Aurochsgenetics,
acornerstoneofEuropeanbiodiversity
Picture:ManoloUno(c)StaffanWidstrand
Authors:
• drs.RonaldGoderie(TaurusFoundation);• dr.JohannesA.Lenstra(UtrechtUniversity,FacultyofVeterinaryMedicine);• MaulikUpadhyay(pHDWageningenUniversity);• dr.RichardCrooijmans(AnimalBreedingandGenomicsCentre,Wageningen
University);• ir.LeoLinnartz(ArkNature)
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PrefaceIn2015areportiswrittenonAurochsgenetics,madepossiblebyagrantfromtheDutchLibertyWildlifefund.ThisfundprovidedtheTaurusfoundationwithagrantofEUR20.000toconductgeneticresearchonaurochsanditsrelationwithnowadaysso-called‘primitive’breeds.Thisisthesummaryofthatreport.Thissummaryshortlydescribesthecurrentstateofaffairs,whatwedoknowearly2015abouttheaurochs,aboutdomesticcattleandtherelationshipofaurochsandtheprimitivebreedsusedintheTaurosProgramme.
Nijmegen,December2015.
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TableofcontentsPreface2Tableofcontents.........................................................................................................3Summary.....................................................................................................................41 Introduction..........................................................................................................62 Aurochs:ashortdescription..................................................................................83 Thedomesticationofcattle.................................................................................124 Results.................................................................................................................155 ThemeaningofgeneticresearchfortheTaurosProgramme...............................19Glossary.....................................................................................................................21Literature...................................................................................................................22Annex1:LONGLISTOF‘ORIGINAL’BREEDS.........................................................................24Annex2:SetwithcriteriafortheTaurosProgramme.................................................25Annex3:Briefsummaryofthebreedsinvolved.........................................................27Annex4:Overviewofconductedresearch.................................................................29Annex5:Theresultsoftheinheritancecheck............................................................30
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SummaryTheTaurosProgrammeisaninitiativeoftheDutchTaurusfoundation,since2011inclosecooperationwithRewildingEuropeandARKNature.Theprogrammeaimsatrewildingcattleandwantstodevelopacrediblereplacementfortheextinctaurochs.ThisreplacementiscalledtheTauros.TheTauroswillbebasedonthebestavailablescientificknowledge,involvingarcheo-zoology,history,ecology,ethologyandgenetics.
In2013afundingofEUR20.000fromtheDutchLibertyWildlifeFund,providedbyRewildingEurope,wasanewstimulustotheresearch.WealreadyhadbeenabletomakeacomparisonbetweenAurochsandalistof30EuropeanprimitivebreedsbasedonthemitochondrialDNA(thisisasmallDNA-particleoutsideofthecell-nucleus).ButwiththefundingfromtheLibertyWildlifeFundwenow–forthefirsttime-wereabletomakethecomparisonbetweenAurochsandour30‘primitive’breedsbasedontheDNAasitisfoundinthecellnucleus(autosomal1DNA).ThistypeofDNAinsidethecell-nucleuscodesforthephysicalproperties.ThisuniqueresultcouldonlybeachievedbecauseofthecooperationwithaDublin-basedresearchconsortiumwhoveryrecently(2015)wasabletosequencethecompletegenomeof(oneindividualof)Aurochs.
Ashortsummaryoftheresults–atthismoment–wouldbe:
§ Wealreadyknewthat-basedontheearlieranalysisofmitochondrialDNA(mtDNA)whichspecificallyinheritsfrommothertochild-certainso-calledPodolianbreeds(breedsfromItalyandtheBalkancountries)showacloserelationshipwithAurochs.Theseresultsarebasedontheanalysesofagreaternumberofaurochsindividuals(sinceit’sfareasiertoreconstructthesesmallfragmentsofDNA,thantoreconstructthecompletegenome);
§ TheinformationwenowgotfromthecomparisonofAurochswithour30primitivebreedsbasedontheautosomalDNA,showsthatIberianbreedsingeneralshowacloserrelationshiptoAurochs.SincetheinformationbasedonnuclearDNAtellsusfarmoreaboutinheritance,atthismomentweconsidertheanalysesoftheautosomalDNAasthemostvaluable;
§ ‘Rankingthestars’:Oneanalyses(NEIgeneticdistance)calculatedthedistancebetweentheanalysedbreedsincomparisontotheoneanalysedAurochs.ItwasfoundthatfourofthebreedsthatdoplayamajorroleintheTaurosbreedingprogramme(threeIberianandonePodolianbreed)areinthetop10andsixofourbreedsareinthetop15.
Theseresultsleadtothefollowingconclusionsandrecommendationsforthebreedingprogrammeandfollow-upresearch:
Conclusions
§ Theleadingpractiseinthebreedingprogrammesofar–combiningIberianandpodolianbreeds/individualswithAurochs-likephysicalproperties–isbeingconfirmed(bythecombinedresultsofautosomalandmtDNAandtheNEI-analyses).HoweveroneshouldtakeinconsiderationthatatthistimeonlyonespecimenofAurochshasbeenfullysequenced(autosomalDNA),sonewadditionalinformation
1Geneticjargonismarkedorangewhenfirstused.Ashortexplanationcanbefoundintheglossary.
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couldleadtoadjustmentsofthisconclusion.Itisthereforethatthebreedingprogrammestillusesaratherwidevarietyofcrossings(withtheselectedbreeds);
§ Inthe‘top10’anumberofbreedsisapparentthatarenot(yet)usedinthebreedingProgramme.Thereasonforthiswasthatonfenotypicalfeatures(thewaytheanimalslook)betterbreedsareavailable.Howeverinfuturestudiesweneedtolookbetteratsomeofthesebreeds(suchasthehighrankingBusha(notusedintheprogrammebecauseofitslimitedsize)
§ AsthePodolicawasrankedlow,weshouldreconsidertheuseofthisbreed,whichalreadywasusedinaverylimitednumber.
Recommendationsare:
FutureresearchneedstodivedeeperintothegeneticfeaturesofAurochstobeabletoidentifyspecificgenesfordomestication(andthusgettingmore‘grip’ongenesthatcouldbecrucialfortheoppositedirection:therewildingofthespecies).
MorefullysequencedspecimensofAurochsthatcoverasubstantialpartoftheEuropeancontinentareneeded,tobeabletogetanideaofthegeneticdiversitywithinAurochs.
Theresearchneedstoresultinpracticalguidelines–ontheshortrun–asthebaseforfutureselection.
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1 Introduction
TheTaurosProgrammeisaninitiativeoftheDutchTaurusfoundation,since2011incooperationwithRewildingEuropeandARKNature.Theprogrammeaimsatrewildingcattleandwantstodevelopacrediblereplacementfortheextinctaurochs.ThisreplacementiscalledtheTauros.InthenearandmoredistantfutureherdsofTauroswillliveaswildcattleandwillcontributetothesustainableconservationofEurope’s,openlandscapesandbiodiversity.Atthismoment,breedinggroupsofTaurosarealreadygrazingareasintheNetherlands,Spain,Portugal,Croatia,CzechRepublicandRomania.
TheTauroswillbebasedonthebestavailablescientificknowledge,involvingarcheo-zoology,history,ecology,ethologyandgenetics.Duringmillionsofyearsaurochsanditspredecessorsbecamegeneticallyadaptedtotheirenvironment.Therefore,theaurochsgeneswilltellushowtobreedaneffectivereplacementonthebasisofthegeneticvariationoftheprimitivedomesticcattleusedforbreedingtheaurochs.
In2013afundingofEUR20.000fromtheDutchLibertyWildlifeFund,providedthroughRewildingEurope,wasanewstimulustotheresearch.Thisallowedacomparisonoftheautosomal2DNA’sfrom30EuropeanprimitivebreedswiththeDNAoftheaurochs.Inadditionotherbreeds–suchastheCroatianBoskarinandMalteseCattle–wereaddedtothedatabase.ThebudgetoftheLibertyWildlifeFundwasalsousedforanalysingtheoffspringoftheTauroscrossbreds,avitalpartofthetotalresearch,sincewithoutfundingofthispartoftheresearchitwouldbeimpossibletojudgewhetherornotthebreedingisintherightdirection.ThefundingoftheDutchLibertyWildlifeFundwasalsousedtowritethisreportandbydoingsoprovidingbroaderaccesstotheTaurosresearch.
On-goinggeneticresearch
Indifferentpartsoftheworldresearchgroupsareworkingonthegeneticstructureofaurochs.ThisstartedwiththesequencingofmitochondrialDNA(mtDNA),whichindicatesthematernallineage(i.e.,itistransmittedfrommothertocalve)andisbecauseofitsabundanceinacellrelativelyeasytoanalyse.Onemilestonehasbeenreachedrecently:theresearchgroupofDavidMacHugh(UniversityCollegeofDublin)hasbeenabletosequencethecompletegenomeofaBritishaurochsfrom6700BPTheseresultshavebeenpublishedrecently3.Sograduallythecontoursofaurochsgenesareemerging.However,takenintoaccountthatoncetheremusthavebeenmillionsoftheseanimalsroamingforseveralhundredthousandsyearsaroundEurope,AsiaandNorthernAfrica,theremusthavebeenasubstantialgeneticdiversitywithinthepopulations.Withthegenomeofonlyoneindividualbeingavailableatthismoment,itmustbeclearthatwe’restillmissingalotofinformation.
Theaurochsgenomicsequencecanbeinterpretedonthebasisofourgrowingunderstandingoftheroleofgenesandgenevariantsofcattleindetermining
2Geneticjargonismarkedorangewhenfirstused.Ashortexplanationcanbefoundintheglossary.3Parketal.GenomeBiology(2015)16:234DOI10.1186/s13059-015-0790-2
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phenotypesassize,appearance(e.g.,coatcolour,shapeofhornsincattle)andadaptation.
SurveyoftheTaurosResearchProgramme
IntheTaurosProgammeanumberofthingshasbeentriedtodelvedeeperintoAurochsgenetics.In2009/2010partsofaurochsskeletonsfrommuseumsalloverEuropewerecollected,togetherwithgeneticmaterialofaround30differentEuropeancattlebreedswith‘primitive’features.DNAwasextractedandthegeneticvariationwasmeasuredat770.000sites(SNPs,seebelow)inthegenome.Thisyieldedmostvaluablenewinformationonthegeneticstructureofca.30primitiveEuropeanbreeds.WefoundthatthemtDNAfromaurochsandtaurinecattlearedifferent,andwecouldconstructafirstgenealogicaltree,sketchingthegeneticrelationbetweenthesebreedsandaurochs.Thereweredisappointmentstoo.DNAextractedfromaurochsbonesprovedtobeunsuitableforthegeneticanalysis,precludingforthemomentadirectcomparisonoftheaurochsandcattleautosomalDNA.ThisearlypartoftheresearchwasfundedandcarriedoutbyWageningenUniversity,aspartoftheirbroaderinterestinthegeneticstructureofproductivecattle-breeds.
In2012anotherattempttosequenceAurochsgeneticmaterialwasmade.PartoftheskinofamedievalspecimenofaPolishaurochswasanalysedbytheresearchgroupofBethShapirooftheUniversityofCalifornia,oneoftheforemostresearchersinancientDNA.Howeverthisleadtonoresult:theDNAprovedtobetoodeteriorated.Fortunately,in2012-2013,DavidMacHughagreedtogiveWageningenUniversityaccesstothe–atthatmomentstillunpublished-sequenceoftheaurochsgenomes,creatingnewopportunities.
InadditiontothegrantoftheLibertyWildlifeFundin2013,in2014theTaurusfoundationandWageningenUniversityweregrantedaEuropeanresearchbudgetofEUR240.000aspartoftheEGS-ABGprogramme.Sointheautumnof2014,aPhDstudenthasstartedonafour-yearprogrammeatWageningenUniversityincooperationwiththeUniversityofUppsala.Thefirstresultsarebeingpublishedearly2015.
AnextcontributiontoourdatasetwillbetheDNAsequenceofsixtotencompletegenomesoftheimportantparent-animalsofthebreedingoftheTaurosProgram.ThiswillbefundedbygrantofEUR10.000providedbytheDutchProvinceofGelderlandaspartoftheir‘Foodvalley’-programmeandanadditionalEUR10.000providedbyREaspartofthebudgetforthetransportofSayaguesaanimalsfromSpaintoCroatia.
Thecomingyearstheknowledgeontheaurochsandonprimitivecattlewillbebuiltupfurther.AdditionalspecimensofaurochswillbeanalysedbytheUniversityCollegeofDublinandDublinUniversity-andourunderstandingofthehistoryofdomesticationofcattleisgrowing(partofaresearchprojectoftheUniversityofLyon).
Becauseimportantgeneticinformationisstillmissing,wemodifiedourinitialchoices.Insteadoffocussingonjusttwoorthreebreedstostartwith,wechoseforawidervarietyofbreeds.Thisgivesflexibility,forinstanceifon-goingandfuturegeneticresearchleadstonewinsights.AdirectandmostusefulapplicationofthegeneticanalysiscarriedoutwiththefundingofLibertyWildlifeFundisthatwewillbeabletodetermineaccuratelythebreedcompositionoftheTaurosoffspringandweareabletocomparetheoffspringwithAurochs.
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2 Aurochs:ashortdescription
Archaeology,archaeozoology,writtenhistoricalsources,cavepaintings,knowledgeaboutcattlegenetics,cattlebreedingandcattlebehaviourhaveallcontributedtoourunderstandingoftheaurochsinthelastdecades.Weknowwhattheanimallookedlike,weknowitsecology,
Thelooks
Thesizeoftheaurochsbullsvariedbetween155cmand180cm,dependingonthelocalcircumstances.Thesizeofcowswasbetween135cmand155cm.Bullsweresignificantlylargerthancows.Thebodyshapeofanaurochswasdifferentfrommanydevelopedcattlebreedstoday.Ithadanathleticbodyshapeandlonglegs,builtforlong-distancewalking.Itwasaswiftandveryagileanimal.Theskull,carryingthelargehorns,waselongatedandbigcomparedtotherestofthebody.Bull’shornscouldreachupto107cminlengthandbetween10and18cmindiameterattheirbase.Thecowshadhornsprobablyreachingupto70cminlength.Thebull’shornswerelarger,withthecurvaturemorestronglypronouncedincows.Bothsexeshadadistinctlight-coloured,oftenwhite,muzzleandadistincteelstripe(especiallywithbulls).
Range,habitat,ecologyandbehaviour
Attheendofthelastglacialperiod(12,000BP)aurochswasendemicovermostofAsia,Europe,NorthAfricaandtheatthetimegreenSahara(figure2.1,fromFeliuset.al.,2014)
Figure2.1:ThepresumedmaximumareawhereAurochshistoricallyroamedaround(mapbyMarleenFelius,Feliuset.al.,2014)
AurochsoriginatedinIndiasome2millionyearsago.DuringthePleistoceneepoch(2,5millionto12,000yearsago)thespeciesspreadacrossotherpartsofAsia,alsointonorthernAfricaandEurope.TheearliestremainsofaurochsfoundinGermanydate
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fromabout400,000yearsago,butitisestimatedtheanimalreachedEuropelongbeforethat,probablyaround700,000yearsago.
Aurochshabitat:forestdwellerorkingofthesavannah?
Untilrecently,theprevailingtheorywasthataurochsmainlylivedinclosedcanopyforestsandforestedriverinehabitatsorswamps.Thisviewhasbeenadjustedbyecologicalevidenceandthestudiesonaurochsteeth,andskeletons,suchastherecentstudyofBakkeret.al.,20154.Althoughthere’sstillsomescientificdebate,theviewpointofRewildingEuropeandTaurusfoundationisthatAurochswasagrass-eatingplainanimal.Duringthe‘end-game’theanimalwasdriventoasuboptimalhabitat,themoreorlessclosedforests,becausemenoccupiedthefertileplains.Bytheirgrazingactivitiesaurochsopenedupthevegetationandcreatedbiodiversemosaicsofvegetation.
Aurochsbehaviour
Cattledon’trunawayfromthreats,butgrouptogetheranddefendthemselvesifneeded.Therefore,cattlearenotevenlydistributedoverthelandscape,butformfamilygroupsandsometimesroamaroundinlargergroups.Thisresultsinatemporaryandlocallyhighgrazingpressure,dependingontheseasonalfoodavailability.Thelargerthearea,thegreatertheireffectisondifferentiatingthelandscape.
Figure2.1:Preferredhabitatofaurochs
Undernaturalconditions,aurochsprobablywithdrewinsummertothecoolmountains,wheregrasswasmoreavailablethaninthehotlowlands,whilesnowandcoldforcedthemdownagainduringthewinter.
4www.pnas.org/cgi/doi/10.1073/pnas.1502545112
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Aurochsgenetics
Alotofinformationisavailableonthegeneticsofcattleandthevariabilitywithinandacrossbreeds.ThecompletegenomeofaHerefordcowisavailablesince20095,Onthegenomeabout22,000genescodingforproteinshavebeenidentified.Theroleofmanygenesarenowbeingdiscovered,withanemphasisontheeconomicallyimportantgenesformilkinggift,meatqualitye.g.
ThefirststudiesofancientAurochsDNAfocusedonmtDNA.In2010thecompletesequenceofAurochsmtDNAwasdescribed.BasedonarecentmtDNAstudyofourmoderncattlebreedsithasbeenestimatedaround80femalesfromtheFertileCrescentstandatthebasisofallournowadayscattle.RecentlythecompletegenomeofoneBritishAurochshasbeenanalysedpublished(Parket.al.,2015).
In2010forthefirsttimethecompletemtDNAsequenceofanaurochs(wasestablished(Edwardsetal2010).ThissamplebelongedtotheaurochshaplogroupP(derivedfrom‘primigenius’).InItalianAurochsanotherhaplogroup‘T’wasfound.Figure2.1showsthetreeofthecommoncattleTmtDNA(hereindicatedasBRS),theaurochsPmtDNAandthesporadicQandRmtDNA.
Figure2.3:AtreeofBovinehaplogroupsofmtDNA6
AncientDNAanalyses
Amongothermethods,suchasSNPgenotypingandwholegenomeanalyses,thecurrentstudyusesancientDNA-analysesasasourceofinformation.Box1givesanoverviewofhowthisisdone.
5Elsik,C.G.etal,(BovineGenomeSequencingandAnalysisConsortium)(2009)."Thegenomesequenceoftaurinecattle:awindowtoruminantbiologyandevolution".Science324(5926):522–528.doi:10.1126/science.1169588.PMC2943200.PMID19390049.6source:http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0015760
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Box1AncientDNA(fromMageeet.al.,2014)
ThestudyofancientDNAhasthepotentialtoanswerquestionsaboutthepastdirectly,thusprovidingatemporalaspecttomoderngeneticstudies.AncientstudieshaveshownthatDNAcansurviveintactformorethan500,000years,enablingmanyevolutionaryquestionstobeaddressedoverthistimeperiod,includingbiogeography,phylogeny,taxonomyandselectiveprocessesatthepopulationlevel,aswellasdomesticationorigins.AncientDNAcanbeextractedfromavarietyofanimalremains,ranginginconditionandage,themostpopularofwhicharebones,teeth,andhair.ThecytoplasmiclocationandhighcopynumberofmitochondrialchromosomeswithineacheukaryoticcellhasmademtDNAtheprimarygeneticmarkerforancientDNAstudies.However,recentadvancesingenomictechnologieshaveseenanincreaseinthenumberofancientDNAstudiesthatinvolvepartialorwholenucleargenomesequencesfromarchaeologicalsamples(ShapiroandHofreiter,2014).AnalyzingancientDNAisaveryspecializedtechnologyandrequiresdedicatedfacilitiestopreventcontaminationwithmodernDNA.
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3 Thedomesticationofcattle
OneofthemaingoalsoftheTaurosProgrammeisthede-domesticationofcattle.Thereforeitisimportanttosummarizewhatweknowonthedomesticationitself.
Earlydomestication:the‘covenantofthewild’
Untilrecentlymuchofourknowledgeondomesticationwasbasedonarchaeologicalexcavations,osteologicalanalysesandradiocarbondating.Nowadaysourknowledgeofearlydomesticationofcattlehasmadebigleapsbecauseoftheuseofmoderngenetics.
Wethinkthatafteralongperiodofco-evolutionofman,aurochs(andwolf)(co-)domesticationwasthenextstepinthisprocess.Domesticationledtoselectiveadvantagesforbothpartners:animalswereprovidedwithfoodandshelterhumanswithmilk,meatandworksupply.Domesticationhasbeendescribedasa“covenantofthewild”(Budiansky,1994inMageeet.al.,2014).Cattlewereprobablyoneofthemoredifficultspeciesfordomesticationbecauseoftheirsize,butmoreeasybecauseoftheirtendencytoholdgroundinsteadoffleeing(suchashorses).Theywereamongstthefirstwildgrazerstobedomesticated.
10,000BP:thestartofitall
Around10,750-10,250yearsBPthedomesticationofTaurinecattlestartedintheFertileCrescent(Mesopotamia).MolecularresearchhasindicatedthatthenumberofaurochsfirstdomesticatedintheMiddleEastwasverysmall,possiblyonly80individuals.Alongthedomesticationrouteswild-matingswithwildaurochsbullshavetakenplaceinAfricaandEurope.Thefirstselectiononcattle-traitswasdirectedonmanageablesizeanddocility.Asaresultofdomestication,cattlebegantodecreaseinsizethroughoutthemillenniathatfollowed.Cattleweremultipurposeandbredformeat,milksupplyandworkingpower.Bullswerecastrated.
Routesofdomestication
Domesticationprovedtobeaverysuccessfulinnovation.Thefirstfarmerssoonneedednewareasandtooktheiranimalsalongwiththem.ThisiswherethemythofthefoundingofEurope-ZeusdisguisedasabullwithEuropeonhisbackswimmingfromtheLibanesecoasttoCrete-meetsreality.Around10,000BPandfrom8,500BP,startingintheFertileCrescent,awestwardexpansionofagriculturalsocietiesbroughtthefirstdomestictaurinecattletocentralAnatoliaintoEurope7.AnotherroutewasalongtheNorth-AfricanMediterraneancoastandthenalongtheWest-AfricanAtlanticcoast.ThetypicalAfricanT1mtDNAhaplotypeisalsofoundinIberiancattle,anindicationofimmigrationofAfricancattleinEurope(Spain,Italy,Malta)acrosstheseastraits.ThemigrationfurtherintoEuropefollowedprobablytworoutes:viatheMediterraneancoastsandalongtheDanuberiver.Approximately7,500BPdomesticcattlereachedCentralEuropeviatheDanubianroute.NorthernEuropefollowedca.1,000yearslater.
7MainsourceFeliusetal2014
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DuringfurtherdomesticationinEuropeintrogressionofaurochs(bulls)mayverywellhavetakenplace(andinfacthastakenplaceasisproveninParket.al.,2015).Groupsofcattlewereherdedinremoteareasandwerenotprotectedbybarbedwire.
AccordingtoDeckeretal(2014),indicine(fromBosIndicus)influenceisfoundinthreePodolianbreeds(Chianina,RomagnolaandMarchigiana),whichprobablydescendfromHungariansteppebullsimportedduringthe14thtothe18thcentury.Lenstraet.al.(2014)findsindicineinfluenceinallPodolianbreeds.
Figure3.1:DomesticationroutesintoAfricaandEurope(fromFeliuset.al.,2014)
Bovinebreeds
Themigrationofcattletoallinhabitedcontinents,localadaptationandartificialselectionhasresultedintovarietiesofcattlebreedsweobservetoday.Sincethelast100-200years,manycattlebreedshavebeenintentionallymodifiedtoimproveandproducethetraitsbeneficialforhuman,resultingintohighoutputcattlebreedsspecializedforspecificpurposesTheprofitablespecializedcattlebreedshavereplacedseverallocalandoftenextremelywell-adaptedtraditionalbreeds.Asaresult,verymanycattlebreedsareatrisk,mostofthembeingtheprimitivecattlebreeds.Themechanizationofagriculturehasunderminedthemaintenanceofauthenticdraughtcattlewithprimitivegenevariants,leadingtoafurtherdriftfromthewell-adaptednaturalgrazer,theaurochs.AmajorconcernisthecrossbreedingoflocallyadaptedbullswithhighlyproductivebreedssuchastheLimousin.
ClassificationofEuropeanbreeds
Deckeretal.,(2014)givesaclassificationoffivemajorgroupsofbreedsandwithineachgroupseveralclusters.
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1. North-Europeancattle,comprisingeightbreedclusters.2. CentralEuropeancattle,withmanydualpurpose(beef-workordairy-work)and
triple-purposebreeds,comprisingthreefivebreedclusters:3. Iberiancattle,withalargevarietyofcoatcoloursandhornmorphologyandmainly
usedasbeef,workandfightingcattle;4. GreyPodoliancattle,themostlylong-hornedand,primarilydevelopedasbeefand
workanimals;5. BalkanandAnatolianbreed,agroupofgeneticallydiversebreedsstillrepresenting
theundevelopedtaurinecattle.
ThelonglistofbreedsfortheTaurosProgramme
IntheTaurosProgrammeafirstselectionwasmadefromthehundredsofbreedsstillpreservedinEurope,thelonglist.ThisselectionwasperformedbythescientificcommitteeoftheTaurosProgrammein2009andbaseduponspecificfeatures,history,purposeandmanagement-system.Basedonthesecriteria38cattle-breedswereselectedandalsousedinthegeneticanalyses,includingthebreeds((suchastheBoskarinandMalteseCattle)thatwereaddedlatertothislist.Annex1givesthetotallist.
Basedonmoredetailedknowledgeofthebreedsasub-selectionofthislonglistwasusedintheinitialstageoftheTaurosProgramme,theshortlist.ThesebreedsandthereasonsfortheirinitialselectionfortheTaurosProgrammearelistedinAnnex3.AbreedastheItalianChianinaforexamplehasbeenleftoutsincethePodolianMaremmanaprimitivawithlargerhorns,lessmeattypical,bettercolorationandamoreferalwayoflivingisthebetteralternative.
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4 Results
Researchquestions
IntheendtheTaurosProgrammeneedsanswersonthefollowingquestions:
§ Whatwasthegeneticconstitutionofandvariationwithinaurochs?§ Whichgenesincattlehavebeenchangedinresponsetodomestication?§ Whichgenesencodetheessentialaurochs-likefeatures?§ Whatisthegeneticconstitutionofthe(primitive)cattle-breedsbeingusedandhow
dotheyrelatetoeachotherandtotheaurochs?§ HowcanweusethisgeneticinformationintheactualTaurosbreedingprogramme?§ DoestheTaurosProgrammegeneticallymoveintherightdirection:doesthe
offspringapproachthegeneticallyaurochs?§ WilltheresultingTaurospopulationshaveenoughgeneticvariation?
Toprovidetheseanswers(atleastthebeginningoftheanswers)WageningenUniversitycarriesoutthegeneticresearch,inclosecooperationwithotherscientificinstitutes.Annex4givesanoverviewofthedifferentresearch-stepstakensofar.Theamountofresearchhasbeenlimitedbytheavailablebudget.Intheperiod2010-2015chronologicallythefollowingstepshavebeentakenaspartofthegeneticresearchspecificallyorientedtowardtheTaurosProgramme.TheresearchspecificallyfundedbytheLibertyWildlifefundwasstartedin2013andconcludedin2015:
§ Analysesof100crossbredsfromtheTaurosprogramme(inheritancecheck);§ 770kSNPanalysesofadditionalbreeds(24individuals);
ThisresearchismostrelevanttotheTaurosProgramme,sincewithouttheinheritancecheckandthefurtheranalysisoftheoffspringgenerations,wewillnotbeabletocheckiftheprogrammemovesintherightdirection.
Methodsfollowed
Anumberofdifferentmethodsofanalysishavebeencarriedout:ingeneralquitespecialistgenetictoolsandmethods,hardlyunderstandablefornon-geneticists.Thereforethemethodsarenotdescribedinthesummary.
Results
Theresultsofthedifferentanalysesarerepresentedinthreedifferentways.InMauliketal(2015)8aditionalanalyseshavebeencarriedoutanditsresultspresented,howeverthatdoesnotchangethegeneralconclusions.
1. CalculationoftheNeigeneticdistance;2. PopulationstructureanalysisbyModelbasedclustering;3. Parentagecheck.
NEIGeneticdistance:Rankingthestars
8Mauliket.al.,2015.Geneticorigin,admixtureandpopulationhistoryofprimitiveEuropeancattleandAurochs
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NeiGeneticdistanceestimatesthedifferencebetweenbreeds,akindof‘rankingthestars’.NeiGeneticdistanceshowshowclosedifferentbreedsaretothe(oneBritish)aurochs.Table4.1givestheresults.
TABLE4.1:GENETICDISTANCETOAUROCHS
Ranking NEIDistance Code Breed Country 4.1.1 0 4.1.2 AU4.1.3 Aurochs 4.1.4 Britain1 0,1245 PA Pajuna Spain2 0,1249 BU Busha Balkanregion3 0,1307 CA Cardena Spain4 0,1312 SA Sayaguesa Spain5 0,1365 FL Fleckvieh Switzerland6 0,1374 BN Berrendaennegro Spain7 0,1377 LM Limia Spain8 0,1385 BC BerrendaenColorado Spain9 0,1386 LI Lidia Spain
4.1.5 10 4.1.6 0,13904.1.7 MA4.1.8 Maremmana 4.1.9 Italy11 0,1391 AR Arouquesa Portugal12 0,1412 ME Maronesa Portugal13 0,1443 BS Brownswiss Switzerland14 0,1457 HL Highlander Scotland15 0,1465 JE Jersey JerseyIsland16 0,1465 KC Kerrycattle Ireland17 0,1481 GA Galloway Britain18 0,1486 HE Heck -19 0,1515 CH Chianina Italy20 0,1520 RO Romeniangrey Romania21 0,1533 WP Whitepark Britain22 0,1536 BK Boskarin Croatia23 0,1548 MN Marondesa Spain24 0,1553 CC Cachea Portugal25 0,1563 AL Alentejana Portugal26 0,1582 TU Tudanca Spain27 0,1587 DB Dutchbelted TheNetherlands28 0,1600 MT Maltese Malta29 0,1628 EL EnglishLonghorn Britain30 0,1638 MI Mirandesa Portugal31 0,1923 AN Angler Germany32 0,1925 CL Caldela Portugal34 0,1978 PO Podolica Italy35 0,2239 NE Nelore India
Table4.1showsthatfromthe34primitivebreedsanalysedsofarthefourmostfrequentlyusedbreeds(marked)intheTaurosProgrammeareinthetop10.Thisisaremarkableresult,sincebefore2015thechoiceofbreedswasmadeonthebasisofthephenotypeandthehistoryofabreed.Especiallythenr1rankingofthePajunabreedisencouraging.Withregardtotheotherbreedsinthetopten:thereweresoundphenotypicalreasonsnotwantingtousethemintheinitialphase,presentedherebelow.Howeverthisdoesnotmeanthatthosebreedswillnotbeusedatall,because
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apparentlytheypossessgenesthatratethemhighintherankingcomparedtotheonespecimenofAurochs.
Modelbasedclustering
Modelbasedclusteringgivesinsightintheancestry.Figure4.1showstheresultofthecalculations,assumingthreedifferentclusters:zebu(blue)northEuropeancattle(green)andsouth/Europeancattle.Theaurochsspecimen(therightmostcolumnoffigure4.1)containselementsfromallthreeclusters.ThepatternsuggestszebuinfluenceintheEast-European(Podolian,Busha)butnotintheIberianclusters.Fromthedomesticationhistorythismakessense,sincetheEast-EuropeancattlehasbeenincontactwithAnatoliancattle,whichunderwentalreadyzebuinfluencesince5000BP.
Parentagecheck
SinceTauros-herdsoftenhavemorematingbulls,it’simportanttohaveagoodparentagecheck.Theparentageofthefirst100offspringoftheTaurosProgrammehasbeencheckedwitha40kSNParray.The40kanalysesinawayareanoverkillfortheparentagecheck,butthesamedatawillbeusefultocheckthesegregationofspecifictraits,onceweknowforwhichspecificgenesandtraitswehavetolook.
TheresultsofthehereditycheckarepresentedinAnnex5.From126outofthe130sampledindividualsitwaspossibletopositivelydeterminetheidentityofatleastoneoftheparents.Inmostcasesthemotherwasalreadyknownfromobservationsinthefield,butinothercasesmaterialofthemotherwasnotavailableanymore.Mostfatherswereknownasdonorsofartificialinseminations,butseveralfathersofimportantanimalshavenotbeensampledandremain(asyet)unknown.OncewewillhavesampledmoreindividualswewillbeabletopositivelyidentifytheparentageofallTaurosoffspring.
TheparentagetestrevealedthatintheherdofKeentnearlyallmatingswerebytheMaremannaxPajunabull(ManoloUno,nrNL896492389)andthatasecondbullinline–nrNL896492390,oneyearyounger–didnotperform.IntheotherlargeareaintheNetherlands(Kempen-Broek)thediversityofreproducingbullsislarger,becausetheareaissplitintoseveralsmallerareaseachwithadifferentbullandifavailablemorebullsinonegroup.
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Figure4.1:AncestryassignmentofalltheindividualswithK=39
9ProbablytheWhiteParkCattleandRomanianGreysampleshavebeenswitchedinthisfigure.ThiswillbecorrectedinthefinalpaperMauliket.al.,2015.
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5 ThemeaningofgeneticresearchfortheTaurosProgramme
ThegeneticresearchisoneofthepillarsoftheTaurosProgramme.Ourinitialideaswereprettystraightforward:sampleafairnumberofAurochsindividualsandcomparethesewiththebreedsyouwanttouseandalreadyareusing,andchoosethemostaurochs-likeanimals.Ofcoursetheroadappearedbumpierthanweexpectedwithamajorset-backourfailureofanalysingaurochsmaterial.Thisapproachinitselfwasnotunrealistichowever.
Meaningoftheresults
Forthefirsttimeagenealogicaltableofaurochsandprimitivecattlebreeds,basedon770KSNPautosomalDNA,hasbeenconstructed(table4.1).Althoughstillpreliminary,itisencouragingthattheoutcomeoftheresearchconfirmsourbreedingprogrammesofar.Itroughlyconfirmsthatfourofthebreedsthatdoplayamajorroleinourbreedingprogrammeareinthetop10andsixofourbreedsareinthetop15.Weexpecttobeabletodrawmoredefiniteconclusionswhenmorespecimensofaurochswillhavebeenanalysed.
ThereareotherprimitivebreedswedidnotconsideryetsuchasUkranianGrey,HungarianSteppecattle,ortheoffspringofelderIberiancattlebreedsinSouthAmerica(Criollo).Especiallythelattercouldbeverypromisingsince,someoftheseherdsstemdirectlyfromoldIberianbreedswithouttoomuchexternalinfluence.
Thespeedofourresearchwasn’tashighaswehopedforinitially.ButtheinitialslowprogressmeantthatwewereabletoincludenewinsightssuchasFeliuset.al.(2014)andLenstraet.al.(2015).Theyprovidedthebigpictureofcattledomesticationandthespreadofgenes.Thisdetailedpicturewasearliernotavailable.Wecannowconcludethatourresultsareinlinewiththosetwopublications.
ThestrategyofcombiningIberianandPodolianprimitivebreedsisinlinewiththeoutcomeofthegeneticresearch.Bythisstrategytworoutesofcattle-domesticationandtwodifferentresourcesof‘primitive’genesarecombined.TheIberianbreedshaveretainedmuchgeneticdiversity,whichwasincreasedbytheinfluenceofAfricancattlewiththeAfricanaurochsancestry.Incontrast,PodolianbreedshaveadifferentoriginintheBalkansandmayverywellhavepickedupgenevariantsofEuropeanaurochs.
Nextsteps
Obviously,wewillhavetocontinueoutinvestigationsandalsocontinueourinternationalcollaborations.Ourplansarethefollowing:
§ Attheshortterm,aphenotypicaldescriptionofindividualsaswellasmorepracticalguidelinesforselection(basedonthegross-listofbreedingcriteria).Nowthatthefirst100offspringhavebeenanalysed,weneedtodescribethemostessentialfeatureoftheanimalsaswell.Wewillnowworkoutapracticalprogrammetomakemeasurementsatindividualanimals.Thiswillenableustopredictthedevelopmentofanimalsinanearlystageandtospeeduptheselectionprocess;
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§ Wewillcarryoutwholegenomesequencingofapproximately10Taurosfounderanimals;
§ Mostessentialwillbegenotypesofotheraurochsindividualstoassessthegeneticvariationwithinaurochs.ThisisbeingworkedonbytheresearchgroupsofMacHughandBradleyandalsoontheUniversityofLyon;
§ Onthelongtermweneedtohavemoreinsightinidentityofthegenesforrelevanttraits,sothatwewillbeabletopredicttoacertainextentwhatthedevelopmentofayoungindividualwillbe.ThisanalysiswillbeperformedbythepHDinhisnextyears.
SincetheTaurusFoundationhasnottherequiredresources,additionalbudgetswillbenecessarytocarryoutthisresearchandtosetpriorities.
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Glossary• Alleles:AlternativeformsofaDNAsequence• Autosomalchromosomes:AllchromosomesexcepttheXandYsexchromosomes.• CNV:Copynumbervariation.AcertainDNAsegmentcancontainmorecopiesofa
gene,eachcopynotnecessarilybeingaperfectcopy.Thenumberofcopiescandependontheindividualandthusisapolymorphism:theCNV.
• Chromosome:Alongdouble-strandedandfoldedDNAmoleculethatcontainsgeneticinformationandispassedtonextgeneration
• Diploid:Anorganismwithtwocopiesofeachchromosome,onefromeachparent.• DNA:Deoxyribonucleicacid;themoleculethatcarriesgeneticinformation• Dominant:Aphenotypeoritscorrespondingallelethatinaheterozygotemasksthe
presenceofthealternative,recessivephenotypeofthesamelocusintheheterozygote;
• F1generationFirst-generationoffspringofacross;• F2generationSecond-generationoffspringofacross;• Gene:Thefundamentalunitofheredity;aspecificandfunctionalsectionofDNA
withinachromosomethatcodesforaproteinorRNAmolecule• Geneticdistance:Astatistictoolusedtomeasurethedifferencesinallele
frequenciesbetweenspeciesorbreeds.Thesmallerthedistancethemoretwobreedsorspeciesarealike.AstandardtoolingeneticsisNEIgeneticsdistance.
• Genotype:Thegeneticmakeupofanindividual;theallelecombinationatagivenlocus
• Haplotype:Thecombinationofallelesinachromosome,geneormtDNAmoleculeiscalledahaplotype
• Haplogroup:Anumberofsimilarhaplotypesiscalledahaplogroup.Haplotypesandhaplogroupswithinspeciesoftenhaveageographicdistributionindicatinghistoriccolonization-routes
• Heterozygote:Anindividualwithtwodifferentallelesatagivengeneorlocus• Homozygote:Anindividualwithtwoidenticalallelesofagivengeneorlocus• Locus:Aspecificlocationonachromosome• mtDNA:MitochondrialDNAisoutsidethenucleus.Itdoesnottakepartinthesexual
reproduction,butistransmittedfrommothertooffspring.• mRNA:MessengerRNA(mRNA)directsthesynthesisofproteins• Phenotype:Thephysicalcharacteristicsofanindividual• Recessive:Aphenotypeexhibitedonlywhenhomozygousortheallelecorresponding
tothisphenotype.• Sexchromosomes:Chromosomescontrollingthegender.InmammalsthesearetheX
andYchromosomeswithtwoXchromosomes(XX)infemalesandoneXandoneY(XY)inmales.
• Sexlinked:phenotypecontrolledbyageneononeofthesexchromosomes
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Literature• BollonginoR.,J.Elsner,J.D.VigneandJ.Burger,2008:Y-SNPsdonotindicate
hybridisationbetweenEuropeanaurochsanddomesticcattle.PLoSONE2008,3(10).
• E.Wright,S.Viner-Daniels,2015.GeographicalvariationinthesizeandshapeoftheEuropeanaurochs(Bosprimigenius).JournalofArchaeologicalScience54(2015).
• EdwardsC.J.,D.A.Magee,S.D.E.Park,P.A.McGettigan,A.J.Lohan,A.Murphy,E.K.Finlay,B.Shapiro,A.T.Chamberlain,M.B.Richards,D.G.Bradley,B.J.Loftus,D.E.MacHugh,2011.ACompleteMitochondrialGenomeSequencefromaMesolithicWildAurochs(Bosprimigenius).PLoSONE5(2):e9255.
• Edwards,C.J,D.A.Magee.,S.D.E.Park,P.A.McGettigan,A.J.Lohan,2010.ACompleteMitochondrialGenomeSequencefromaMesolithicWildAurochs(Bosprimigenius).PLoSONE5(2):e9255.
• FeliusM.,M.L.Beerling,D.S.Buchanan,B.Theunissen,P.A.Koolmees,J.A.Lenstra5,2014.OntheHistoryofCattleGeneticResources.Diversity2014,6,705-750;
• Felius.M.,CattleBreeds:AnEncyclopedia,Doetinchem,1995.• GoderieR.,W.Helmer,H.Kerkdijk-Otten,S.Widstrand,2013.Borntobewild.
ThecomebackofaEuropeanicon.RoodbontPublishers,Zutphen.• LariM.,E.Rizzi,S.Mona,G.Corti,G.Catalano,K.Chen,C.Vernesi,G.Larson,P.
Boscato,G.DeBellis,A.Cooper,D.Caramelli,andG.Bertorelle,2011.TheCompleteMitochondrialGenomeofan11,450-year-oldAurochsen(Bosprimigenius)fromCentralItaly.BMCEvolutionaryBiology2011,11:32
• LenstraJ.A.,P.Ajmone-Marsan,A.Beja-Pereira,R.Bollongino,D.G.Bradley,L.Colli,A.deGaetano,C.J.Edwards,M.Felius,L.Ferretti,C.Ginja,P.Hristov,J.Kantanen,J.PedroLirón,D.A.Magee,R.NegriniandG.A.Radoslavov,2014.Meta-AnalysisofMitochondrialDNARevealsSeveralPopulationBottlenecksduringWorldwideMigrationsofCattle.Diversity2014,6,178-187.
• Lynch,A.H.,Hamilton,J.&Hedges,R.E.M.(2008).Wherethewildthingsare:aurochsandcattleinEngland.Antiquity82(318):1025-1039
• MageeD.A.,D.E.MacHughandCeiridwenJ.Edwards,2014.Interrogationofmodernandancientgenomesrevealsthecomplexdomestichistoryofcattle.AnimalFrontiers,Volume4,Issue3Pages7-22.
• Mauliket.al.,2015(inprep).• McInerney,Jeremy,(2010).‘TheCattleoftheSun.CowsandCultureinthe
WorldoftheAncientGreeks’.PrincetonUniversityPress.• Perez-PardalL.,L.J.Royo,A.Beja-Pereira,S.Chen,R.J.C.Cantet,A.Traore,I.
Curik,J.Sölkner,R.Bozzi,I.Fernandez,I.Alvarez,J.P.Gutierrez,E.Gomez,F.A.PoncedeLeonandF.Goyache,2009.MultiplepaternaloriginsofdomesticcattlerevealedbyY-specificinterspersedmultilocusmicrosatellites.Heredity(2010)105,511–519.
• Shapiro,B.,andM.Hofreiter.2014.Apaleogenomicperspectiveonevolutionandgenefunction:NewinsightsfromancientDNA.Science343:1236573.
• StefanoMona,G.Catalano,M.Lari,G.Larson,P.Boscato,A.Casoli,L.Sineo,C.DiPatti,E.Pecchioli,D.Caramelli,G.Bertorelle,2010.Populationdynamicofthe
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extinctEuropeanaurochs:geneticevidenceofanorth-southdifferentiationpatternandnoevidenceofpost-glacialexpansion.BMCEvolutionaryBiology10:83,2010.
• Vera,F.W.M.,1997.‘MetaphorsfortheWilderness:Oak,Hazel,CattleandHorse.’Ph.D.diss.,AgriculturalUniversity,Wageningen,TheNetherlands.
• Vuure,C.van,2005.RetracingtheAurochs:History,MorphologyandEcologyofanExtinctWildOx.PensoftPublishers.Sofia-Moscow,2005.
• Wright,E.(2013).ThehistoryoftheEuropeanaurochs(Bosprimigenius)fromtheMiddlePleistocenetoitsextinction:anarchaeologicalinvestigationofitsevolution,morphologicalvariabilityandresponsetohumanexploitation.Thesis.DepartmentofArchaeology,UniversityofSheffield.
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Annex1:LONGLISTOF‘ORIGINAL’BREEDS
Breed Code Country RegionAlentejana AL Portugal Iberian
Angler AN Germany Dutch
Arouquesa AR Portugal Iberian
Aurochs AU Britain BosprimigeniusBerrendaenColorado BC Spain IberianBerrendaennegro BN Spain Iberian
Boskarin BK Croatia BalkanandItalyBrownSwiss BS Switzerland Alps
Busha BU Balkanregion BalkanandItalyCachea CC Portugal Iberian
Caldela CL Portugal Iberian
Cardena CA Spain Iberian
Chianina CH Italy BalkanandItaly
EnglishLonghorn EL Britain English
Fleckvieh FL Switzerland AlpsGalloway GA Britain English
Heck HE Germany Heck
Jersey JE JerseyIsland Jersey
Kerrycattle KC Ireland English
Lidia LI Spain Iberian
Limia LM Spain Iberian
Maremmana MA Italy BalkanandItalyMarondesa MN Spain Iberian
Mirandesa MI Portugal Iberian
MT Maltese Malta BalkanandItalyNelore NE India BosIndicus
Pajuna PA Spain Iberian
Podolica PO Ukraine BalkanandItaly
Romeniangrey RO Romania BalkanandItaly
Sayaguesa SA Spain Iberian
ScottishHighlander HL Scoland English
Tudanca TU Spain Iberian
Whitepark WP Britain English
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Annex2:SetwithcriteriafortheTaurosProgrammeCharacteristics Bull Cow
Shoulderheight 160to185cm 145to150cm
Overallbuild • Narrow/slender• Forehand:dorsalprotrusionsofthevertebrae,attachedtothebackoftheskull.Thisisneededforthemusclestocarryalargeheadwithhorns
• S-shapedback/spinecurvatureasaresultofdorsalprotrusionsoftheshouldervertebraetoconnectneckmuscles
• Squarebuildregardingtrunklengthandleglength.
• Narrow/slender• • • • • S-shapedback/spinecurvature• • • Squarebuildregardingtrunklengthandleg
lengthHindquarters • Slender
• Straightlegs• Hipjoint:sloping
• Slender• Straightlegs• Hipjoint:sloping(tofacilitatenaturalcalvingease)
Muscling • Nodoublemuscling• Musclescanbesomewhatpronounced,butfunctional
• Neckismuscled• Notypicalmeatcattleconformation
• Nodoublemuscling• Musclescanbesomewhatpronounced,butfunctional
Coloration • Calvesarered• Colorationchangestoblack/darkbrownafterafewtosixmonths
• Selectionagainsttoolightlycoloredindividuals
• Lightersaddleontheback(shadingingeneral)ispermitted
• Lighteelstripe• Whiteorlightmuzzle(cangetlessdistinctwhenanimalisolderthan2years)
• Darknose,nopinknosecolorpermitted
• Calvesarered• Redtobrownishwhenmature• Selectionagainsttoodarkortoolightlycoloredindividuals;Cowsshouldbelightercoloredthanbulls
• Lightersaddleontheback(shadingingeneral)ispermitted
• Canhavealighteelstripe,butthisisnotmandatory
• Whiteorlightmuzzle• Darknose,nopinknosecolorpermitted
Legs • Comparativelylonglegs.• Squarebuildregardingtrunklengthandleglength.
• Comparativelylonglegs.• Squarebuildregardingtrunklengthandleglength.
Head • Longandslender.• Bigincomparisonwiththerestofthebody.
• Longandslender.• Bigincomparisonwiththerestofthebody.
Horns • Length:60to107cm• Circumferenceonthebase:10to18cm• Form:variable,butonaverageasfollows:inthebeginninggrowingsomewhatbackwardsandupwardsinanangleof0to30degrees,thenincurvingforwardandinwardsinanangleof60degreesinrelationtotheskullandfinallyupwardsandoutwards.
• Length:40to70cm• Circumferenceonthebase:7to10cm• Form:variable,butonaverageasfollows:inthebeginninggrowingsomewhatbackwardsandupwardsinanangleof0to30degrees,thenincurvingforwardandinwardsinanangleof60degreesinrelationtotheskullandfinallyupwardsandoutwards.
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Characteristics Bull Cow
Coat • Singleshortcoatinsummer• Doublelongercoatinwinter• Fastsheddingofwintercoatinspring
• Singleshortcoatinsummer• Doublelongercoatinwinter• Fastsheddingofwintercoatinspring
Dewlap NotoverlypresentlikeinZebu NotoverlypresentlikeinZebu
Hump(neck) NohumplikeinZebu,exceptforfunctionaldorsalprotrusionsofthevertebraetoconnectneckmuscles.
NotpresentlikeinZebu
Udder Small,hairyandhardlyvisible
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Annex3:BriefsummaryofthebreedsinvolvedMaremmana‘Primitiva’(centralItaly)
TheMaremmanabreedcomesfromMaremma,aprairieandcoastalduneregioninwesterncentralItaly.Maremmanacattleareadaptedtowetandmarshyareas,butdowellinmountainsaswell.TheMaremmanaprimitivahasstrongaurochsfeatureslikethesize,coloursetting,thickhorns,cleardifferenceincolourbetweenmalesandfemalesandanaturalherdstructure.Long-legged,largecattle,withbullsreachingupto170cmwitherheight,swung-backlineinbulls,causedbythestronglyexpressedshoulderarea,extremelyhardy.Maremmanaprimitivaareextremelyhardycattleandcansurvivewellonpoorforage.InItaly,theMaremmanaliveinnaturalsemi-feralherds.
Pajuna(southernSpain)
ThePajunaisararecattlebreedfromAndalucíainSouthernSpain,withalong-leggedathleticbody,allowingthemtomovearoundtheterrainwithgreatease.Pajunabullscanreachupto165cmshoulderheight.Thehorns,especiallyofthebulls,arefacingforwardsandhavealightcolourwithadarktip.Bullsaredarkbrowntoblackwithalighteelstripe,whilethecowsarereddish-brownwithalongface.Theyareabletosurviveunderharshconditionsandonpoorforage.HighintheSpanishSierraNevadatheylivemostlyfromthornyshrubs.Andinthelowlandvalleystheyareknowntosurviveonsemi-desert-likevegetation,ifneeded.Thebreedadaptseasilytochangingcircumstances,whichisalsoanancienttrait.InAndalucíatheyarefrequentlykeptinlargerareas,withanaturalherdstructure.
Limia(north-westernSpain)
TheLimiacattlearefromGalícia,innorthwesternSpainandarealarge-sizedbreedaswell.Limiacattlealsohavecleardifferenceincoatcolourandshoulderheightbetweenmalesandfemales.Withtheirslenderbodiestheymovewithgreatease,andtheyareabletosurviveonpoorforage.Galíciaisrelativelycoldandwet,sotheanimalsareusedtolivinginawetcountry,withcoldandsnowywinters,likeinthemorenorthernpartsofEurope.
Sayaguesa(north-centralSpain)
TheSayaguesaorZamoranaisanoldcattlebreedfromSayagointheprovinceofZamorawestofSalamancainSpain.Traditionally,Sayaguesaswereusedforpullingploughs,cartsandwagons.It’salarge-sizedbreed,withbullsweighingonaveragebetween1000and1100kgandcowsbetween650and700kg.Thebullsareblackandsometimesstillhavealighteelstripe.Thecowsoftenarejustslightlylighterthanthebulls,sincefarmersselectonblackcolour,butreddish-brownSayaguesacowsstillcanbefound.Theyalsohavethewhitecolouredmuzzleoftheaurochs.Thehornsareaurochs-likeandpointforward,especiallyinthebulls.Sayaguesaisalong-leggedbreedwithacurvedbacklineandalongface.Thebreedisknowntobehardyandrobustand
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doesn’tneedanyshelterorsupplementaryfeedingintheSpanishdehesa-countryside.Theirbehaviouriswatchfulandshy,andthecowswillstronglydefendtheircalves.
Maronesa(northernPortugal)
MaronesaisahardybreedfromtheTrás-os-MontesregioninnorthernPortugalamongwolves.Somelineagesareveryaurochs-like.Thebullsaremuchlargerthanthecows,withadarkbrowntoblackcoatcolourandalighteelstripe,alightlycolouredmuzzleandverylargehorns.Maronesacowsarereddish-brown.Theheadsofbullsarelargeandtheyarewellmuscledwithacurvedbacklinefromthestronglypronouncedshoulderpart.Thehornsarethickandpointforward.Maronesasareusedtoharshconditionswithsnowywintersandhotsummersandabletolivewithoutanyhumanmanagement,withcalvesgrowingupsuccessfullyalsowithwolvesaround.
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Annex4:OverviewofconductedresearchTheamountofresearchhasbeenlimitedbytheavailablebudget.Intheperiod2010-2015chronologicallythefollowingstepshavebeentakenaspartofthegeneticresearchspecificallyorientedtowardtheTaurosProgramme.
1. In2010,aspartofandfundedbyWageningenUniversity,departmentAnimalBreedingandGenetics(ABG)thefollowingresearchhasbeendone:§ Thecollectionandgeneticanalysisof30originalbreeds(togetherwith
commerciallyinterestingbreedssuchasHolstein-FrisianandMRY)withthe770KSNParraytechnology
§ Thecollectionandanalysiswiththe770KSNParrayofaround25differentmuseumspecimensofaurochs;
2. In2012wetriedviaCaliforniaUniversitytoobtainacompletegenomesequencefromamedievalPolishaurochs(fundedbyTaurusFoundation);
3. In2013-2015additionalresearchcarriedoutwiththefundingoftheLibertywildlifefundhasbeendone:§ Analysesof100crossbredsfromtheTaurosprogramme(inheritancecheck);§ 770kSNPanalysesofadditionalbreeds(24individuals);Althoughthispartoftheresearchisfinanciallynotthemostsubstantial,itismostrelevanttotheTaurosProgramme,sincewithouttheinheritancecheckandthefurtheranalysisoftheoffspringgenerations,wewillnotbeabletocheckiftheProgrammemovesintherightdirection;
4. In2014-2015thecompletegenomesofanumberofimportantanimalsatthebaseoftheTaurosProgrammewillbesequenced.ThispartoftheresearchhasbeenmadepossiblebythefundingbytheFood-ValleyprogrammeoftheProvinceofGelderlandwith50%co-financingoftheTaurusfoundation;
Overtheperiod2014-2018furtherresearchandanalyseswilltakeplace,whichwasmadepossiblebyEuropeanfundingoftheEGS-ABGprogramme(PhDstudent)andwillultimatelyleadtopracticalrecommendationsforthebreedingprogramme;
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Annex5:TheresultsoftheinheritancecheckResultsoftheinheritancecheck.Insomecasesnotenoughgeneticmaterialwasavailable,inthatcasethecellinthistableisablank.Thiswillbesolvedinlateranalyses.
rund moeder vader
1 545195012 376909631 53990031794
2 545195454 53990019996 53990031794
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9 925696414 2110745436 896492389
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13 925696856 539900199995 896492389
14 925696902 71107310549 896492389
15 925696940 53990060204 896492389
16 925697082 896492341 50106187461
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19 529492238 440854133 53990031819
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21 545195005 533491768 53990031794
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23 709292298 323445106 53990031794
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25 719994582 522603112
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29 745492858 524990573 53990031794
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32 925696337 60807600739
33 925696438 414107057
34 925696584 713403378 90105426310
35 925696894 323445106 896492390
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36 925696926 50807262136
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53 545195245 709292274 90807606401
54 545195461 479830100 53990052783
55 545195562 353842274 896492389
56 545195788 370304537 896492389
57 545195795 399720891 53990052783
58 545196008 883794304 53990052783
59 545196008 61105912075 53990052783
60 545196015 284949088 53990052783
61 545196022 472356731 53990052783
62 655303439 21107310624 53990052783
63 713403417 80807262457
64 721393342 53990019995 limia-lacon
65 743694849 71107310549 31104103451
66 743694856 470099997 896492389
67 743694902 2110745436 31104103451
68 743694971 21107310759 31104103451
69 743694995 379111246 896492389
70 807502790 60807262148
71 883794304 61105912075 53990052783
72 887603509 425202502
73 896492341 376909631 53990031794
74 896492411 370304520 53990031794
75 925696320 896492389
76 925696344 479689812 896492389
Summaryof:AurochsGenetics,acornerstoneofbiodiversity
page32
77 925696351 527791195 896492389
78 925696382 379111260 896492389
79 925696399 896492389
80 925696421 497390343 896492389
81 925696445 379111246 896492389
82 925696452 745492997 896492389
83 925696469 470099911 896492389
84 925696577 738993793 90105426310
85 925696623 53990063275 90105426310
86 925696700 83148 90105426310
87 925696779 470099997 896492389
88 925696786 479689766 896492389
89 925696793 433089850 896492389
90 925696801 339089693 896492389
91 925696870 546492510 896492389
92 925696919 525590963 896492389
93 925696964 53990019996 90105426310
94 925696971 745492740 896492389
95 925696995 370304520 896492389
96 925697006 370304537 896492389
97 925697013 496590168 896492389
98 925697037 284418052 896492389
99 925697044 527791311 896492389
100 925697099 745492841 50106187461
101 925697121 896492341 50106187461
102 925697152 533491814 50106187461
103 925697152 896492411 50106187461
104 925697169 526991390 896492389
105 925697176 241297191 896492389
106 925697121 896492341 50106187461
107 925697152 533491814 50106187461
108 925697152 896492411 50106187461
109 925697169 526991390 896492389
110 925697176 241297191 896492389
111 20807502792 30807606416
112 21107310624 91105237441 9716
113 21107310624 91105237441 9718
114 21107310624 91105237441 9814
115 30809822607 30807606416
116 31104103451 81104103138 limon
117 61105912075 61103323901 lacon
Summaryof:AurochsGenetics,acornerstoneofbiodiversity
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118 90104784771 90104054854 PAJ23
119 53990052783 11GR148C007 9716
120 53990052783 11GR148C007 9718
121 53990052783 11GR148C007 9814
122 527791126 370304520 225690381
123 545194824 53990060204 53990031794
124 545194949 377628465 53990031794
125 545195601 284418076 53990052783
126 883794506 379111246 53990031794
127 887603561 425202502 Julio
128 896492396 241297191 53990031819
129 925697114 896492411 50106187461
130 60807262148 50807618816