IfthehistoryofthesePortoSantorabbitshadnotbeenknown,mostnaturalists,onobservingtheirmuchreducedsize,theircolour,reddishaboveandgreybeneath,theirtailsandearsnottippedwithblack,wouldhaverankedthemasadistinctspecies。TheywouldhavebeenstronglyconfirmedinthisviewbyseeingthemaliveintheZoologicalGardens,andhearingthattheyrefusedtocouplewithotherrabbits。Yetthisrabbit,whichtherecanbelittledoubtwouldthushavebeenrankedasadistinctspecies,ascertainlyoriginatedsincetheyear1420。Finally,fromthethreecasesoftherabbitswhichhaverunwildinPortoSanto,Jamaica,andtheFalklandIslands,weseethattheseanimalsdonot,undernewconditionsoflife,reverttoorretaintheiraboriginalcharacter,asissogenerallyassertedtobethecasebymostauthors。
OSTEOLOGICALCHARACTERS。
Whenweremember,ontheonehand,howfrequentlyitisstatedthatimportantpartsofthestructurenevervary;and,ontheotherhand,onwhatsmalldifferencesintheskeletonfossilspecieshaveoftenbeenfounded,thevariabilityoftheskullandofsomeotherbonesinthedomesticatedrabbitwelldeservesattention。Itmustnotbesupposedthatthemoreimportantdifferencesimmediatelytobedescribedstrictlycharacteriseanyonebreed;allthatcanbesaidis,thattheyaregenerallypresentincertainbreeds。Weshouldbearinmindthatselectionhasnotbeenappliedtofixanycharacterintheskeleton,andthattheanimalshavenothadtosupportthemselvesunderuniformhabitsoflife。Wecannotaccountformostofthedifferencesintheskeleton;butweshallseethattheincreasedsizeofthebody,duetocarefulnurtureandcontinuedselection,hasaffectedtheheadinaparticularmanner。Eventheelongationandloppingoftheearshaveinfluencedinasmalldegreetheformofthewholeskull。Thewantofexercisehasapparentlymodifiedtheproportionallengthofthelimbsincomparisonwiththatofthebody。
[Asastandardofcomparison,IpreparedskeletonsoftwowildrabbitsfromKent,onefromtheShetlandIslands,andonefromAntriminIreland。Asallthebonesinthesefourspecimensfromsuchdistantlocalitiescloselyresembledeachother,presentingscarcelyanyappreciabledifference,itmaybeconcludedthatthebonesofthewildrabbitaregenerallyuniformincharacter。
SKULL。
Ihavecarefullyexaminedskullsoftenlargelop—earedrabbits,andoffivecommondomesticrabbits,whichlatterdifferfromthelop—earedonlyinnothavingsuchlargebodiesorears,yetbothlargerthaninthewildrabbit。Firstforthetenlop—earedrabbits:inallthesetheskullisremarkablyelongatedincomparisonwithitsbreadth。Inawildrabbitthelengthwas3。15inches,inalargefancyrabbit4。3;whilstthebreadthofthecraniumenclosingthebrainwasinbothalmostexactlythesame。Evenbytakingasthestandardofcomparisonthewidestpartofthezygomaticarch,theskullsofthelop—earedareproportionallytotheirbreadththree—quartersofaninchtoolong。Thedepthoftheheadhasincreasedalmostinthesameproportionwiththelength;itisthebreadthalonewhichhasnotincreased。Theparietalandoccipitalbonesenclosingthebrainarelessarched,bothinalongitudinalandtransverseline,thaninthewildrabbit,sothattheshapeofthecraniumissomewhatdifferent。
Thesurfaceisrougher,lesscleanlysculptured,andthelinesofsuturesaremoreprominent。
Althoughtheskullsofthelargelop—earedrabbitsincomparisonwiththoseofthewildrabbitaremuchelongatedrelativelytotheirbreadth,yet,relativelytothesizeofbody,theyarefarfromelongated。Thelop—earedrabbitswhichIexaminedwere,thoughnotfat,morethantwiceasheavyasthewildspecimens;buttheskullwasveryfarfrombeingtwiceaslong。
Evenifwetakethefairerstandardofthelengthofbody,fromthenosetotheanus,theskullisnotonanaverageaslongasitoughttobebyathirdofaninch。InthesmallferalPortoSantorabbit,ontheotherhand,theheadrelativelytothelengthofbodyisaboutaquarterofaninchtoolong。
Thiselongationoftheskullrelativelytoitsbreadth,Ifindauniversalcharacter,notonlywiththelargelop—earedrabbits,butinalltheartificialbreeds;asiswellseenintheskulloftheAngora。Iwasatfirstmuchsurprisedatthefact,andcouldnotimaginewhydomesticationcouldproducethisuniformresult;buttheexplanationseemstolieinthecircumstancethatduringanumberofgenerationstheartificialraceshavebeencloselyconfined,andhavehadlittleoccasiontoexerteithertheirsenses,orintellect,orvoluntarymuscles;consequentlythebrain,asweshallpresentlymorefullysee,hasnotincreasedrelativelywiththesizeofbody。Asthebrainhasnotincreased,thebonycaseenclosingithasnotincreased,andthishasevidentlyaffectedthroughcorrelationthebreadthoftheentireskullfromendtoend。
(FIGURE6。SKULLOFWILDRABBIT,ofnaturalsize。
FIGURE7。SKULLOFLARGELOP—EAREDRABBIT,ofnaturalsize。
FIGURE8。PARTOFZYGOMATICARCH,showingtheprojectingendofthemalarboneoftheauditorymeatus:ofnaturalsize。Upperfigure,WildRabbit。
Lowerfigure,Lop—eared,hare—colouredRabbit。
FIGURE9。POSTERIORENDOFSKULL,ofnaturalsize,showingtheinter—
parietalbone。A。WildRabbit。B。FeralRabbitfromislandofP。Santo,nearMadeira。C。LargeLop—earedRabbit。)
Inalltheskullsofthelargelop—earedrabbits,thesupra—orbitalplatesorprocessesofthefrontalbonesaremuchbroaderthaninthewildrabbit,andtheygenerallyprojectmoreupwards。Inthezygomaticarchtheposteriororprojectingpointofthemalar—boneisbroaderandblunter;andinthespecimen,figure8,itissoinaremarkabledegree。Thispointapproachesnearertotheauditorymeatusthaninthewildrabbit,asmaybebestseeninfigure8;butthiscircumstancemainlydependsonthechangeddirectionofthemeatus。Theinter—parietalbone(seefigure9)differsmuchinshapeintheseveralskulls;generallyitismoreoval,thatismoreextendedinthelineofthelongitudinalaxisoftheskull,thaninthewildrabbit。Theposteriormarginof\"thesquareraisedplatform\"
(4/25。Waterhouse’Nat。Hist。Mammalia’volume2page36。)oftheocciput,insteadofbeingtruncated,orprojectingslightlyasinthewildrabbit,isinmostlop—earedrabbitspointed,asinfigure9,C。Theparamastoidsrelativelytothesizeoftheskullaregenerallymuchthickerthaninthewildrabbit。
(FIGURE10。OCCIPITALFORAMEN,ofnaturalsize,in——A。WildRabbit;B。
LargeLop—earedRabbit。)
Theoccipitalforamen(figure10)presentssomeremarkabledifferences:inthewildrabbit,theloweredgebetweenthecondylesisconsiderablyandalmostangularlyhollowedout,andtheupperedgeisdeeplyandsquarelynotched;hencethelongitudinalaxisexceedsthetransverseaxis。Intheskullsofthelop—earedrabbitsthetransverseaxisexceedsthelongitudinal;forinnoneoftheseskullswastheloweredgebetweenthecondylessodeeplyhollowedout;infiveofthemtherewasnouppersquarenotch,inthreetherewasatraceofthenotch,andintwoaloneitwaswelldeveloped。Thesedifferencesintheshapeoftheforamenareremarkable,consideringthatitgivespassagetosoimportantastructureasthespinalmarrow,thoughapparentlytheoutlineofthelatterisnotaffectedbytheshapeofthepassage。
(FIGURE11。SKULL,ofnaturalsize,ofHalf—lopRabbit,showingthedifferentdirectionoftheauditorymeatusonthetwosides,andtheconsequentgeneraldistortionoftheskull。Theleftearoftheanimal(orrightsideoffigure)loppedforwards。)
Inalltheskullsofthelargelop—earedrabbits,thebonyauditorymeatusisconspicuouslylargerthaninthewildrabbit。Inaskull4。3inchesinlength,andwhichbarelyexceededinbreadththeskullofawildrabbit(whichwas3。15inchesinlength),thelongerdiameterofthemeatuswasexactlytwiceasgreat。Theorificeismorecompressed,anditsmarginonthesidenearesttheskullstandsuphigherthantheouterside。Thewholemeatusisdirectedmoreforwards。Asinbreedinglop—earedrabbitsthelengthoftheears,andtheirconsequentloppingandlyingflatontheface,arethechiefpointsofexcellence,therecanhardlybeadoubtthatthegreatchangeinthesize,form,anddirectionofthebonymeatus,relativelytothissamepartinthewildrabbit,isduetothecontinuedselectionofindividualshavinglargerandlargerears。Theinfluenceoftheexternalearonthebonymeatusiswellshownintheskulls(Ihaveexaminedthree)ofhalf—lops(seefigure5),inwhichoneearstandsupright,andtheotherandlongerearhangsdown;forintheseskullstherewasaplaindifferenceintheformanddirectionofthebonymeatusonthetwosides。Butitisamuchmoreinterestingfact,thatthechangeddirectionandincreasedsizeofthebonymeatushaveslightlyaffectedonthesamesidethestructureofthewholeskull。Iheregiveadrawing(figure11)oftheskullofahalf—lop;anditmaybeobservedthatthesuturebetweentheparietalandfrontalbonesdoesnotrunstrictlyatrightanglestothelongitudinalaxisoftheskull;theleftfrontalboneprojectsbeyondtherightone;boththeposteriorandanteriormarginsoftheleftzygomaticarchonthesideoftheloppingearstandalittleinadvanceofthecorrespondingbonesontheoppositeside。Eventhelowerjawisaffected,andthecondylesarenotquitesymmetrical,thatontheleftstandingalittleinadvanceofthatontheright。Thisseemstomearemarkablecaseofcorrelationofgrowth。Whowouldhavesurmisedthatbykeepingananimalduringmanygenerationsunderconfinement,andsoleadingtothedisuseofthemusclesoftheears,andbycontinuallyselectingindividualswiththelongestandlargestears,hewouldthusindirectlyhaveaffectedalmosteverysutureintheskullandtheformofthelowerjaw!
Inthelargelop—earedrabbitstheonlydifferenceinthelowerjaw,incomparisonwiththatofthewildrabbit,isthattheposteriormarginoftheascendingramusisbroaderandmoreinflected。Theteethinneitherjawpresentanydifference,exceptthatthesmallincisors,beneaththelargeones,areproportionatelyalittlelonger。Themolarteethhaveincreasedinsizeproportionatelywiththeincreasedwidthoftheskull,measuredacrossthezygomaticarch,andnotproportionallywithitsincreasedlength。Theinnerlineofthesocketsofthemolarteethintheupperjawofthewildrabbitformsaperfectlystraightline;butinsomeofthelargestskullsofthelop—earedthislinewasplainlybowedinwards。Inonespecimentherewasanadditionalmolartoothoneachsideoftheupperjaw,betweenthemolarsandpremolars;butthesetwoteethdidnotcorrespondinsize;andasnorodenthassevenmolars,thisismerelyamonstrosity,thoughacuriousone。
Thefiveotherskullsofcommondomesticrabbits,someofwhichapproachinsizetheabove—describedlargestskulls,whilsttheothersexceedbutlittlethoseofthewildrabbit,areonlyworthnoticeaspresentingaperfectgradationinalltheabove—specifieddifferencesbetweentheskullsofthelargestlop—earedandwildrabbits。Inall,however,thesupra—
orbitalplatesareratherlarger,andinalltheauditorymeatusislarger,inconformitywiththeincreasedsizeoftheexternalears,thaninthewildrabbit。Thelowernotchintheoccipitalforameninsomewasnotsodeepasinthewildrabbit,butinallfiveskullstheuppernotchwaswelldeveloped。
TheskulloftheAngorarabbit,likethelatterfiveskulls,isintermediateingeneralproportions,andinmostothercharacters,betweenthoseofthelargestlop—earedandwildrabbits。Itpresentsonlyonesingularcharacter:thoughconsiderablylongerthantheskullofthewildrabbit,thebreadthmeasuredwithintheposteriorsupra—orbitalfissuresisnearlyathirdlessthaninthewild。Theskullsofthesilver—grey,andchinchillaandHimalayanrabbitsaremoreelongatedthaninthewild,withbroadersupra—orbitalplates,butdifferlittleinanyotherrespect,exceptingthattheupperandlowernotchesoftheoccipitalforamenarenotsodeeporsowelldeveloped。TheskulloftheMoscowrabbitscarcelydiffersatallfromthatofthewildrabbit。InthePortoSantoferalrabbitsthesupra—orbitalplatesaregenerallynarrowerandmorepointedthaninourwildrabbits。
Assomeofthelargestlop—earedrabbitsofwhichIpreparedskeletonswerecolouredalmostlikehares,andastheselatteranimalsandrabbitshave,asitisaffirmed,beenrecentlycrossedinFrance,itmightbethoughtthatsomeoftheabove—describedcharactershadbeenderivedfromacrossataremoteperiodwiththehare。ConsequentlyIexaminedskullsofthehare,butnolightcouldthusbethrownonthepeculiaritiesoftheskullsofthelargerrabbits。Itis,however,aninterestingfact,asillustratingthelawthatvarietiesofonespeciesoftenassumethecharactersofotherspeciesofthesamegenus,thatIfound,oncomparingtheskullsoftenspeciesofharesintheBritishMuseum,thattheydifferedfromeachotherchieflyintheverysamepointsinwhichdomesticrabbitsvary,——namely,ingeneralproportions,intheformandsizeofthesupra—orbitalplates,intheformofthefreeendofthemalarbone,andinthelineofsutureseparatingtheoccipitalandfrontalbones。Moreovertwoeminentlyvariablecharactersinthedomesticrabbit,namely,theoutlineoftheoccipitalforamenandtheshapeofthe\"raisedplatform\"oftheocciput,werelikewisevariableintwoinstancesinthesamespeciesofhare。
VERTEBRAE。
ThenumberisuniforminalltheskeletonswhichIhaveexamined,withtwoexceptions,namely,inoneofthesmallferalPortoSantorabbitsandinoneofthelargestlop—earedkinds;bothofthesehadasusualsevencervical,twelvedorsalwithribs,but,insteadofsevenlumbar,bothhadeightlumbarvertebrae。Thisisremarkable,asGervaisgivessevenasthenumberforthewholegenusLepus。Thecaudalvertebraeapparentlydifferbytwoorthree,butIdidnotattendtothem,andtheyaredifficulttocountwithcertainty。
(FIGURE12。ATLASVERTEBRAE,ofnaturalsize;inferiorsurfaceviewedobliquely。Upperfigure,WildRabbit。Lowerfigure,Hare—coloured,large,Lop—earedRabbit,a,supra—median,atlantoidprocess;b,infra—medianprocess。)
Inthefirstcervicalvertebra,oratlas,theanteriormarginoftheneuralarchvariesalittleinwildspecimens,beingeithernearlysmooth,orfurnishedwithasmallsupra—medianatlantoidprocess;Ihavefiguredaspecimenwiththelargestprocess(a)whichIhaveseen;butitwillbeobservedhowinferiorthisisinsizeanddifferentinshapetothatinalargelop—earedrabbit。Inthelatter,theinfra—medianprocess(b)isalsoproportionallymuchthickerandlonger。Thealaearealittlesquarerinoutline。
(FIGURE13。THIRDCERVICALVERTEBRAE,ofnaturalsize,of:A。WildRabbit;
B。Hare—coloured,large,Lop—earedRabbit,a,a,inferiorsurface;b,b,anteriorarticularsurfaces。)
THIRDCERVICALVERTEBRA。
Inthewildrabbit(figure13,Aa)thisvertebra,viewedontheinferiorsurface,hasatransverseprocess,whichisdirectedobliquelybackwards,andconsistsofasinglepointedbar;inthefourthvertebrathisprocessisslightlyforkedinthemiddle。Inthelargelop—earedrabbitsthisprocess(Ba)isforkedinthethirdvertebra,asinthefourthofthewildrabbit。Butthethirdcervicalvertebraeofthewildandlop—eared(Ab,B
b)rabbitsdiffermoreconspicuouslywhentheiranteriorarticularsurfacesarecompared;fortheextremitiesoftheantero—dorsalprocessesinthewildrabbitaresimplyrounded,whilstinthelop—earedtheyaretrifid,withadeepcentralpit。Thecanalforthespinalmarrowinthelop—eared(Bb)ismoreelongatedinatransversedirectionthaninthewildrabbit;
andthepassagesforthearteriesareofaslightlydifferentshape。Theseseveraldifferencesinthisvertebraseemtomewelldeservingattention。
FIRSTDORSALVERTEBRA。
Itsneuralspinevariesinlengthinthewildrabbit;beingsometimesveryshort,butgenerallymorethanhalfaslongasthatoftheseconddorsal;
butIhaveseenitintwolargelop—earedrabbitsthree—fourthsofthelengthofthatoftheseconddorsalvertebra。
(FIGURE14。DORSALVERTEBRAE,fromsixthtotenthinclusive,ofnaturalsize,viewedlaterally。A。WildRabbit。B。Large,Hare—coloured,socalledSpanishRabbit。)
NINTHANDTENTHDORSALVERTEBRAE。
Inthewildrabbittheneuralspineoftheninthvertebraisjustperceptiblythickerthanthatoftheeighth;andtheneuralspineofthetenthisplainlythickerandshorterthanthoseofalltheanteriorvertebrae。Inthelargelop—earedrabbitstheneuralspinesofthetenth,ninth,andeighthvertebrae,andeveninaslightdegreethatoftheseventh,areverymuchthicker,andofsomewhatdifferentshape,incomparisonwiththoseofthewildrabbit。Sothatthispartofthevertebralcolumndiffersconsiderablyinappearancefromthesamepartinthewildrabbit,andcloselyresemblesinaninterestingmannerthesesamevertebraeinsomespeciesofhares。IntheAngora,Chinchilla,andHimalayanrabbits,theneuralspinesoftheeighthandninthvertebraeareinaslightdegreethickerthaninthewild。Ontheotherhand,inoneoftheferalPortoSantorabbits,whichinmostofitscharactersdeviatesfromthecommonwildrabbit,inadirectionexactlyoppositetothatassumedbythelargelop—earedrabbits,theneuralspinesoftheninthandtenthvertebraewerenotatalllargerthanthoseoftheseveralanteriorvertebra。InthissamePortoSantospecimentherewasnotraceintheninthvertebraoftheanteriorlateralprocesses(seefigure14),whichareplainlydevelopedinallBritishwildrabbits,andstillmoreplainlydevelopedinthelargelop—earedrabbits。Inahalf—wildrabbitfromSandonPark(4/26。TheserabbitshaverunwildforaconsiderabletimeinSandonPark,andinotherplacesinStaffordshireandShropshire。Theyoriginated,asIhavebeeninformedbythegamekeeper,fromvariously—coloureddomesticrabbitswhichhadbeenturnedout。Theyvaryincolour;butmanyaresymmetricallycoloured,beingwhitewithastreakalongthespine,andwiththeearsandcertainmarksabouttheheadofablackish—greytint。Theyhaveratherlongerbodiesthancommonrabbits),ahaemalspinewasmoderatelywelldevelopedontheundersideofthetwelfthdorsalvertebra,andIhaveseenthisinnootherspecimen。
LUMBARVERTEBRAE。
Ihavestatedthatintwocasestherewereeightinsteadofsevenlumbarvertebrae。ThethirdlumbarvertebraeinoneskeletonofawildBritishrabbit,andinoneofthePortoSantoferalrabbits,hadahaemalspine;
whilstinfourskeletonsoflargelop—earedrabbits,andintheHimalayanrabbit,thissamevertebrahadawelldevelopedhaemalspine。
PELVIS。
Infourwildspecimensthisbonewasalmostabsolutelyidenticalinshape;
butinseveraldomesticatedbreedsshadesofdifferencescouldbedistinguished。Inthelargelop—earedrabbits,thewholeupperpartoftheiliumisstraighter,orlesssplayedoutwards,thaninthewildrabbit;andthetuberosityontheinnerlipoftheanteriorandupperpartoftheiliumisproportionallymoreprominent。
(FIGURE15。TERMINALBONEOFSTERNUM,ofnaturalsize,A。WildRabbit。B。
Hare—coloured,Lop—earedRabbit。C。Hare—colouredSpanishRabbit。(N。B。Theleft—handangleoftheupperarticularextremityofBwasbroken,andhasbeenaccidentallythusrepresented。))
STERNUM。
Theposteriorendoftheposteriorsternalboneinthewildrabbit(figure15,A)isthinandslightlyenlarged;insomeofthelargelop—earedrabbits(B)itismuchmoreenlargedtowardstheextremity;whilstinotherspecimens(C)itkeepsnearlyofthesamebreadthfromendtoend,butismuchthickerattheextremity。
(FIGURE16。ACROMIONOFSCAPULA,ofnaturalsize。A。WildRabbit。B,C,D,Large,Lop—earedRabbits。)
SCAPULA。
Theacromionsendsoutarectangularbar,endinginanobliqueknob,whichlatterinthewildrabbit(figure16,A)variesalittleinshapeandsize,asdoestheapexoftheacromioninsharpness,andthepartjustbelowtherectangularbarinbreadth。Butthevariationsintheserespectsinthewildrabbitareveryslight:whilstinthelargelop—earedrabbitstheyareconsiderable。Thusinsomespecimens(B)theobliqueterminalknobisdevelopedintoashortbar,forminganobtuseanglewiththerectangularbar。Inanotherspecimen(C)thesetwounequalbarsformnearlyastraightline。Theapexoftheacromionvariesmuchinbreadthandsharpness,asmaybeseenbycomparingfiguresB,C,andD。
LIMBS。
IntheseIcoulddetectnovariation;butthebonesofthefeetweretootroublesometocomparewithmuchcare。]
IhavenowdescribedallthedifferencesintheskeletonswhichIhaveobserved。Itisimpossiblenottobestruckwiththehighdegreeofvariabilityorplasticityofmanyofthebones。Weseehowerroneoustheoften—repeatedstatementis,thatonlythecrestsoftheboneswhichgiveattachmenttomusclesvaryinshape,andthatonlypartsofslightimportancebecomemodifiedunderdomestication。Noonewillsay,forinstance,thattheoccipitalforamen,ortheatlas,orthethirdcervicalvertebraisapartofslightimportance。Iftheseveralvertebraeofthewildandlop—earedrabbits,ofwhichfigureshavebeengiven,hadbeenfoundfossil,palaeontologistswouldhavedeclaredwithouthesitationthattheyhadbelongedtodistinctspecies。
[THEEFFECTSOFTHEUSEANDDISUSEOFPARTS。
Inthelargelop—earedrabbitstherelativeproportionallengthofthebonesofthesameleg,andofthefrontandhindlegscomparedwitheachother,haveremainednearlythesameasinthewildrabbit;butinweight,thebonesofthehindlegsapparentlyhavenotincreasedindueproportionwiththefrontlegs。Theweightofthewholebodyinthelargerabbitsexaminedbymewasfromtwicetotwiceandahalfasgreatasthatofthewildrabbit;andtheweightofthebonesofthefrontandhindlimbstakentogether(excludingthefeet,onaccountofthedifficultyofcleaningsomanysmallbones)hasincreasedinthelargelop—earedrabbitsinnearlythesameproportion;consequentlyindueproportiontotheweightofbodywhichtheyhavetosupport。Ifwetakethelengthofthebodyasthestandardofcomparison,thelimbsofthelargerabbitshavenotincreasedinlengthindueproportionbyoneinchandahalf。Again,ifwetakeasthestandardofcomparisonthelengthoftheskull,which,aswehavebeforeseen,hasnotincreasedinlengthindueproportiontothelengthofbody,thelimbswillbefoundtobe,proportionallywiththoseofthewildrabbit,fromhalftothree—quartersofaninchtooshort。Hence,whateverstandardofcomparisonbetaken,thelimb—bonesofthelargelop—earedrabbitshavenotincreasedinlength,thoughtheyhaveinweight,infullproportiontotheotherpartsoftheframe;andthis,Ipresume,maybeaccountedforbytheinactivelifewhichduringmanygenerationstheyhavespent。Norhasthescapulaincreasedinlengthindueproportiontotheincreasedlengthofthebody。
Thecapacityoftheosseouscaseofthebrainisamoreinterestingpoint,towhichIwasledtoattendbyfinding,aspreviouslystated,thatwithalldomesticatedrabbitsthelengthoftheskullrelativelytoitsbreadthhasgreatlyincreasedincomparisonwiththatofthewildrabbits。Ifwehadpossessedalargenumberofdomesticatedrabbitsofnearlythesamesizewiththewildrabbits,itwouldhavebeenasimpletasktohavemeasuredandcomparedthecapacitiesoftheirskulls。Butthisisnotthecase:almostallthedomesticbreedshavelargerbodiesthanwildrabbits,andthelop—earedkindsaremorethandoubletheirweight。Asasmallanimalhastoexertitssenses,intellect,andinstinctsequallywithalargeanimal,weoughtnotbyanymeanstoexpectananimaltwiceorthriceaslargeasanothertohaveabrainofdoubleortreblethesize。(4/27。
SeeProf。Owen’sremarksonthissubjectinhispaperonthe’ZoologicalSignificanceoftheBrain,etc。,ofMan,etc。’readbeforeBrit。
Association1862:withrespecttoBirdssee’Proc。Zoolog。Soc。’January11,1848page8。)Now,afterweighingthebodiesoffourwildrabbits,andoffourlargebutnotfattenedlop—earedrabbits,Ifindthatonanaveragethewildaretothelop—earedinweightas1to2。17;inaveragelengthofbodyas1to1。41;whilstincapacityofskulltheyareas1to1。15。Henceweseethatthecapacityoftheskull,andconsequentlythesizeofthebrain,hasincreasedbutlittle,relativelytotheincreasedsizeofthebody;andthisfactexplainsthenarrownessoftheskullrelativelytoitslengthinalldomesticrabbits。
TABLE3:MEASUREMENTSOFWILDANDSEMI—WILDRABBITS。
I。LengthofSkull(inches)。
II。LengthofBodyfromIncisorstoAnus(inches)。
III。WeightofwholeBody(poundsandounces)。
IV。CapacityofSkullmeasuredbySmallShot(grains)。
V。CapacitycalculatedaccordingtoLengthofSkullrelativelytothatofNo。1(WildRabbit,Kent)(grains)。
VI。DifferencebetweenactualandcalculatedcapacitiesofSkulls(grains)。
VII。ShowinghowmuchpercenttheBrain,bycalculationaccordingtothelengthoftheSkullistoolight(—)ortooheavy(),relativelytotheBrainoftheWildRabbitNo。1。
NAMEOFBREED:I。II。III。IV。V。VI。
VII。
WILDANDSEMI—WILDRABBITS:
1。WildRabbit,Kent:3。1517。43,5972……
2
2。WildRabbit,ShetlandIslands:3。15……979……
2
3。WildRabbit,Ireland:3。15……992……
2
4。Domesticrabbit,runwild,Sandon:3。1518。5……977……
2
5。Wild,commonvariety,smallspecimen,Kent:2。9617。02,1487591338—
4
6。Wild,fawn—colouredvariety,Scotland:3。1……91895032—
3
7。Silver—grey,smallspecimen,Thetfordwarren:2。9515。52,1193891028
3
8。Feralrabbit,PortoSanto:2。83……89387320
2
9。Feralrabbit,PortoSanto:2。85……756879123—
16
10。Feralrabbit,PortoSanto:2。95……83591075—
9
AverageofthreePortoSantorabbits:2。88……82888860—
7
DOMESTICRABBITS:
11。Himalayan:3。520。5……9631080117—
12
12。Moscow:3。2517。03,88031002199—
24
13。Angora:3。519。53,16971080383—
54
14。Chinchilla:3。6522。0……9951126131—
13
15。Largelop—eared:4。124。57,010651265200—
18
16。Largelop—eared:4。125。07,1311531265112—
9
17。Largelop—eared:4。07……10371255218—
21
18。Largelop—eared:4。125。07,41208126557—
4
19。Largelop—eared:4。3……1232132694—
7
20。Largelop—eared:4。25……11241311187—
16
21。Largehare—coloured:3。8624。06,141131119160—
5
22。Averageofabovesevenlargelop—earedrabbits:4。1124。627,411361268132—
11
23。Hare(L。timidus)
Englishspecimen:3。617,01315
24。Hare(L。timidus)
Germanspecimen:3。827,01455
IntheupperhalfofTable3Ihavegiventhemeasurementsoftheskulloftenwildrabbits;andinthelowerhalf,ofeleventhoroughlydomesticatedkinds。Astheserabbitsdiffersogreatlyinsize,itisnecessarytohavesomestandardbywhichtocomparethecapacitiesoftheirskulls。Ihaveselectedthelengthofskullasthebeststandard,forinthelargerrabbitsithasnot,asalreadystated,increasedinlengthsomuchasthebody;butastheskull,likeeveryotherpart,variesinlength,neitheritnoranyotherpartaffordsaperfectstandard。
Inthefirstcolumnoffigurestheextremelengthoftheskullisgivenininchesanddecimals。Iamawarethatthesemeasurementspretendtogreateraccuracythanispossible;butIhavefoundittheleasttroubletorecordtheexactlengthwhichthecompassgave。Thesecondandthirdcolumnsgivethelengthandweightofbody,whenevertheseobservationsweremade。Thefourthcolumngivesthecapacityoftheskullbytheweightofsmallshotwithwhichtheskullswerefilled;butitisnotpretendedthattheseweightsareaccuratewithinafewgrains。Inthefifthcolumnthecapacityisgivenwhichtheskulloughttohavehadbycalculation,accordingtothelengthofskull,incomparisonwiththatofthewildrabbitNo。1;inthesixthcolumnthedifferencebetweentheactualandcalculatedcapacities,andinthesevenththepercentageofincreaseordecrease,aregiven。Forinstance,asthewildrabbitNo。5hasashorterandlighterbodythanthewildrabbitNo。1,wemighthaveexpectedthatitsskullwouldhavehadlesscapacity;theactualcapacity,asexpressedbytheweightofshot,is875grains,whichis97grainslessthanthatofthefirstrabbit。Butcomparingthesetworabbitsbythelengthoftheirskulls,weseethatinNo。1theskullis3。15inchesinlength,andinNo。52。96inchesinlength;accordingtothisratio,thebrainofNo。5oughttohavehadacapacityof913grainsofshot,whichisabovetheactualcapacity,butonlyby38grains。Or,toputthecaseinanotherway(asincolumnVII),thebrainofthissmallrabbit,No。5,forevery100grainsofweightisonly4grainstoolight,——thatis,itought,accordingtothestandardrabbitNo。1,tohavebeen4percentheavier。IhavetakentherabbitNo。
1asthestandardofcomparisonbecause,oftheskullshavingafullaveragelength,thishastheleastcapacity;sothatitistheleastfavourabletotheresultwhichIwishtoshow,namely,thatthebraininalllong—domesticatedrabbitshasdecreasedinsize,eitheractually,orrelativelytothelengthoftheheadandbody,incomparisonwiththebrainofthewildrabbit。HadItakentheIrishrabbit,No。3,asthestandard,thefollowingresultswouldhavebeensomewhatmorestriking。
TurningtoTable3:thefirstfourwildrabbitshaveskullsofthesamelength,andthesedifferbutlittleincapacity。TheSandonrabbit(No。4)
isinteresting,as,thoughnowwild,itisknowntobedescendedfromadomesticatedbreed,asisstillshownbyitspeculiarcolouringandlongerbody;neverthelesstheskullhasrecovereditsnormallengthandfullcapacity。Thenextthreerabbitsarewild,butofsmallsize,andtheyallhaveskullswithslightlylessenedcapacities。ThethreePortoSantoferalrabbits(Nos。8to10)offeraperplexingcase;theirbodiesaregreatlyreducedinsize,asinalesserdegreearetheirskullsinlengthandinactualcapacity,incomparisonwiththeskullsofwildEnglishrabbits。ButwhenwecomparethecapacitiesoftheskullinthethreePortoSantorabbits,weobserveasurprisingdifference,whichdoesnotstandinanyrelationtotheslightdifferenceinthelengthoftheirskulls,nor,asI
believe,toanydifferenceinthesizeoftheirbodies;butIneglectedweighingseparatelytheirbodies。Icanhardlysupposethatthemedullarymatterofthebraininthesethreerabbits,livingundersimilarconditions,candifferasmuchasisindicatedbytheproportionaldifferenceofcapacityintheirskulls;nordoIknowwhetheritispossiblethatonebrainmaycontainconsiderablymorefluidthananother。
HenceIcanthrownolightonthiscase。
LookingtothelowerhalfofTable3,whichgivesthemeasurementsofdomesticatedrabbits,weseethatinallthecapacityoftheskullisless,butinveryvariousdegrees,thanmighthavebeenanticipatedaccordingtothelengthoftheirskulls,relativelytothatofthewildrabbitNo。1。Inline22theaveragemeasurementsofsevenlargelop—earedrabbitsaregiven。Nowthequestionarises,hastheaveragecapacityoftheskullinthesesevenlargerabbitsincreasedasmuchasmighthavebeenexpectedfromthegreatlyincreasedsizeofbody。Wemayendeavourtoanswerthisquestionintwoways:intheupperhalfoftheTablewehavemeasurementsoftheskullsofsixsmallwildrabbits(Nos。5to10),andwefindthatonanaveragetheskullsare。18ofaninchshorter,andincapacity91grainsless,thantheaveragelengthandcapacityofthethreefirstwildrabbitsonthelist。Thesevenlargelop—earedrabbits,onanaverage,haveskulls4。11inchesinlength,and1136grainsincapacity;sothattheseskullshaveincreasedinlengthmorethanfivetimesasmuchastheskullsofthesixsmallwildrabbitshavedecreasedinlength;hencewemighthaveexpectedthattheskullsofthelargelop—earedrabbitswouldhaveincreasedincapacityfivetimesasmuchastheskullsofthesixsmallrabbitshavedecreasedincapacity;andthiswouldhavegivenanaverageincreasedcapacityof455grains,whilsttherealaverageincreaseisonly155grains。Again,thelargelop—earedrabbitshavebodiesofnearlythesameweightandsizeasthecommonhare,buttheirheadsarelonger;
consequently,ifthelop—earedrabbitshadbeenwild,itmighthavebeenexpectedthattheirskullswouldhavehadnearlythesamecapacityasthatoftheskullofthehare。Butthisisfarfrombeingthecase;fortheaveragecapacityofthetwohare—skulls(Nos。23,24)issomuchlargerthantheaveragecapacityofthesevenlop—earedskulls,thatthelatterwouldhavetobeincreased21percenttocomeuptothestandardofthehare。(4/23。Thisstandardisapparentlyconsiderablytoolow,forDr。
Crisp(’Proc。Zoolog。Soc。’1861page86)gives210grainsastheactualweightofthebrainofaharewhichweighed7pounds,and125grainsastheweightofthebrainofarabbitwhichweighed3pounds5ounces,thatis,thesameweightastherabbitNo。1inmylist。NowthecontentsoftheskullofrabbitNo。1inshotisinmytable972grains;andaccordingtoDr。Crisp’sratioof125to210,theskullofthehareoughttohavecontained1632grainsofshot,insteadofonly(inthelargesthareinmytable)1455grains。)
Ihavepreviouslyremarkedthat,ifwehadpossessedmanydomesticrabbitsofthesameaveragesizewiththewildrabbit,itwouldhavebeeneasytocomparethecapacityoftheirskulls。NowtheHimalayan,Moscow,andAngorarabbits(Nos。11,12,13ofTable3)areonlyalittlelargerinbodyandhaveskullsonlyalittlelonger,thanthewildanimal,andweseethattheactualcapacityoftheirskullsislessthaninthewildanimal,andconsiderablylessbycalculation(column7),accordingtothedifferenceinthelengthoftheirskulls。Thenarrownessofthebrain—caseinthesethreerabbitscouldbeplainlyseenandprovedbyexternalmeasurement。TheChinchillarabbit(No。14)isaconsiderablylargeranimalthanthewildrabbit,yetthecapacityofitsskullonlyslightlyexceedsthatofthewildrabbit。TheAngorarabbit,No。13,offersthemostremarkablecase;
thisanimalinitspurewhitecolourandlengthofsilkyfurbearsthestampoflongdomesticity。Ithasaconsiderablylongerheadandbodythanthewildrabbit,buttheactualcapacityofitsskullislessthanthatofeventhelittlewildPortoSantorabbits。Bythestandardofthelengthofskullthecapacity(seecolumn7)isonlyhalfofwhatitoughttohavebeen!Ikeptthisindividualanimalalive,anditwasnotunhealthynoridiotic。ThiscaseoftheAngorarabbitsomuchsurprisedme,thatI
repeatedallthemeasurementsandfoundthemcorrect。IhavealsocomparedthecapacityoftheskulloftheAngorawiththatofthewildrabbitbyotherstandards,namely,bythelengthandweightofthebody,andbytheweightofthelimb—bones;butbyallthesestandardsthebrainappearstobemuchtoosmall,thoughinalessdegreewhenthestandardofthelimb—
boneswasused;andthislattercircumstancemayprobablybeaccountedforbythelimbsofthisancientlydomesticatedbreedhavingbecomemuchreducedinweight,fromitslong—continuedinactivelife。HenceIinferthatintheAngorabreed,whichissaidtodifferfromotherbreedsinbeingquieterandmoresocial,thecapacityoftheskullhasreallyundergonearemarkableamountofreduction。]
Fromtheseveralfactsabovegiven,——namely,firstly,thattheactualcapacityoftheskullintheHimalayan,Moscow,andAngorabreeds,islessthaninthewildrabbit,thoughtheyareinalltheirdimensionsratherlargeranimals;secondly,thatthecapacityoftheskullofthelargelop—
earedrabbitshasnotbeenincreasedinnearlythesameratioasthecapacityoftheskullofthesmallerwildrabbitshasbeendecreased;andthirdly,thatthecapacityoftheskullinthesesamelargelop—earedrabbitsisveryinferiortothatofthehare,ananimalofnearlythesamesize,——Iconclude,notwithstandingtheremarkabledifferencesincapacityintheskullsofthesmallPortoSantorabbits,andlikewiseinthelargelop—earedkinds,thatinalllong—domesticatedrabbitsthebrainhaseitherbynomeansincreasedindueproportionwiththeincreasedlengthoftheheadandincreasedsizeofthebody,orthatithasactuallydecreasedinsize,relativelytowhatwouldhaveoccurredhadtheseanimalslivedinastateofnature。Whenwerememberthatrabbits,fromhavingbeendomesticatedandcloselyconfinedduringmanygenerations,cannothaveexertedtheirintellect,instincts,senses,andvoluntarymovements,eitherinescapingfromvariousdangersorinsearchingforfood,wemayconcludethattheirbrainswillhavebeenfeeblyexercised,andconsequentlyhavesufferedindevelopment。Wethusseethatthemostimportantandcomplicatedorganinthewholeorganisationissubjecttothelawofdecreaseinsizefromdisuse。
Finally,letussumupthemoreimportantmodificationswhichdomesticrabbitshaveundergone,togetherwiththeircausesasfaraswecanobscurelyseethem。Bythesupplyofabundantandnutritiousfood,togetherwithlittleexercise,andbythecontinuedselectionoftheheaviestindividuals,theweightofthelargerbreedshasbeenmorethandoubled。
Thebonesofthelimbstakentogetherhaveincreasedinweight,indueproportionwiththeincreasedweightofbody,butthehindlegshaveincreasedlessthanthefrontlegs;butinlengththeyhavenotincreasedindueproportion,andthismayhavebeencausedbythewantofproperexercise。Withtheincreasedsizeofthebodythethirdcervicalhasassumedcharacterspropertothefourthcervicalvertebra;andtheeighthandninthdorsalvertebraehavesimilarlyassumedcharacterspropertothetenthandposteriorvertebrae。Theskullinthelargerbreedshasincreasedinlength,butnotindueproportionwiththeincreasedlengthofbody;thebrainhasnotdulyincreasedindimensions,orhasevenactuallydecreased,andconsequentlythebonycaseforthebrainhasremainednarrow,andbycorrelationhasaffectedthebonesofthefaceandtheentirelengthoftheskull。Theskullhasthusacquireditscharacteristicnarrowness。Fromunknowncausesthesupra—orbitalprocessofthefrontalbonesandthefreeendofthemalarboneshaveincreasedinbreadth;andinthelargerbreedstheoccipitalforamenisgenerallymuchlessdeeplynotchedthaninwildrabbits。Certainpartsofthescapulaandtheterminalsternalboneshavebecomehighlyvariableinshape。Theearshavebeenincreasedenormouslyinlengthandbreadththroughcontinuedselection;theirweight,conjoinedprobablywiththedisuseoftheirmuscles,hascausedthemtolopdownwards;andthishasaffectedthepositionandformofthebonyauditorymeatus;andthisagain,bycorrelation,thepositioninaslightdegreeofalmosteveryboneintheupperpartoftheskull,andeventhepositionofthecondylesofthelowerjaw。
CHAPTER1。V。
DOMESTICPIGEONS。
ENUMERATIONANDDESCRIPTIONOFTHESEVERALBREEDS。
INDIVIDUALVARIABILITY。
VARIATIONSOFAREMARKABLENATURE。
OSTEOLOGICALCHARACTERS:SKULL,LOWERJAW,NUMBEROFVERTEBRAE。
CORRELATIONOFGROWTH:TONGUEWITHBEAK;EYELIDSANDNOSTRILSWITHWATTLED
SKIN。
NUMBEROFWING—FEATHERS,ANDLENGTHOFWING。
COLOURANDDOWN。
WEBBEDANDFEATHEREDFEET。
ONTHEEFFECTSOFDISUSE。
LENGTHOFFEETINCORRELATIONWITHLENGTHOFBEAK。
LENGTHOFSTERNUM,SCAPULA,ANDFURCULUM。
LENGTHOFWINGS。
SUMMARYONTHEPOINTSOFDIFFERENCEINTHESEVERALBREEDS。
Ihavebeenledtostudydomesticpigeonswithparticularcare,becausetheevidencethatallthedomesticracesaredescendedfromoneknownsourceisfarclearerthanwithanyotherancientlydomesticatedanimal。Secondly,becausemanytreatisesinseverallanguages,someofthemold,havebeenwrittenonthepigeon,sothatweareenabledtotracethehistoryofseveralbreeds。Andlastly,because,fromcauseswhichwecanpartlyunderstand,theamountofvariationhasbeenextraordinarilygreat。Thedetailswilloftenbetediouslyminute;butnoonewhoreallywantstounderstandtheprogressofchangeindomesticanimals,andespeciallynoonewhohaskeptpigeonsandhasmarkedthegreatdifferencebetweenthebreedsandthetruenesswithwhichmostofthempropagatetheirkind,willdoubtthatthisminutenessisworthwhile。Notwithstandingtheclearevidencethatallthebreedsarethedescendantsofasinglespecies,I
couldnotpersuademyselfuntilsomeyearshadpassedthatthewholeamountofdifferencebetweenthem,hadarisensincemanfirstdomesticatedthewildrock—pigeon。
Ihavekeptaliveallthemostdistinctbreeds,whichIcouldprocureinEnglandorfromtheContinent;