects,aremorecompletelybird-likethanthoseofotherOrnithoscelida。
Fig。7。——RestorationofCompsognathusLongipesWehavehadtostretchthedefinitionoftheclassofbirdssoastoincludebirdswithteethandbirdswithpaw-likeforelimbsandlongtails。ThereisnoevidencethatCompsognathuspossessedfeathers;but,ifitdid,itwouldbehardindeedtosaywhetheritshouldbecalledareptilianbirdoranavianreptile。
AsCompsognathuswalkeduponitshindlegs,itmusthavemadetrackslikethoseofbirds。AndasthestructureofthelimbsofseveralofthegiganticOrnithoscelida,suchasIguanodon,leadstotheconclusionthattheyalsomayhaveconstantly,oroccasionally,assumedthesameattitude,apeculiarinterestattachestothefactthat,intheWealdenstrataofEngland,therearetobefoundgiganticfootsteps,arrangedinorderlikethoseoftheBrontozoum,andwhichtherecanbenoreasonabledoubtweremadebysomeoftheOrnithoscelida,theremainsofwhicharefoundinthesamerocks。And,knowingthatreptilesthatwalkedupontheirhindlegsandsharedmanyoftheanatomicalcharactersofbirdsdidonceexist,itbecomesaveryimportantquestionwhetherthetracksintheTriasofMassachusetts,towhichIreferredsometimeago,andwhichformerlyusedtobeunhesitatinglyascribedtobirds,maynotallhavebeenmadebyornithoscelidanreptiles;andwhether,ifwecouldobtaintheskeletonsoftheanimalswhichmadethesetracks,weshouldnotfindinthemtheactualstepsoftheevolutionalprocessbywhichreptilesgaverisetobirds。
TheevidentialvalueofthefactsIhavebroughtforwardinthisLecturemustbeneitherovernorunderestimated。Itisnothistoricalproofoftheoccurrenceoftheevolutionofbirdsfromreptiles,forwehavenosafegroundforassumingthattruebirdshadnotmadetheirappearanceatthecommencementoftheMesozoicepoch。Itis,infact,quitepossiblethatallthesemoreorlessavi-formreptilesoftheMesozoicepochsarenottermsintheseriesofprogressionfrombirdstoreptilesatall,butsimplythemoreorlessmodifieddescendantsofPalaeozoicformsthroughwhichthattransitionwasactuallyeffected。
WearenotinapositiontosaythattheknownOrnithoscelidaareintermediateintheorderoftheirappearanceontheearthbetweenreptilesandbirds。Allthatcanbesaidisthat,ifindependentevidenceoftheactualoccurrenceofevolutionisproducible,thentheseintercalaryformsremoveeverydifficultyinthewayofunderstandingwhattheactualstepsoftheprocess,inthecaseofbirds,mayhavebeen。
Thatintercalaryformsshouldhaveexistedinancienttimesisanecessaryconsequenceofthetruthofthehypothesisofevolution;and,hence,theevidenceIhavelaidbeforeyouinproofoftheexistenceofsuchforms,is,sofarasitgoes,infavourofthathypothesis。
Thereisanotherseriesofextinctreptileswhichmaybesaidtobeintercalarybetweenreptilesandbirds,insofarastheycombinesomeofthecharactersofboththesegroups;andwhich,astheypossessedthepowerofflight,mayseem,atfirstsight,tobenearerrepresentativesoftheformsbywhichthetransitionfromthereptiletothebirdwaseffected,thantheOrnithoscelida。
ThesearethePterosauria,orPterodactyles,theremainsofwhicharemetwiththroughouttheseriesofMesozoicrocks,fromtheliastothechalk,andsomeofwhichattainedagreatsize,theirwingshavingaspanofeighteenortwentyfeet。
Theseanimals,intheformandproportionsoftheheadandneckrelativelytothebody,andinthefactthattheendsofthejawswereoften,ifnotalways,moreorlessextensivelyensheathedinhornybeaks,remindusofbirds。Moreover,theirbonescontainedaircavities,renderingthemspecificallylighter,asisthecaseinmostbirds。Thebreastbonewaslargeandkeeled,asinmostbirdsandinbats,andtheshouldergirdleisstrikinglysimilartothatofordinarybirds。But,itseemstome,thatthespecialresemblanceofpterodactylestobirdsendshere,unlessImayaddtheentireabsenceofteethwhichcharacterisesthegreatpterodactyles(Pteranodon)
discoveredbyProfessorMarsh。Allotherknownpterodactyleshaveteethlodgedinsockets。Inthevertebralcolumnandthehindlimbstherearenospecialresemblancestobirds,andwhenweturntothewingstheyarefoundtobeconstructedonatotallydifferentprinciplefromthoseofbirds。
Fig。8。——PterodactylusSpectabilis(VonMeyer)。
Therearefourfingers。Thesefourfingersarelarge,andthreeofthem,thosewhichanswertothethumbandtwofollowingfingersinmyhand——areterminatedbyclaws,whilethefourthisenormouslyprolongedandconvertedintoagreatjointedstyle。
Youseeatonce,fromwhatIhavestatedaboutabird’swing,thattherecouldbenothinglesslikeabird’swingthanthisis。Itwasconcludedbygeneralreasoningthatthisfingerhadtheofficeofsupportingawebwhichextendedbetweenitandthebody。Anexistingspecimenprovesthatsuchwasreallythecase,andthatthepterodactylesweredevoidoffeathers,butthatthefingerssupportedavastweblikethatofabat’swing;infact,therecanbenodoubtthatthisancientreptileflewafterthefashionofabat。
Thus,thoughthepterodactyleisareptilewhichhasbecomemodifiedinsuchamannerastoenableittofly,andtherefore,asmightbeexpected,presentssomepointsofresemblancetootheranimalswhichfly;ithas,sotospeak,goneoffthelinewhichleadsdirectlyfromreptilestobirds,andhasbecomedisqualifiedforthechangeswhichleadtothecharacteristicorganisationofthelatterclass。Therefore,viewedinrelationtotheclassesofreptilesandbirds,thepterodactylesappeartometobe,inalimitedsense,intercalaryforms;buttheyarenotevenapproximatelylinear,inthesenseofexemplifyingthosemodificationsofstructurethroughwhichthepassagefromthereptiletothebirdtookplace。
LECTURESONEVOLUTION
III
THEDEMONSTRATIVEEVIDENCEOFEVOLUTION
Theoccurrenceofhistoricalfactsissaidtobedemonstrated,whentheevidencethattheyhappenedisofsuchacharacterastorendertheassumptionthattheydidnothappeninthehighestdegreeimprobable;andthequestionInowhavetodealwithis,whetherevidenceinfavouroftheevolutionofanimalsofthisdegreeofcogencyis,orisnot,obtainablefromtherecordofthesuccessionoflivingformswhichispresentedtousbyfossilremains。
ThosewhohaveattendedtotheprogressofpalaeontologyareawarethatevidenceofthecharacterwhichIhavedefinedhasbeenproducedinconsiderableandcontinually-increasingquantityduringthelastfewyears。Indeed,theamountandthesatisfactorynatureofthatevidencearesomewhatsurprising,whenweconsidertheconditionsunderwhichalonewecanhopetoobtainit。
Itisobviouslyuselesstoseekforsuchevidenceexceptinlocalitiesinwhichthephysicalconditionshavebeensuchastopermitofthedepositofanunbroken,orbutrarelyinterrupted,seriesofstratathroughalongperiodoftime;inwhichthegroupofanimalstobeinvestigatedhasexistedinsuchabundanceastofurnishtherequisitesupplyofremains;andinwhich,finally,thematerialscomposingthestrataaresuchastoensurethepreservationoftheseremainsinatolerablyperfectandundisturbedstate。
Itsohappensthatthecasewhich,atpresent,mostnearlyfulfilsalltheseconditionsisthatoftheseriesofextinctanimalswhichculminatesinthehorses;bywhichtermImeantodenotenotmerelythedomesticanimalswithwhichweareallsowellacquainted,buttheirallies,theass,zebra,quagga,andthelike。Inshort,Iuse"horses"astheequivalentofthetechnicalnameEquidae,whichisappliedtothewholegroupofexistingequineanimals。
Thehorseisinmanywaysaremarkableanimal;notleastsointhefactthatitpresentsuswithanexampleofoneofthemostperfectpiecesofmachineryinthelivingworld。Intruth,amongtheworksofhumaningenuityitcannotbesaidthatthereisanylocomotivesoperfectlyadaptedtoitspurposes,doingsomuchworkwithsosmallaquantityoffuel,asthismachineofnature’smanufacture——thehorse。And,asanecessaryconsequenceofanysortofperfection,ofmechanicalperfectionasofothers,youfindthatthehorseisabeautifulcreature,oneofthemostbeautifulofallland-animals。Lookattheperfectbalanceofitsform,andtherhythmandforceofitsaction。
Thelocomotivemachineryis,asyouareaware,residentinitsslenderforeandhindlimbs;theyareflexibleandelasticlevers,capableofbeingmovedbyverypowerfulmuscles;and,inordertosupplytheengineswhichworktheseleverswiththeforcewhichtheyexpend,thehorseisprovidedwithaveryperfectapparatusforgrindingitsfoodandextractingtherefromtherequisitefuel。
Withoutattemptingtotakeyouveryfarintotheregionofosteologicaldetail,Imustneverthelesstroubleyouwithsomestatementsrespectingtheanatomicalstructureofthehorse;
and,moreespecially,willitbeneedfultoobtainageneralconceptionofthestructureofitsforeandhindlimbs,andofitsteeth。ButIshallonlytouchuponthosepointswhichareabsolutelyessentialtoourinquiry。
Letusturninthefirstplacetothefore-limb。Inmostquadrupeds,asinourselves,thefore-armcontainsdistinctbonescalledtheradiusandtheulna。Thecorrespondingregioninthehorseseemsatfirsttopossessbutonebone。Carefulobservation,however,enablesustodistinguishinthisboneapartwhichclearlyanswerstotheupperendoftheulna。Thisiscloselyunitedwiththechiefmassofthebonewhichrepresentstheradius,andrunsoutintoaslendershaftwhichmaybetracedforsomedistancedownwardsuponthebackoftheradius,andtheninmostcasesthinsoutandvanishes。Ittakesstillmoretroubletomakesureofwhatisneverthelessthefact,thatasmallpartofthelowerendoftheboneofthehorse’sforearm,whichisonlydistinctinaveryyoungfoal,isreallythelowerextremityoftheulna。
Whatiscommonlycalledthekneeofahorseisitswrist。
The"cannonbone"answerstothemiddleboneofthefivemetacarpalbones,whichsupportthepalmofthehandinourselves。The"pastern,""coronary,"and"coffin"bonesofveterinariansanswertothejointsofourmiddlefingers,whilethehoofissimplyagreatlyenlargedandthickenednail。Butifwhatliesbelowthehorse’s"knee"thuscorrespondstothemiddlefingerinourselves,whathasbecomeofthefourotherfingersordigits?Wefindintheplacesofthesecondandfourthdigitsonlytwoslendersplint-likebones,abouttwo-
thirdsaslongasthecannonbone,whichgraduallytapertotheirlowerendsandbearnofingerjoints,or,astheyaretermed,phalanges。Sometimes,smallbonyorgristlynodulesaretobefoundatthebasesofthesetwometacarpalsplints,anditisprobablethattheserepresentrudimentsofthefirstandfifthtoes。Thus,thepartofthehorse’sskeleton,whichcorrespondswiththatofthehumanhand,containsoneovergrownmiddledigit,andatleasttwoimperfectlateraldigits;andtheseanswer,respectively,tothethird,thesecond,andthefourthfingersinman。
Correspondingmodificationsarefoundinthehindlimb。
Inourselves,andinmostquadrupeds,thelegcontainstwodistinctbones,alargebone,thetibia,andasmallerandmoreslenderbone,thefibula。But,inthehorse,thefibulaseems,atfirst,tobereducedtoitsupperend;ashortslenderboneunitedwiththetibia,andendinginapointbelow,occupyingitsplace。Examinationofthelowerendofayoungfoal’sshinbone,however,showsadistinctportionofosseousmatter,whichisthelowerendofthefibula;sothattheapparentlysingle,lowerendoftheshinboneisreallymadeupofthecoalescedendsofthetibiaandfibula,justasthe,apparentlysingle,lowerendofthefore-armboneiscomposedofthecoalescedradiusandulna。
Theheelofthehorseisthepartcommonlyknownasthehock。
Thehindercannonboneanswerstothemiddlemetatarsalboneofthehumanfoot,thepastern,coronary,andcoffinbones,tothemiddletoebones;thehindhooftothenail;asinthefore-
foot。And,asinthefore-foot,therearemerelytwosplintstorepresentthesecondandthefourthtoes。Sometimesarudimentofafifthtoeappearstobetraceable。
Theteethofahorsearenotlesspeculiarthanitslimbs。Thelivingengine,likeallothers,mustbewellstokedifitistodoitswork;andthehorse,ifitistomakegooditswearandtear,andtoexerttheenormousamountofforcerequiredforitspropulsion,mustbewellandrapidlyfed。Tothisend,goodcuttinginstrumentsandpowerfulandlastingcrushersareneedful。Accordingly,thetwelvecuttingteethofahorseareclose-setandconcentratedinthefore-partofitsmouth,likesomanyadzesorchisels。Thegrindersormolarsarelarge,andhaveanextremelycomplicatedstructure,beingcomposedofanumberofdifferentsubstancesofunequalhardness。Theconsequenceofthisisthattheywearawayatdifferentrates;
and,hence,thesurfaceofeachgrinderisalwaysasunevenasthatofagoodmillstone。
Ihavesaidthatthestructureofthegrindingteethisverycomplicated,theharderandthesofterpartsbeing,asitwere,interlacedwithoneanother。Theresultofthisisthat,asthetoothwears,thecrownpresentsapeculiarpattern,thenatureofwhichisnotveryeasilydecipheredatfirst;butwhichitisimportantweshouldunderstandclearly。Eachgrindingtoothoftheupperjawhasanouterwallsoshapedthat,ontheworncrown,itexhibitstheformoftwocrescents,oneinfrontandonebehind,withtheirconcavesidesturnedoutwards。Fromtheinnersideofthefrontcrescent,acrescenticfrontridgepassesinwardsandbackwards,anditsinnerfaceenlargesintoastronglongitudinalfoldorpillar。
Fromthefrontpartofthehindercrescent,abackridge
takesalikedirection,andalsohasitspillar。
Thedeepinterspacesorvalleysbetweentheseridgesandtheouterwallarefilledbybonysubstance,whichiscalledcement,andcoatsthewholetooth。
Thepatternofthewornfaceofeachgrindingtoothofthelowerjawisquitedifferent。Itappearstobeformedoftwocrescent-
shapedridges,theconvexitiesofwhichareturnedoutwards。
Thefreeextremityofeachcrescenthasapillar,andthereisalargedoublepillarwherethetwocrescentsmeet。Thewholestructureis,asitwere,imbeddedincement,whichfillsupthevalleys,asintheuppergrinders。
Ifthegrindingfacesofanupperandofalowermolarofthesamesideareappliedtogether,itwillbeseenthattheopposedridgesarenowhereparallel,butthattheyfrequentlycross;
andthatthus,intheactofmastication,ahardsurfaceintheoneisconstantlyappliedtoasoftsurfaceintheother,andviceversa。Theythusconstituteagrindingapparatusofgreatefficiency,andonewhichisrepairedasfastasitwears,owingtothelong-continuedgrowthoftheteeth。
Someotherpeculiaritiesofthedentitionofthehorsemustbenoticed,astheybearuponwhatIshallhavetosaybyandby。
Thusthecrownsofthecuttingteethhaveapeculiardeeppit,whichgivesrisetothewell-known"mark"ofthehorse。Thereisalargespacebetweentheouterincisorsandthefrontgrinder。
Inthisspacetheadultmalehorsepresents,neartheincisorsoneachside,aboveandbelow,acanineor"tush,"whichiscommonlyabsentinmares。Inayounghorse,moreover,thereisnotunfrequentlytobeseeninfrontofthefirstgrinder,averysmalltooth,whichsoonfallsout。Ifthissmalltoothbecountedasone,itwillbefoundthatthereareseventeethbehindthecanineoneachside;namely,thesmalltoothinquestion,andthesixgreatgrinders,amongwhich,byanunusualpeculiarity,theforemosttoothisratherlargerthanthosewhichfollowit。
Ihavenowenumeratedthosecharacteristicstructuresofthehorsewhichareofmostimportanceforthepurposewehaveinview。
Toanyonewhoisacquaintedwiththemorphologyofvertebratedanimals,theyshowthatthehorsedeviateswidelyfromthegeneralstructureofmammals;andthatthehorsetypeis,inmanyrespects,anextrememodificationofthegeneralmammalianplan。Theleastmodifiedmammals,infact,havetheradiusandulna,thetibiaandfibula,distinctandseparate。Theyhavefivedistinctandcompletedigitsoneachfoot,andnooneofthesedigitsisverymuchlargerthantherest。Moreover,intheleastmodifiedmammals,thetotalnumberoftheteethisverygenerallyforty-four,whileinhorses,theusualnumberisforty,andintheabsenceofthecanines,itmaybereducedtothirty-six;theincisorteetharedevoidofthefoldseeninthoseofthehorse:thegrindersregularlydiminishinsizefromthemiddleoftheseriestoitsfrontend;whiletheircrownsareshort,earlyattaintheirfulllength,andexhibitsimpleridgesortubercles,inplaceofthecomplexfoldingsofthehorse’sgrinders。
Hencethegeneralprinciplesofthehypothesisofevolutionleadtotheconclusionthatthehorsemusthavebeenderivedfromsomequadrupedwhichpossessedfivecompletedigitsoneachfoot;whichhadthebonesofthefore-armandofthelegcompleteandseparate;andwhichpossessedforty-fourteeth,amongwhichthecrownsoftheincisorsandgrindershadasimplestructure;whilethelattergraduallyincreasedinsizefrombeforebackwards,atanyrateintheanteriorpartoftheseries,andhadshortcrowns。
Andifthehorsehasbeenthusevolved,andtheremainsofthedifferentstagesofitsevolutionhavebeenpreserved,theyoughttopresentuswithaseriesofformsinwhichthenumberofthedigitsbecomesreduced;thebonesofthefore-armandleggraduallytakeontheequinecondition;andtheformandarrangementoftheteethsuccessivelyapproximatetothosewhichobtaininexistinghorses。
Letusturntothefacts,andseehowfartheyfulfiltheserequirementsofthedoctrineofevolution。
InEuropeabundantremainsofhorsesarefoundintheQuaternaryandlaterTertiarystrataasfarasthePlioceneformation。
Butthesehorses,whicharesocommoninthecave-depositsandinthegravelsofEurope,areinallessentialrespectslikeexistinghorses。AndthatistrueofallthehorsesofthelatterpartofthePlioceneepoch。But,indepositswhichbelongtotheearlierPlioceneandlaterMioceneepochs,andwhichoccurinBritain,inFrance,inGermany,inGreece,inIndia,wefindanimalswhichareextremelylikehorses——which,infact,aresosimilartohorses,thatyoumayfollowdescriptionsgiveninworksupontheanatomyofthehorseupontheskeletonsoftheseanimals——butwhichdifferinsomeimportantparticulars。
Forexample,thestructureoftheirforeandhindlimbsissomewhatdifferent。Theboneswhich,inthehorse,arerepresentedbytwosplints,imperfectbelow,areaslongasthemiddlemetacarpalandmetatarsalbones;and,attachedtotheextremityofeach,isadigitwiththreejointsofthesamegeneralcharacterasthoseofthemiddledigit,onlyverymuchsmaller。Thesesmalldigitsaresodisposedthattheycouldhavehadbutverylittlefunctionalimportance,andtheymusthavebeenratherofthenatureofthedew-claws,suchasaretobefoundinmanyruminantanimals。TheHipparion,astheextinctEuropeanthree-toedhorseiscalled,infact,presentsafootsimilartothatoftheAmericanProtohippus
(Fig。9),exceptthat,intheHipparion,thesmallerdigitsaresituatedfartherback,andareofsmallerproportionalsize,thanintheProtohippus。
Theulnaisslightlymoredistinctthaninthehorse;andthewholelengthofit,asaveryslendershaft,intimatelyunitedwiththeradius,iscompletelytraceable。Thefibulaappearstobeinthesameconditionasinthehorse。TheteethoftheHipparionareessentiallysimilartothoseofthehorse,butthepatternofthegrindersisinsomerespectsalittlemorecomplex,andthereisadepressiononthefaceoftheskullinfrontoftheorbit,whichisnotseeninexistinghorses。
IntheearlierMiocene,andperhapsthelaterEocenedepositsofsomepartsofEurope,anotherextinctanimalhasbeendiscovered,whichCuvier,whofirstdescribedsomefragmentsofit,consideredtobeaPalaeotherim。Butasfurtherdiscoveriesthrewnewlightuponitsstructure,itwasrecognisedasadistinctgenus,underthenameofAnchitherium。
Initsgeneralcharacters,theskeletonofAnchitherium
isverysimilartothatofthehorse。Infact,LartetandDeBlainvillecalleditPal?otheriumequinumorhippoides;andDeChristol,in1847,saidthatitdifferedfromHipparioninlittlemorethanthecharactersofitsteeth,andgaveitthenameofHipparitherium。Eachfootpossessesthreecompletetoes;
whilethelateraltoesaremuchlargerinproportiontothemiddletoethaninHipparion,anddoubtlessrestedonthegroundinordinarylocomotion。
Theulnaiscompleteandquitedistinctfromtheradius,thoughfirmlyunitedwiththelatter。Thefibulaseemsalsotohavebeencomplete。Itslowerend,thoughintimatelyunitedwiththatofthetibia,isclearlymarkedofffromthelatterbone。
Thereareforty-fourteeth。Theincisorshavenostrongpit。
Thecaninesseemtohavebeenwelldevelopedinbothsexes。
Thefirstofthesevengrinders,which,asIhavesaid,isfrequentlyabsent,and,whenitdoesexist,issmallinthehorse,isagood-sizedandpermanenttooth,whilethegrinderwhichfollowsitisbutlittlelargerthanthehinderones。
Thecrownsofthegrindersareshort,andthoughthefundamentalpatternofthehorse-toothisdiscernible,thefrontandbackridgesarelesscurved,theaccessorypillarsarewanting,andthevalleys,muchshallower,arenotfilledupwithcement。
Sevenyearsago,whenIhappenedtobelookingcriticallyintothebearingofpalaentologicalfactsuponthedoctrineofevolution,itappearedtomethattheAnchitherium,theHipparion,andthemodernhorses,constituteaseriesinwhichthemodificationsofstructurecoincidewiththeorderofchronologicaloccurrence,inthemannerinwhichtheymustcoincide,ifthemodernhorsesreallyaretheresultofthegradualmetamorphosis,inthecourseoftheTertiaryepoch,ofalessspecialisedancestralform。AndIfoundbycorrespondencewiththelateeminentFrenchanatomistandpalaeontologist,M。Lartet,thathehadarrivedatthesameconclusionfromthesamedata。
ThattheAnchitheriumtypehadbecomemetamorphosedintotheHippariontype,andthelatterintotheEquine
type,inthecourseofthatperiodoftimewhichisrepresentedbythelatterhalfoftheTertiarydeposits,seemedtometobetheonlyexplanationofthefactsforwhichtherewasevenashadowofprobability。
And,hence,Ihaveeversinceheldthatthesefactsaffordevidenceoftheoccurrenceofevolution,which,inthesensealreadydefined,maybetermeddemonstrative。
AllwhohaveoccupiedthemselveswiththestructureofAnchitherium,fromCuvieronwards,haveacknowledgeditsmanypointsoflikenesstoawell-knowngenusofextinctEocenemammals,Palaeotherium。Indeed,aswehaveseen,CuvierregardedhisremainsofAnchitheriumasthoseofaspeciesofPalaeotherium。Hence,inattemptingtotracethepedigreeofthehorsebeyondtheMioceneepochandtheAnchitheroidform,InaturallysoughtamongthevariousspeciesofPalaeotheroidanimalsforitsnearestally,andIwasledtoconcludethatthePalaeotheriumminus(Plagiolophus)
representedthenextstepmorenearlythananyformthenknown。
Ithinkthatthisopinionwasfullyjustifiable;buttheprogressofinvestigationhasthrownanunexpectedlightonthequestion,andhasbroughtusmuchnearerthancouldhavebeenanticipatedtoaknowledgeofthetrueseriesoftheprogenitorsofthehorse。
Youareallawarethat,whenyourcountrywasfirstdiscoveredbyEuropeans,therewerenotracesoftheexistenceofthehorseinanypartoftheAmericanContinent。TheaccountsoftheconquestofMexicodwellupontheastonishmentofthenativesofthatcountrywhentheyfirstbecameacquaintedwiththatastoundingphenomenon——amanseateduponahorse。
Nevertheless,theinvestigationsofAmericangeologistshaveprovedthattheremainsofhorsesoccurinthemostsuperficialdepositsofbothNorthandSouthAmerica,justastheydoinEurope。Therefore,forsomereasonorother——nofeasiblesuggestiononthatsubject,sofarasIknow,hasbeenmade——thehorsemusthavediedoutonthiscontinentatsomeperiodprecedingthediscoveryofAmerica。OflateyearstherehasbeendiscoveredinyourWesternTerritoriesthatmarvellousaccumulationofdeposits,admirablyadaptedforthepreservationoforganicremains,towhichIreferredtheotherevening,andwhichfurnishesuswithaconsecutiveseriesofrecordsofthefaunaoftheolderhalfoftheTertiaryepoch,forwhichwehavenoparallelinEurope。Theyhaveyieldedfossilsinanexcellentstateofconservationandinunexamplednumberandvariety。
TheresearchesofLeidyandothershaveshownthatformsalliedtotheHipparionandtheAnchitheriumaretobefoundamongtheseremains。Butitisonlyrecentlythattheadmirablyconceivedandmostthoroughlyandpatientlyworked-outinvestigationsofProfessorMarshhavegivenusajustideaofthevastfossilwealth,andofthescientificimportance,ofthesedeposits。IhavehadtheadvantageofglancingoverthecollectionsinYaleMuseum;andIcantrulysaythat,sofarasmyknowledgeextends,thereisnocollectionfromanyoneregionandseriesofstratacomparable,forextent,orforthecarewithwhichtheremainshavebeengottogether,orfortheirscientificimportance,totheseriesoffossilswhichhehasdepositedthere。Thisvastcollectionhasyieldedevidencebearinguponthequestionofthepedigreeofthehorseofthemoststrikingcharacter。IttendstoshowthatwemustlooktoAmerica,ratherthantoEurope,fortheoriginalseatoftheequineseries;andthatthearchaicformsandsuccessivemodificationsofthehorse’sancestryarefarbetterpreservedherethaninEurope。
ProfessorMarsh’skindnesshasenabledmetoputbeforeyouadiagram,everyfigureinwhichisanactualrepresentationofsomespecimenwhichistobeseenatYaleatthispresenttime(Fig。9)。
Fig。9。
ThesuccessionofformswhichhehasbroughttogethercarriesusfromthetoptothebottomoftheTertiaries。Firstly,thereisthetruehorse。NextwehavetheAmericanPlioceneformofthehorse(Pliohippus);intheconformationofitslimbsitpresentssomeveryslightdeviationsfromtheordinaryhorse,andthecrownsofthegrindingteethareshorter。ThencomestheProtohippus,whichrepresentstheEuropeanHipparion,havingonelargedigitandtwosmallonesoneachfoot,andthegeneralcharactersofthefore-armandlegtowhichIhavereferred。ButitismorevaluablethantheEuropeanHipparionforthereasonthatitisdevoidofsomeofthepeculiaritiesofthatform——peculiaritieswhichtendtoshowthattheEuropeanHipparionisratheramemberofacollateralbranch,thanaforminthedirectlineofsuccession。
Next,inthebackwardorderintime,istheMiohippus,
whichcorrespondsprettynearlywiththeAnchitheriumofEurope。Itpresentsthreecompletetoes——onelargemedianandtwosmallerlateralones;andthereisarudimentofthatdigit,whichanswerstothelittlefingerofthehumanhand。
TheEuropeanrecordofthepedigreeofthehorsestopshere;
intheAmericanTertiaries,onthecontrary,theseriesofancestralequineformsiscontinuedintotheEoceneformations。
AnolderMioceneform,termedMesohippus,hasthreetoesinfront,withalargesplint-likerudimentrepresentingthelittlefinger;andthreetoesbehind。Theradiusandulna,thetibiaandthefibula,aredistinct,andtheshortcrownedmolarteethareanchitherioidinpattern。
ButthemostimportantdiscoveryofallistheOrohippus,
whichcomesfromtheEoceneformation,andistheoldestmemberoftheequineseries,asyetknown。Herewefindfourcompletetoesonthefrontlimb,threetoesonthehindlimb,awell-
developedulna,awell-developedfibula,andshort-crownedgrindersofsimplepattern。
Thus,thankstotheseimportantresearches,ithasbecomeevidentthat,sofarasourpresentknowledgeextends,thehistoryofthehorse-typeisexactlyandpreciselythatwhichcouldhavebeenpredictedfromaknowledgeoftheprinciplesofevolution。Andtheknowledgewenowpossessjustifiesuscompletelyintheanticipation,thatwhenthestilllowerEocenedeposits,andthosewhichbelongtotheCretaceousepoch,haveyieldeduptheirremainsofancestralequineanimals,weshallfind,first,aformwithfourcompletetoesandarudimentoftheinnermostorfirstdigitinfront,with,probably,arudimentofthefifthdigitinthehindfoot;while,instillolderforms,theseriesofthedigitswillbemoreandmorecomplete,untilwecometothefive-toedanimals,inwhich,ifthedoctrineofevolutioniswellfounded,thewholeseriesmusthavetakenitsorigin。
ThatiswhatImeanbydemonstrativeevidenceofevolution。
Aninductivehypothesisissaidtobedemonstratedwhenthefactsareshowntobeinentireaccordancewithit。Ifthatisnotscientificproof,therearenomerelyinductiveconclusionswhichcanbesaidtobeproved。Andthedoctrineofevolution,atthepresenttime,restsuponexactlyassecureafoundationastheCopernicantheoryofthemotionsoftheheavenlybodiesdidatthetimeofitspromulgation。Itslogicalbasisispreciselyofthesamecharacter——thecoincidenceoftheobservedfactswiththeoreticalrequirements。
Theonlywayofescape,ifitbeawayofescape,fromtheconclusionswhichIhavejustindicated,isthesuppositionthatallthesedifferentequineformshavebeencreatedseparatelyatseparateepochsoftime;and,Irepeat,thatofsuchanhypothesisasthisthereneitheris,norcanbe,anyscientificevidence;and,assuredly,sofarasIknow,thereisnonewhichissupported,orpretendstobesupported,byevidenceorauthorityofanyotherkind。Icanbutthinkthatthetimewillcomewhensuchsuggestionsasthese,suchobviousattemptstoescapetheforceofdemonstration,willbeputuponthesamefootingasthesuppositionmadebysomewriters,whoareI
believenotcompletelyextinctatpresent,thatfossilsaremeresimulacra,arenoindicationsoftheformerexistenceoftheanimalstowhichtheyseemtobelong;butthattheyareeithersportsofnature,orspecialcreations,intended——asIheardsuggestedtheotherday——totestourfaith。
Infact,thewholeevidenceisinfavourofevolution,andthereisnoneagainstit。AndIsaythis,althoughperfectlywellawareoftheseemingdifficultieswhichhavebeenbuiltupuponwhatappearstotheuninformedtobeasolidfoundation。Imeetconstantlywiththeargumentthatthedoctrineofevolutioncannotbewellfounded,becauseitrequiresthelapseofaveryvastperiodoftime;whilethedurationoflifeupontheearththusimpliedisinconsistentwiththeconclusionsarrivedatbytheastronomerandthephysicist。ImayventuretosaythatIamfamiliarwiththoseconclusions,inasmuchassomeyearsago,whenPresidentoftheGeologicalSocietyofLondon,Itookthelibertyofcriticisingthem,andofshowinginwhatrespects,asitappearedtome,theylackedcompleteandthoroughdemonstration。But,puttingthatpointaside,supposethat,astheastronomers,orsomeofthem,andsomephysicalphilosophers,tellus,itisimpossiblethatlifecouldhaveenduredupontheearthforaslongaperiodasisrequiredbythedoctrineofevolution——supposingthattobeproved——Idesiretobeinformed,whatisthefoundationforthestatementthatevolutiondoesrequiresogreatatime?Thebiologistknowsnothingwhateveroftheamountoftimewhichmayberequiredfortheprocessofevolution。ItisamatteroffactthattheequineformswhichIhavedescribedtoyouoccur,intheorderstated,intheTertiaryformations。ButIhavenottheslightestmeansofguessingwhetherittookamillionofyears,ortenmillions,orahundredmillions,orathousandmillionsofyears,togiverisetothatseriesofchanges。Abiologisthasnomeansofarrivingatanyconclusionastotheamountoftimewhichmaybeneededforacertainquantityoforganicchange。Hetakeshistimefromthegeologist。Thegeologist,consideringtherateatwhichdepositsareformedandtherateatwhichdenudationgoesonuponthesurfaceoftheearth,arrivesatmoreorlessjustifiableconclusionsastothetimewhichisrequiredforthedepositofacertainthicknessofrocks;andifhetellsmethattheTertiaryformationsrequired500,000,000yearsfortheirdeposit,Isupposehehasgoodgroundforwhathesays,andI
takethatasameasureofthedurationoftheevolutionofthehorsefromtheOrohippusuptoitspresentcondition。
And,ifheisright,undoubtedlyevolutionisaveryslowprocess,andrequiresagreatdealoftime。Butsuppose,now,thatanastronomeroraphysicist——forinstance,myfriendSirWilliamThomson——tellsmethatmygeologicalauthorityisquitewrong;andthathehasweightyevidencetoshowthatlifecouldnotpossiblyhaveexisteduponthesurfaceoftheearth500,000,000yearsago,becausetheearthwouldhavethenbeentoohottoallowoflife,myreplyis:"Thatisnotmyaffair;
settlethatwiththegeologist,andwhenyouhavecometoanagreementamongyourselvesIwilladoptyourconclusion。"
Wetakeourtimefromthegeologistsandphysicists;anditismonstrousthat,havingtakenourtimefromthephysicalphilosopher’sclock,thephysicalphilosophershouldturnrounduponus,andsaywearetoofastortooslow。Whatwedesiretoknowis,isitafactthatevolutiontookplace?Astotheamountoftimewhichevolutionmayhaveoccupied,weareinthehandsofthephysicistandtheastronomer,whosebusinessitistodealwiththosequestions。
Ihavenow,ladiesandgentlemen,arrivedattheconclusionofthetaskwhichIsetbeforemyselfwhenIundertooktodelivertheselectures。Mypurposehasbeen,nottoenablethoseamongyouwhohavepaidnoattentiontothesesubjectsbefore,toleavethisroominaconditiontodecideuponthevalidityortheinvalidityofthehypothesisofevolution;butIhavedesiredtoputbeforeyoutheprinciplesuponwhichallhypothesesrespectingthehistoryofNaturemustbejudged;
andfurthermore,tomakeapparentthenatureoftheevidenceandtheamountofcogencywhichistobeexpectedandmaybeobtainedfromit。Tothisend,Ihavenothesitatedtoregardyouasgenuinestudentsandpersonsdesirousofknowingthetruth。Ihavenotshrunkfromtakingyouthroughlongdiscussions,thatIfearmayhavesometimestriedyourpatience;
andIhaveinflicteduponyoudetailswhichwereindispensable,butwhichmaywellhavebeenwearisome。ButIshallrejoice——I
shallconsiderthatIhavedoneyouthegreatestservicewhichitwasinmypowertodo——ifIhavethusconvincedyouthatthegreatquestionwhichwehavebeendiscussingisnotonetobedealtwithbyrhetoricalflourishes,orbylooseandsuperficialtalk;butthatitrequiresthekeenattentionofthetrainedintellectandthepatienceoftheaccurateobserver。
WhenIcommencedthisseriesoflectures,Ididnotthinkitnecessarytoprefacethemwithaprologue,suchasmightbeexpectedfromastrangerandaforeigner;forduringmybriefstayinyourcountry,Ihavefounditveryhardtobelievethatastrangercouldbepossessedofsomanyfriends,andalmostharderthataforeignercouldexpresshimselfinyourlanguageinsuchawayastobe,toallappearance,soreadilyintelligible。SofarasIcanjudge,thatmostintelligent,andperhaps,Imayadd,mostsingularlyactiveandenterprisingbody,yourpressreporters,donotseemtohavebeendeterredbymyaccentfromgivingthefullestaccountofeverythingthatI
happentohavesaid。
ButthevesselinwhichItakemydepartureto-morrowmorningisevennowreadytosliphermoorings;IawakefrommydelusionthatIamotherthanastrangerandaforeigner。Iamreadytogobacktomyplaceandcountry;but,beforedoingso,letme,bywayofepilogue,tendertoyoumymostheartythanksforthekindandcordialreceptionwhichyouhaveaccordedtome;
andletmethankyoustillmoreforthatwhichisthegreatestcomplimentwhichcanbeaffordedtoanypersoninmyposition——
thecontinuousandundisturbedattentionwhichyouhavebestoweduponthelongargumentwhichIhavehadthehonourtolaybeforeyou。
FOOTNOTES
(1)Theabsenceofanykeelonthebreast-boneandsomeotherosteologicalpeculiarities,observedbyProfessorMarsh,however,suggestthatHesperornismaybeamodificationofalessspecialisedgroupofbirdsthanthattowhichtheseexistingaquaticbirdsbelong。
(2)Asecondspecimen,discoveredin1877,andatpresentintheBerlinmuseum,showsanexcellentlypreservedskullwithteeth;
andthreedigits,allterminatedbyclaws,intheforelimb。
1893。
(3)Iusetheword"type"becauseitishighlyprobablethatmanyformsofAnchitherium-likeandHipparion-likeanimalsexistedintheMioceneandPlioceneepochs,justasmanyspeciesofthehorsetribeexistnow,anditishighlyimprobablethattheparticularspeciesofAnchitheriumorHipparion,whichhappentohavebeendiscovered,shouldbepreciselythosewhichhaveformedpartofthedirectlineofthehorse’spedigree。
(4)Sincethislecturewasdelivered,ProfessorMarshhasdiscoveredanewgenusofequinemammals(Eohippus)fromthelowestEocenedepositsoftheWest,whichcorrespondsverynearlytothisdescription。——AmericanJournalofScience,
November,1876。