Introduction

WhenonboardH.M.S.Beagle,asnaturalist,Iwasmuchstruck
withcertainfactsinthedistributionoftheinhabitantsofSouth
America,andinthegeologicalrelationsofthepresenttothepast
inhabitantsofthatcontinent.Thesefactsseemedtometothrowsome
lightontheoriginofspeciesthatmysteryofmysteries,asithas
beencalledbyoneofourgreatestphilosophers.Onmyreturnhome,
itoccurredtome,in1837,thatsomethingmightperhapsbemadeout
onthisquestionbypatientlyaccumulatingandreflectingonallsorts
offactswhichcouldpossiblyhaveanybearingonit.Afterfiveyears
workIallowedmyselftospeculateonthesubject,anddrewupsome
shortnotes;theseIenlargedin1844intoasketchoftheconclusions,
whichthenseemedtomeprobable:fromthatperiodtothepresent
dayIhavesteadilypursuedthesameobject.IhopethatImaybe
excusedforenteringonthesepersonaldetails,asIgivethemtoshow
thatIhavenotbeenhastyincomingtoadecision.

Althoughmuchremainsobscure,andwilllongremain
obscure,Icanentertainnodoubt,afterthemostdeliberatestudyand
dispassionatejudgementofwhichIamcapable,thattheviewwhich
mostnaturalistsentertain,andwhichIformerlyentertained
namely,thateachspecieshasbeenindependentlycreatedis
erroneous.Iamfullyconvincedthatspeciesarenotimmutable;but
thatthosebelongingtowhatarecalledthesamegeneraarelineal
descendantsofsomeotherandgenerallyextinctspecies,inthesame
mannerastheacknowledgedvarietiesofanyonespeciesarethe
descendantsofthatspecies.Furthermore,IamconvincedthatNatural
Selectionhasbeenthemainbutnotexclusivemeansofmodification.

ChapterIII.StruggleforExistence
Beforeenteringonthesubjectofthischapter,Imustmakea
fewpreliminaryremarks,toshowhowthestruggleforexistence
bearsonNaturalSelection.Ithasbeenseeninthelastchapterthat

CharlesDarwin
OntheOriginofSpecies
byMeansofNaturalSelection(1859)

CharlesDarwinsOriginofSpecies,inwhichheoutlined
histheoryofevolutionthroughnaturalselection,
remainsoneofthemostimportantworksofscientific
studyeverwritten.Afterconceivingthetheoryduring
hisvoyagewiththeHMSBeagle,Darwintooktwenty
yearstowritethisbook.Helaysouthistheoryslowly,
patiently,andmethodicallysosimplythatDarwins
youngcolleagueThomasHuxleyrespondedtothe
theorybyexclaiming,Howextremelystupid[Iam]not
tohavethoughtofthat!

amongstorganicbeingsinastateofnaturethereissomeindividual
variability;indeedIamnotawarethatthishaseverbeendisputed.
Howhaveallthoseexquisiteadaptationsofonepartofthe
organisationtoanotherpart,andtotheconditionsoflife,andofone
distinctorganicbeingtoanotherbeing,beenperfected?Weseethese
beautifulcoadaptationsmostplainlyinthewoodpeckerand
missletoe;andonlyalittlelessplainlyinthehumblestparasitewhich
clingstothehairsofaquadrupedorfeathersofabird;inthestructure
ofthebeetlewhichdivesthroughthewater;intheplumedseed
whichiswaftedbythegentlestbreeze;inshort,weseebeautiful
adaptationseverywhereandineverypartoftheorganicworld.

IshouldpremisethatIusethetermStruggleforExistenceina
largeandmetaphoricalsense,includingdependenceofonebeingon
another,andincluding(whichismoreimportant)notonlythelifeof
theindividual,butsuccessinleavingprogeny.Twocanineanimalsin
atimeofdearth,maybetrulysaidtostrugglewitheachotherwhich
shallgetfoodandlive.Butaplantontheedgeofadesertissaidto
struggleforlifeagainstthedrought,thoughmoreproperlyitshould
besaidtobedependentonthemoisture.Aplantwhichannually
producesathousandseeds,ofwhichonanaverageonlyonecomesto
maturity,maybemoretrulysaidtostrugglewiththeplantsofthe
sameandotherkindswhichalreadyclothetheground.Inthese
severalsenses,whichpassintoeachother,Iuseforconveniencesake
thegeneraltermofstruggleforexistence.
Astruggleforexistenceinevitablyfollowsfromthehighrate
atwhichallorganicbeingstendtoincrease.Everybeing,which
duringitsnaturallifetimeproducesseveraleggsorseeds,mustsuffer
destructionduringsomeperiodofitslife,andduringsomeseasonor
occasionalyear,otherwise,ontheprincipleofgeometricalincrease,its
numberswouldquicklybecomesoinordinatelygreatthatnocountry
couldsupporttheproduct.Hence,asmoreindividualsareproduced
thancanpossiblysurvive,theremustineverycasebeastrugglefor
existence,eitheroneindividualwithanotherofthesamespecies,or

withtheindividualsofdistinctspecies,orwiththephysical
conditionsoflife.ItisthedoctrineofMalthus1appliedwithmanifold
forcetothewholeanimalandvegetablekingdoms;forinthiscase
therecanbenoartificialincreaseoffood,andnoprudentialrestraint
frommarriage.Althoughsomespeciesmaybenowincreasingin
numbers,allcannotdoso,fortheworldwouldnotholdthem.

ChapterIV.NaturalSelection
Howwillthestruggleforexistence,discussedtoobrieflyin
thelastchapter,actinregardtovariation?Cantheprincipleof
selection,whichwehaveseenissopotentinthehandsofman,apply
innature?Ithinkweshallseethatitcanactmosteffectually.Letitbe
borneinmindinwhatanendlessnumberofstrangepeculiaritiesour
domesticproductions,and,inalesserdegree,thoseundernature,
vary;andhowstrongthehereditarytendencyis.Canit,then,be
thoughtimprobable,seeingthatvariationsusefultomanhave
undoubtedlyoccurred,thatothervariationsusefulinsomewayto
eachbeinginthegreatandcomplexbattleoflife,shouldsometimes
occurinthecourseofthousandsofgenerations?Ifsuchdooccur,can
wedoubt(rememberingthatmanymoreindividualsarebornthan
canpossiblysurvive)thatindividualshavinganyadvantage,
howeverslight,overothers,wouldhavethebestchanceofsurviving
andofprocreatingtheirkind?Ontheotherhand,wemayfeelsure
thatanyvariationintheleastdegreeinjuriouswouldberigidly
destroyed.Thispreservationoffavourablevariationsandtherejection
ofinjuriousvariations,IcallNaturalSelection.Variationsneither
usefulnorinjuriouswouldnotbeaffectedbynaturalselection,and
wouldbeleftafluctuatingelement.

ThomasRobertMalthus(17661834),Britishpoliticaleconomistwhoargued
thatallpopulationgrowthmustalwaysbelimitedbystarvationordisease.

ThisleadsmetosayafewwordsonwhatIcallSexual
Selection.Thisdepends,notonastruggleforexistence,butona
strugglebetweenthemalesforpossessionofthefemales;theresultis
notdeathtotheunsuccessfulcompetitor,butfewornooffspring.
Sexualselectionis,therefore,lessrigorousthannaturalselection.
Generally,themostvigorousmales,thosewhicharebestfittedfor
theirplacesinnature,willleavemostprogeny.Butinmanycases,
victorywilldependnotongeneralvigour,butonhavingspecial
weapons,confinedtothemalesex.Ahornlessstagorspurlesscock
wouldhaveapoorchanceofleavingoffspring.Sexualselectionby
alwaysallowingthevictortobreedmightsurelygiveindomitable
courage,lengthtothespur,andstrengthtothewingtostrikeinthe
spurredleg,aswellasthebrutalcockfighter,whoknowswellthathe
canimprovehisbreedbycarefulselectionofthebestcocks.Howlow
inthescaleofnaturethislawofbattledescends,Iknownot;male
alligatorshavebeendescribedasfighting,bellowing,andwhirling
round,likeIndiansinawardance,forthepossessionofthefemales;
malesalmonshavebeenseenfightingalldaylong;malestagbeetles
oftenbearwoundsfromthehugemandiblesofothermales.
Amongstbirds,thecontestisoftenofamorepeaceful
character.Allthosewhohaveattendedtothesubject,believethat
thereistheseverestrivalrybetweenthemalesofmanyspeciesto
attractbysingingthefemales.TherockthrushofGuiana,birdsof
paradise,andsomeothers,congregate;andsuccessivemalesdisplay
theirgorgeousplumageandperformstrangeanticsbeforethe
females,whichstandingbyasspectators,atlastchoosethemost
attractivepartner.Thosewhohavecloselyattendedtobirdsin
confinementwellknowthattheyoftentakeindividualpreferences
anddislikes:thusSirR.Heronhasdescribedhowonepiedpeacock
waseminentlyattractivetoallhishenbirds.Itmayappearchildishto
attributeanyeffecttosuchapparentlyweakmeans:Icannothere
enteronthedetailsnecessarytosupportthisview;butifmancanina
shorttimegiveelegantcarriageandbeautytohisbantams,according
tohisstandardofbeauty,Icanseenogoodreasontodoubtthat

femalebirds,byselecting,duringthousandsofgenerations,themost
melodiousorbeautifulmales,accordingtotheirstandardofbeauty,
mightproduceamarkedeffect.
Thusitis,asIbelieve,thatwhenthemalesandfemalesofany
animalhavethesamegeneralhabitsoflife,butdifferinstructure,
colour,orornament,suchdifferenceshavebeenmainlycausedby
sexualselection;thatis,individualmaleshavehad,insuccessive
generations,someslightadvantageoverothermales,intheir
weapons,meansofdefence,orcharms;andhavetransmittedthese
advantagestotheirmaleoffspring.

IllustrationsoftheActionofNaturalSelection.
Inordertomakeitclearhow,asIbelieve,naturalselection
acts,Imustbegpermissiontogiveoneortwoimaginaryillustrations.
Letustakethecaseofawolf,whichpreysonvariousanimals,
securingsomebycraft,somebystrength,andsomebyfleetness;and
letussupposethatthefleetestprey,adeerforinstance,hadfromany
changeinthecountryincreasedinnumbers,orthatotherpreyhad
decreasedinnumbers,duringthatseasonoftheyearwhenthewolfis
hardestpressedforfood.Icanundersuchcircumstancesseeno
reasontodoubtthattheswiftestandslimmestwolveswouldhavethe
bestchanceofsurviving,andsobepreservedorselected,provided
alwaysthattheyretainedstrengthtomastertheirpreyatthisorat
someotherperiodoftheyear,whentheymightbecompelledtoprey
onotheranimals.Icanseenomorereasontodoubtthis,thanthat
mancanimprovethefleetnessofhisgreyhoundsbycarefuland
methodicalselection,orbythatunconsciousselectionwhichresults
fromeachmantryingtokeepthebestdogswithoutanythoughtof
modifyingthebreed.
Evenwithoutanychangeintheproportionalnumbersofthe
animalsonwhichourwolfpreyed,acubmightbebornwithan
innatetendencytopursuecertainkindsofprey.Norcanthisbe
thoughtveryimprobable;forweoftenobservegreatdifferencesinthe
naturaltendenciesofourdomesticanimals;onecat,forinstance,
3

takingtocatchrats,anothermice;onecat,accordingtoMr.St.John,
bringinghomewingedgame,anotherharesorrabbits,andanother
huntingonmarshygroundandalmostnightlycatchingwoodcocksor
snipes.Thetendencytocatchratsratherthanmiceisknowntobe
inherited.Now,ifanyslightinnatechangeofhabitorofstructure
benefitedanindividualwolf,itwouldhavethebestchanceof
survivingandofleavingoffspring.Someofitsyoungwouldprobably
inheritthesamehabitsorstructure,andbytherepetitionofthis
process,anewvarietymightbeformedwhichwouldeithersupplant
orcoexistwiththeparentformofwolf.Or,again,thewolves
inhabitingamountainousdistrict,andthosefrequentingthe
lowlands,wouldnaturallybeforcedtohuntdifferentprey;andfrom
thecontinuedpreservationoftheindividualsbestfittedforthetwo
sites,twovarietiesmightslowlybeformed.Imayadd,that,
accordingtoMr.Pierce,therearetwovarietiesofthewolfinhabiting
theCatskillMountainsintheUnitedStates,onewithalight
greyhoundlikeform,whichpursuesdeer,andtheothermorebulky,
withshorterlegs,whichmorefrequentlyattackstheshepherds
flocks.

Letusnowturntothenectarfeedinginsects.Icouldgive
manyfacts,showinghowanxiousbeesaretosavetime;forinstance,
theirhabitofcuttingholesandsuckingthenectaratthebasesof
certainflowers,whichtheycan,withaverylittlemoretrouble,enter
bythemouth.Bearingsuchfactsinmind,Icanseenoreasontodoubt
thatanaccidentaldeviationinthesizeandformofthebody,orinthe
curvatureandlengthoftheproboscis,&c.,fartooslighttobe
appreciatedbyus,mightprofitabeeorotherinsect,sothatan
individualsocharacterisedwouldbeabletoobtainitsfoodmore
quickly,andsohaveabetterchanceoflivingandleaving
descendants.Itsdescendantswouldprobablyinheritatendencytoa
similarslightdeviationofstructure.Ontheotherhand,Ihave
foundbyexperimentthatthefertilityofclovergreatlydependson
beesvisitingandmovingpartsofthecorolla,soastopushthepollen

ontothestigmaticsurface.Hence,againitmightbeagreat
advantagetotheredclovertohaveashorterormoredeeplydivided
tubetoitscorolla,sothatthehivebeecouldvisititsflowers.ThusI
canunderstandhowaflowerandabeemightslowlybecome,either
simultaneouslyoroneaftertheother,modifiedandadaptedinthe
mostperfectmannertoeachother,bythecontinuedpreservationof
individualspresentingmutualandslightlyfavourabledeviationsof
structure.
Iamwellawarethatthisdoctrineofnaturalselection,
exemplifiedintheaboveimaginaryinstances,isopentothesame
objectionswhichwereatfirsturgedagainstSirCharlesLyellsnoble
viewsonthemodernchangesoftheearth,asillustrativeofgeology;
butwenowveryseldomheartheaction,forinstance,ofthecoast
waves,calledatriflingandinsignificantcause,whenappliedtothe
excavationofgiganticvalleysortotheformationofthelongestlines
ofinlandcliffs.Thatnaturalselectionwillalwaysactwithextreme
slowness,Ifullyadmit.Nothingcanbeeffected,unlessfavourable
variationsoccur,andvariationitselfisapparentlyalwaysaveryslow
process.Theprocesswilloftenbegreatlyretardedbyfree
intercrossing.Manywillexclaimthattheseseveralcausesareamply
sufficientwhollytostoptheactionofnaturalselection.Idonot
believeso.Ontheotherhand,Idobelievethatnaturalselectionwill
alwaysactveryslowly,oftenonlyatlongintervalsoftime,and
generallyononlyaveryfewoftheinhabitantsofthesameregionat
thesametime.Ifurtherbelieve,thatthisveryslow,intermittent
actionofnaturalselectionaccordsperfectlywellwithwhatgeology
tellsusoftherateandmanneratwhichtheinhabitantsofthisworld
havechanged.

ChapterVI.DifficultiesonTheory
Longbeforehavingarrivedatthispartofmywork,acrowdof
difficultieswillhaveoccurredtothereader.Someofthemareso
gravethattothisdayIcanneverreflectonthemwithoutbeing
4

staggered;but,tothebestofmyjudgment,thegreaternumberare
onlyapparent,andthosethatarerealarenot,Ithink,fataltomy
theory.
Thesedifficultiesandobjectionsmaybeclassedunderthe
followingheads:Firstly,why,ifspecieshavedescendedfromother
speciesbyinsensiblyfinegradations,dowenoteverywheresee
innumerabletransitionalforms?Secondly,isitpossiblethatan
animalhaving,forinstance,thestructureandhabitsofabat,could
havebeenformedbythemodificationofsomeanimalwithwholly
differenthabits?Canwebelievethatnaturalselectioncouldproduce,
ontheonehand,organsoftriflingimportance,suchasthetailofa
giraffe,whichservesasaflyflapper,and,ontheotherhand,organs
ofsuchwonderfulstructure,astheeye,ofwhichwehardlyasyet
fullyunderstandtheinimitableperfection?Thirdly,caninstinctsbe
acquiredandmodifiedthroughnaturalselection?

OrgansofExtremePerfectionandComplication.
Tosupposethattheeye,withallitsinimitablecontrivancesfor
adjustingthefocustodifferentdistances,foradmittingdifferent
amountsoflight,andforthecorrectionofsphericalandchromatic
aberration,couldhavebeenformedbynaturalselection,seems,I
freelyconfess,absurdinthehighestpossibledegree.Yetreasontells
me,thatifnumerousgradationsfromaperfectandcomplexeyeto
oneveryimperfectandsimple,eachgradebeingusefultoits
possessor,canbeshowntoexist;iffurther,theeyedoesvaryeverso
slightly,andthevariationsbeinherited,whichiscertainlythecase;
andifanyvariationormodificationintheorganbeeverusefultoan
animalunderchangingconditionsoflife,thenthedifficultyof
believingthataperfectandcomplexeyecouldbeformedbynatural
selection,thoughinsuperablebyourimagination,canhardlybe
consideredreal.

IntheArticulata2weseeanopticnervemerelycoatedwith
pigment,andwithoutanyothermechanism;andfromthislowstage,
numerousgradationsofstructure,branchingoffintwo
fundamentallydifferentlines,canbeshowntoexist,untilwereacha
moderatelyhighstageofperfection.Incertaincrustaceans,for
instance,thereisadoublecornea,theinneronedividedintofacets,
withineachofwhichthereisalensshapedswelling.Inother
crustaceansthetransparentconeswhicharecoatedbypigment,and
whichproperlyactonlybyexcludinglateralpencilsoflightWith
thesefacts,herefartoobrieflyandimperfectlygiven,whichshow
thatthereismuchgraduateddiversityintheeyesofliving
crustaceans,andbearinginmindhowsmallthenumberofliving
animalsisinproportiontothosewhichhavebecomeextinct,Icansee
noverygreatdifficulty(notmorethaninthecaseofmanyother
structures)inbelievingthatnaturalselectionhasconvertedthesimple
apparatusofanopticnervemerelycoatedwithpigmentandinvested
bytransparentmembrane,intoanopticalinstrumentasperfectasis
possessedbyanymemberofthegreatarticulateclass.
Hewhowillgothusfaroughtnottohesitatetogofurther,
andtoadmitthatastructureevenasperfectastheeyeofaneagle
mightbeformedbynaturalselection,althoughinthiscasehedoes
notknowanyofthetransitionalgrades.Hisreasonoughttoconquer
hisimagination;thoughIhavefeltthedifficultyfartookeenlytobe
surprisedatanydegreeofhesitationinextendingtheprincipleof
naturalselectiontosuchstartlinglengths.
Itisscarcelypossibletoavoidcomparingtheeyetoa
telescope.Weknowthatthisinstrumenthasbeenperfectedbythe
longcontinuedeffortsofthehighesthumanintellects;andwe
naturallyinferthattheeyehasbeenformedbyasomewhatanalogous
process.Butmaynotthisinferencebepresumptuous?Haveweany
righttoassumethattheCreatorworksbyintellectualpowerslike
thoseofman?

Asubclassofmarinelifesimilartostarfishorsealilies.

Weshouldbeextremelycautiousinconcludingthatanorgan
couldnothavebeenformedbytransitionalgradationsofsomekind.
Numerouscasescouldbegivenamongsttheloweranimalsofthe
sameorganperformingatthesametimewhollydistinctfunctions;
thusthealimentarycanalrespires,digests,andexcretesinthelarvaof
thedragonflyTwodistinctorganssometimesperform
simultaneouslythesamefunctioninthesameindividual;togiveone
instance,therearefishwithgillsorbranchiaethatbreathetheair
dissolvedinthewater,atthesametimethattheybreathefreeairin
theirswimbladders.Theillustrationoftheswimbladderinfishesis
agoodone,becauseitshowsusclearlythehighlyimportantfactthat
anorganoriginallyconstructedforonepurpose,namelyflotation,
maybeconvertedintooneforawhollydifferentpurpose,namely
respiration.Allphysiologistsadmitthattheswimbladderis
homologous,orideallysimilar,inpositionandstructurewiththe
lungsofthehighervertebrateanimals:hencethereseemstometobe
nogreatdifficultyinbelievingthatnaturalselectionhasactually
convertedaswimbladderintoalung,anorganusedexclusivelyfor
respiration.

ChapterXIV.RecapitulationandConclusion

Thatmanyandgraveobjectionsmaybeadvancedagainstthe
theoryofdescentwithmodificationthroughnaturalselection,Ido
notdeny.Ihaveendeavouredtogivetothemtheirfullforce.Nothing
atfirstcanappearmoredifficulttobelievethanthatthemore
complexorgansandinstinctsshouldhavebeenperfectednotby
meanssuperiorto,thoughanalogouswith,humanreason,butbythe
accumulationofinnumerableslightvariations,eachgoodforthe
individualpossessor.Nevertheless,thisdifficulty,thoughappearing
toourimaginationinsuperablygreat,cannotbeconsideredrealifwe
admitthefollowingpropositions,namely,thatgradationsinthe
perfectionofanyorganorinstinct,whichwemayconsider,eitherdo
nowexistorcouldhaveexisted,eachgoodofitskind,thatall

organsandinstinctsare,ineversoslightadegree,variable,and,
lastly,thatthereisastruggleforexistenceleadingtothepreservation
ofeachprofitabledeviationofstructureorinstinct.Thetruthofthese
propositionscannot,Ithink,bedisputed.
Authorsofthehighesteminenceseemtobefullysatisfied
withtheviewthateachspecieshasbeenindependentlycreated.To
myminditaccordsbetterwithwhatweknowofthelawsimpressed
onmatterbytheCreator,thattheproductionandextinctionofthe
pastandpresentinhabitantsoftheworldshouldhavebeendueto
secondarycauses,likethosedeterminingthebirthanddeathofthe
individual.WhenIviewallbeingsnotasspecialcreations,butasthe
linealdescendantsofsomefewbeingswhichlivedlongbeforethey
seemtometobecomeennobled.
Itisinterestingtocontemplateanentangledbank,clothed
withmanyplantsofmanykinds,withbirdssingingonthebushes,
withvariousinsectsflittingabout,andwithwormscrawlingthrough
thedampearth,andtoreflectthattheseelaboratelyconstructed
forms,sodifferentfromeachother,anddependentoneachotherin
socomplexamanner,haveallbeenproducedbylawsactingaround
us.Theselaws,takeninthelargestsense,beingGrowthwith
Reproduction;inheritancewhichisalmostimpliedbyreproduction;
Variabilityfromtheindirectanddirectactionoftheexternal
conditionsoflife,andfromuseanddisuse;aRatioofIncreasesohigh
astoleadtoaStruggleforLife,andasaconsequencetoNatural
Selection,entailingDivergenceofCharacterandtheExtinctionofless
improvedforms.Thus,fromthewarofnature,fromfamineand
death,themostexaltedobjectwhichwearecapableofconceiving,
namely,theproductionofthehigheranimals,directlyfollows.There
isgrandeurinthisviewoflife,withitsseveralpowers,havingbeen
originallybreathedintoafewformsorintoone;andthat,whilstthis
planethasgonecyclingonaccordingtothefixedlawofgravity,from
sosimpleabeginningendlessformsmostbeautifulandmost
wonderfulhavebeen,andarebeing,evolved.