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Impacts of Removing Badgers on Localised Counts of Hedgehogs

IainD.Trewby

, RichardYoung, RobbieA.McDonald, GavinJ.Wilson, JohnDavison, NeilWalker, AndrewRobertson,

C.PatrickDoncaster, RichardJ.Delahay
Published:April15,2014

DOI:10.1371/journal.pone.0095477

Abstract

Experimentalevidenceoftheinteractionsamongmammalianpredatorsthateatorcompetewithoneanotherisrare,duetothe
ethicalandlogisticalchallengesofmanagingwildpopulationsinacontrolledandreplicatedway.Here,wereportonthe
opportunisticuseofareplicatedandcontrolledcullingexperiment(theRandomisedBadgerCullingTrial)toinvestigatethe
relationshipbetweentwosympatricpredators:EuropeanbadgersMelesmelesandwesternEuropeanhedgehogsErinaceus
europaeus.Inareasofpreferredhabitat(amenitygrassland),countsofhedgehogsmorethandoubledovera5yearperiodfrom
thestartofbadgerculling(from0.9ha1precullto2.4ha1postcull),whereashedgehogcountsdidnotchangewheretherewas
nobadgerculling(0.30.3hedgehogsha1).Thistrialprovidesexperimentalevidenceformesopredatorreleaseasanoutcomeof
managementofatoppredator.
Citation:TrewbyID,YoungR,McDonaldRA,WilsonGJ,DavisonJ,WalkerN,etal.(2014)ImpactsofRemovingBadgerson
LocalisedCountsofHedgehogs.PLoSONE9(4):e95477.doi:10.1371/journal.pone.0095477
Editor:FranoisCriscuolo,CNRS,France
Received:November27,2013Accepted:March27,2014Published:April15,2014
Copyright:2014Trewbyetal.ThisisanopenaccessarticledistributedunderthetermsoftheCreativeCommons
AttributionLicense,whichpermitsunrestricteduse,distribution,andreproductioninanymedium,providedtheoriginalauthor
andsourcearecredited.
Funding:ThestudywasfundedbytheUnitedKingdomGovernmentsDepartmentforEnvironment,FoodandRuralAffairs
(http://www.defra.gov.uk).Thefundershadnoroleinstudydesign,datacollectionandanalysis,decisiontopublish,or
preparationofthemanuscript.
Competinginterests:Theauthorshavedeclaredthatnocompetinginterestsexist.
Introduction

Toppredatorsmayhavefarreachingimpactsontheecosystemstheyinhabit[1],[2].Asaconsequence,anthropogenicactivities
whichreduceorremovetoppredatorspeciesmayhavemajorandoftenunintentionaleffectsonthestructure,productivityor
diversityofthewiderecosystem[1],[3].Inparticular,toppredatorsmaysuppresssmallermesopredators,eitherbydirectintraguild
killingorpredationorviachangesinbehavioursuchthatthesmallerpredatorsavoidlocationsorhabitatsutilisedbythetop
predator[1],[4].Adeclineintheabundanceofatoppredatormaythereforeleadtoanincreaseinabundanceand/orapparent
abundanceviamesopredatorrelease[5][7],sometimesextendingfurthertogreaterpredationpressureonlowertrophiclevels[5],
[8][10].Forexample,insouthernCalifornia,CoyotesCanislatranssuppressmesopredators(GrayfoxesUrocyon
cinereoargenteus,StripedskunksMephitismephitisanddomesticcatsFeliscatus)suchthatinhabitatpatcheswherecoyotesare
rareorabsent,mesopredatorabundanceishigher,resultinginthedeclineofscrubbreedingbirds[5].Toppredatorsand
mesopredatorsmaythereforeinteracttoshapecommunitystructureinawiderangeofecosystems,withimportantimplicationsfor
bothpredatorandecosystemmanagement[1].
Thereisagrowingbodyofresearch,whichidentifiesinteractionsbetweenapexpredatorsandmesopredatorsthatareconsistent
withthemesopredatorreleasehypothesis[1],[6].However,themajorityofstudieshavenotprovidedexperimentalmeasuresof
howchangesintheabundanceoftoppredatorsresultinchangesinmesopredatorabundance,butratherdescribeinteractionsor
associationsbetweenspecies[1],[6].Thepaucityoffielddatarelatesinparttothelogisticalandethicalproblemsassociatedwith
accuratelyestimatingandmanipulatingpredatorpopulations[11].
Inthisstudyweinvestigatedtherelationshipbetweenatoppredator,theEuropeanbadgerMelesmelesandasympatric
mesopredator,thewesternEuropeanhedgehogErinaceuseuropaeus,intheUK.TheEuropeanbadgerisamediumsizedmustelid
carnivoreandhasbecomeanapexpredatorinpartsofitsrange,duetotheextirpationoflargerterrestrialcarnivores[12].Badgers
haveabroadomnivorousdiet,primarilyconsistingofinvertebratesandplantmatter[12],thoughtheyalsoeatsmallermammals
includinghedgehogs.Hedgehogsarethemselvesmesopredatorspredatinguponinvertebrates,smallmammals,reptiles,
amphibiansandtheeggsofgroundnestingbirds[13],[14].Previoussurveysandmanipulationsofhedgehogabundanceindicate
thatfoodavailabilityandbadgerpredationplaykeyrolesindeterminingtheabundance,distributionandbehaviourofhedgehogs
[15][18].Badgersandhedgehogsarenotonlypredatorandprey,butalsosharemanyofthesamefoodresourcesandhave
thereforebeenconsideredtointeractviaintraguildpredation,aswellascompetingforfood[18].Thusthereisthepotentialfor
badgerstoexertastronginfluenceonhedgehogabundance,astheformercanbesupportedathighdensitythroughalternative
foodresources,evenashedgehognumbersdecline[19],[20].
Theopportunitytoexperimentallytesttheeffectsofareductioninpredatorabundanceonpopulationsofacompetingpreyspecies
arosefromtheRandomisedBadgerCullingTrial(RBCT)whichwasareplicated,controlledfieldexperimenttoinvestigatetheeffect
ofcullingbadgersontheincidenceofbovinetuberculosis(TB)incattle[21].Previousresearchhasshownthatthereductionin
badgerabundancebycullingwasassociatedwithincreasesinthedensityofredfoxesVulpesvulpes[22].Hencewidescale

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badgercullingmayaffectotherspeciesthatalsointeractwithbadgers.Wetestedthehypothesisthathedgehogabundanceand/or
behaviourwouldchange,inlinewithpredictionsofmesopredatorrelease,asaresultofreductionsinbadgerabundanceafter
culling.
MaterialsandMethods

(a)ExperimentalDesign

ThedesignandimplementationoftheRBCTarefullydescribedelsewhere[23].Briefly,10tripletswereestablished,each
consistingofthreematchedtrialareasofapproximately100km2andwhichwererandomlyassignedtoproactivebadgerculling,
localizedreactivecullingfollowingtheidentificationofTBincattle,orexperimentalcontrolswithnobadgerculling.Westudiedfour
ofthe10triplets:A(Herefordshire),E(Wiltshire),G(Staffordshire/Derbyshire)andI(Cotswolds).Ineachtriplet,thestudyranfor4
to6years,including3to5yearsofsuccessiveannualbadgerculling(Table1).Forlogisticalreasonsitwasnotpossibletosurvey
forhedgehogspriortocullingintripletsAandE.Hedgehogsurveyswerealsocarriedoutinreactivecullingareasbeforebadger
cullingwasimplementedbutnotafterwardsandsoheretheyaretreatedasadditionalexperimentalcontrols.

Table1.Thenumberofyearsofhedgehogsurveysthatwerecarriedoutineachtriplet.

doi:10.1371/journal.pone.0095477.t001
Withintriplets,trialareasexhibitedsimilardensitiesofbadgerspriortotheonsetofculling[21].Theefficacyofbadgercullinginthe
RBCThasbeenestimatedpreviouslybyusingtrappingdata[24],[25]andbyusingsignsofbadgeractivityasanindex[26].There
wasasubstantialreductioninbadgerpopulationintheculledareascomparedtoexperimentalcontrolareasinalltriplets(Table1
[25],[26]).
(b)DataCollection

HedgehogsurveyswerecarriedoutannuallybetweenJulyandSeptember(followingDoncaster[18]).Withineachtrialarea,12
fieldswereselectedforsurvey.Ninepasturefieldswereselectedrandomlyfromallfieldsavailablewithina1kmradiusofavillage
andthreefieldsofamenitygrassland,whichiscommonlyapreferredhabitatforhedgehogs[15],wereselectedinorontheedgeof
villages[27].
Ineachyear,fieldsweresurveyedoverthreeseparatevisitsbetweenthehoursof23:00and03:00[16],[27].Eachfieldwas
systematicallysearchedforhedgehogsusingspotlightsandhedgehogswereuniquelybuttemporarilymarked[27].Itwasassumed
thathedgehogswouldlosetheirmarksbetweenyears.Thetotalnumberofindividualhedgehogscaughtateachsiteoverthethree
repeatvisitswastakenasanindexofrelativehedgehogabundance/activity.
(c)DataAnalysis

Thecountofindividualhedgehogsineachfieldoverthreevisitsforagivenyearwastreatedastheresponsevariable.Toanalyse
variationinthecountofhedgehogs,wefittedageneralisedlinearmixedmodel(GLMM)withtripletandtreatmentasfixed
categoricalvariablesandtreatmentyearasacontinuousvariable.ThemodelwasfittedwithanIterativeReweightedRestricted
MaximumLikelihood(IRREML)procedurewithanegativebinomialerrorstructureandalogarithmlinkfunction.Thearea(m2)of
eachfieldwaslogtransformedandenteredasanoffsetintotheIRREMLmodel,totakeaccountofvariabilityinfieldsize(i.e.
surveyeffort).Treatmenthadtwolevels:culled(anareaaftertheinitiationofbadgerculling)ornotculled(treatmentareasbefore
theinitiationofbadgercullingandexperimentalcontrolareaswithnoculling).Thetermfield,nestedwithintripletandtreatment,
wasenteredasarandomterm.
Results

Inamenitygrassland,therewasasignificanteffectoftheinteractionbetweenbadgercullingandtheyearofcullingonhedgehog
count(2=8.61,d.f.1,p=0.004)(Table2).Nootherfactorswerefoundtohaveasignificanteffect(Table2).Bytheendofculling
operations,hedgehogcountsonamenitygrasslandshadmorethandoubledinbadgercullingareascomparedtoareaswithno
culling(Figure1).Meanhedgehogcountsrangedfrom0.21.0hedgehogha1wherebadgerswerenotculledto0.92.4
hedgehogsha1wherebadgerswereculled(Figure1).Inpasturefields,only12individualhedgehogswerefoundin22%offields
andsothereweretoofewobservationstocarryoutstatisticalanalyses.

Figure1.MeanhedgehogdensityonamenitygrasslandfieldsduringtheRandomisedBadgerCullingTrial.

Shadedcolumnsshowbadgercullingareasandwhitecolumnsshowexperimentalcontrolareas.
doi:10.1371/journal.pone.0095477.g001

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Table2.SummarisedresultsofGLMMexplainingvarianceinannualcountsofhedgehogsonamenitygrasslandinrelationtoexperimental
badgerculling.

doi:10.1371/journal.pone.0095477.t002

Discussion

Inlinewithpredictionsofthemesopredatorreleasehypothesis,experimentalreductioninthebadgerpopulationresultedinan
increaseinthecountofhedgehogsinamenitygrasslandhabitats.Hedgehogpopulationsand/orbehaviourmay,therefore,be
constrainedduetocompetitionand/orpredation,orthethreatofpredation,byalargerpredator.Thisresult,suggeststhatlethal
controlofbadgerpopulationsmayresultinchangestothestructureofthewiderpredatorcommunity.
Overthecourseofthisstudy,thenumbersofhedgehogscaughtinamenitygrasslandfieldsincreasedbyapproximately100%in
theareaswherebadgerswereculled,butnotinthecontrolareaswherecullingdidnottakeplace.Largerpredatorsmaysuppress
smallermesopredatorseitherbydirectpredation/conflict,orbychangingtheirbehavioursuchthattheyavoidhabitatsorlocations
wherethelargerpredatorispresent[1],[28].Itisthereforepossiblethatincreasedcapturesofhedgehogsinthecurrentstudywere
duenottochangesinhedgehognumbers,buttochangesinhedgehogbehaviour,withhedgehogsbeingmorevisibleoractivein
amenitygrasslandsiteswherebadgernumbershadbeenreduced.Femalehedgehogsmayavoidlargergardenhabitats
associatedwithincreasedbadgeractivity,presumablyduetohighriskofpredation[29].However,badgersandhedgehogshave
alsobeenobservedregularlyusingthesameareas[30].Inaddition,aconcurrenttelemetrystudyofhedgehogsintheCotswold
tripletoftheRBCT[31]foundnosignificanteffectsofbadgercullingonrangingbehaviourthatwouldbeconsistentwithincreased
countsinamenitygrasslandsites.Thissuggeststhattheincreaseinhedgehogobservationswasunlikelytobeduetochangesin
behaviour.
Previousstudiesindicatethatbadgerpredationisoneofthemaincausesofhedgehogmortality[15],[17],[18],andthatbadger
densitycorrelatesnegativelywithhedgehogabundance.Itthereforeseemslikelythattheobservedincreaseinthecountsof
hedgehogsinthecurrentstudyreflectsanincreaseinhedgehogabundancefacilitatedbyreducedpredationandhighersurvival.
Thisisalsoconsistentwithpreviousresearchsuggestingthatbadgerpredationhasnegativeimpactsonhedgehogpopulation
growth[15].
Theanalysesinthisstudywerecarriedoutonthenumbersofhedgehogscaughtonamenitygrasslandsites,asveryfew
hedgehogswereobservedinpasturefields.Amenitygrasslandsandfieldsclosetovillagesorhousesmaybekeyhabitatfor
hedgehogsandofferapotentialrefugeagainstpredationbybadgers,whicharetypicallylessactiveintheseareas,presumablydue
tohumandisturbance[16],[17],[27].Hedgehogpresenceonamenitygrasslandshowsthatpreyspeciescancoexistwithpredators
atalandscapescalebyoccupyingareasofhabitatthataremorefavourabletothepreyspecies,perhapsintermsofreduced
predationriskorimprovedfoodavailability[16],[19].
Inthecontextofmesopredatorrelease,interspecificinteractionsareoftenviewedasasimplisticthreelevelinteractionbetweentop
predator,mesopredatorandsmallprey,particularlywhentheapexpredatorinquestionisanobligatecarnivorewithlittledietary
overlapwithmesopredators[1].Insuchcircumstances,mesopredatorreleasemayincreasepredationpressureonthespeciesthat
arepreyeduponbythemesopredator,potentiallyresultinginpopulationdeclines[5],[8].Theconsequencestothewider
ecosystemofchangesinbadgerandhedgehognumbersarehardertopredict,asbothspecieshavebroadandlargelyoverlapping
dietaryniches[12].Theroleofomnivoresinfoodwebdynamicsisnotwellunderstood,althoughtheymayhavestabilisingeffects
byfeedingacrosshabitatsandtrophiclevels[32].Increasesinhedgehognumbersmayresultinincreasedpredationpressureon
certainpreyspecies.Hedgehogsmayoccasionallypredatelargenumbersofsingleinvertebrateandvertebratesspecies[14].For
exampletheyhavebeenshowntohavesignificantimpactonpopulationsofgroundnestingbirdsundercertainconditions,via
predationofnests[33].However,itisalsopossiblethatadeclineinbadgersandresultantincreaseinhedgehogswillhave
negligibleeffectsonlowertrophiclevels,eitherbecausepreyspecieswerealreadybeingconsumedbybadgers,orbecause
specificpreyspeciesconstituteasmallcomponentofhedgehogdiets.Inconclusion,thisstudydemonstratesthatamediumsized,
mustelidomnivoremayacttoconstrainasmallermesopredator.Thisstudyalsoillustratesthevalueoffieldexperimentstoassess
thepotentialeffectsofmanagementstrategiesontheabundanceofwildlifepopulations.Europeanbadgersareawildlifereservoir
forbovinetuberculosisintheUKandIrelandandareconsequentlyofintensemanagementinterest[26].Thisstudyprovides
informationforassessingthepotentialecologicalconsequencesofbadgercullingandfurtherconfirmationthatareductionin
badgernumberswillhavedirectimpactsonothermammalspecies[22].
AuthorContributions

Conceivedanddesignedtheexperiments:ITRYGJWJDCPDRJD.Performedtheexperiments:ITRYGJWJDNWRJD.
Analyzedthedata:ITRYRAMJDNWARCPD.Contributedreagents/materials/analysistools:ITRYJDARCPD.Wrotethepaper:
ITRYRAMGJWJDNWARCPDRJD.
References

1. RitchieEG,JohnsonCN(2009)Predatorinteractions,mesopredatorreleaseandbiodiversityconservation.Ecologyletters12:982998.doi:
10.1111/j.14610248.2009.01347.x
ViewArticle
PubMed/NCBI
GoogleScholar
2. SergioF,CaroT,BrownD(2008)Toppredatorsasconservationtools:ecologicalrationale,assumptions,andefficacy.AnnualReviewofEcology,
Evolution,andSystematics39:119.doi:10.1146/annurev.ecolsys.39.110707.173545
ViewArticle
PubMed/NCBI
GoogleScholar
3. WallachAD,JohnsonCN,RitchieEG,ONeillAJ(2010)Predatorcontrolpromotesinvasivedominatedecologicalstates.Ecologyletters13:10081018.
doi:10.1111/j.14610248.2010.01492.x
ViewArticle
PubMed/NCBI
GoogleScholar
4. ChakarovN,KrgerO(2010)Mesopredatorreleasebyanemergentsuperpredator:anaturalexperimentofpredationinathreelevelguild.PloSone5:

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0095477

3/5

6/6/2015

PLOSONE:ImpactsofRemovingBadgersonLocalisedCountsofHedgehogs

e15229.doi:10.1371/journal.pone.0015229
ViewArticle
PubMed/NCBI
GoogleScholar
5. CrooksK,SoulM(1999)Mesopredatorreleaseandavifaunalextinctionsinafragmentedsystem.Nature:563566.
6. PrughLR,StonerCJ,EppsCW,BeanWT,RippleWJ,etal.(2009)TheRiseoftheMesopredator.BioScience59:779791.doi:10.1525/bio.2009.59.9.9
ViewArticle
PubMed/NCBI
GoogleScholar
7. SoulM,BolgerD,AlbertsA(1988)Reconstructeddynamicsofrapidextinctionsofchaparralrequiringbirdsinurbanhabitatislands.Conservation2:
7592.doi:10.1111/j.15231739.1988.tb00337.x
ViewArticle
PubMed/NCBI
GoogleScholar
8. LetnicM,DworjanynSa(2011)Doesatoppredatorreducethepredatoryimpactofaninvasivemesopredatoronanendangeredrodent?Ecography34:
827835.doi:10.1111/j.16000587.2010.06516.x
ViewArticle
PubMed/NCBI
GoogleScholar
9. RippleWJ,WirsingAJ,WilmersCC,LetnicM(2013)Widespreadmesopredatoreffectsafterwolfextirpation.BiologicalConservation160:7079.doi:
10.1016/j.biocon.2012.12.033
ViewArticle
PubMed/NCBI
GoogleScholar
10. LeviT,WilmersCC(2012)Wolvescoyotesfoxes:acascadeamongcarnivores.Ecology93:921929.doi:10.1890/110165.1
ViewArticle
PubMed/NCBI
GoogleScholar
11. SaloP,BanksPB,DickmanCR,KorpimkiE(2010)Predatormanipulationexperiments:impactsonpopulationsofterrestrialvertebrateprey.Ecological
monographs80:531546.doi:10.1890/091260.1
ViewArticle
PubMed/NCBI
GoogleScholar
12. RoperT(2010)Badger:HarperCollinsUK.
13. ReeveN,LindsayR(1994)Hedgehogs:T.&ADPoyserLondon.
14. JonesC,MossK,SandersM(2005)Dietofhedgehogs(Erinaceuseuropaeus)intheupperWaitakiBasin,NewZealand:implicationsforconservation.
NewZealandJournalofEcology29:2935.
ViewArticle
PubMed/NCBI
GoogleScholar
15. DoncasterCP(1992)Testingtheroleofintraguildpredationinregulatinghedgehogpopulations.ProceedingsBiologicalsciences/TheRoyalSociety249:
113117.doi:10.1098/rspb.1992.0092
ViewArticle
PubMed/NCBI
GoogleScholar
16. MicolT,DoncasterCP,MackinlayLA(1994)CorrelatesoflocalvariationintheabundanceofhedgehogsErinaceuseuropaeus.JournalofAnimal
Ecology63:851860.doi:10.2307/5262
ViewArticle
PubMed/NCBI
GoogleScholar
17. HofaR,BrightPW(2010)Thevalueofagrienvironmentschemesformacroinvertebratefeeders:hedgehogsonarablefarmsinBritain.Animal
Conservation13:467473.doi:10.1111/j.14691795.2010.00359.x
ViewArticle
PubMed/NCBI
GoogleScholar
18. DoncasterCP(1994)FactorsRegulatingLocalVariationsinAbundance:FieldTestsonHedgehogs,Erinaceuseuropaeus.Oikos69:182192.doi:
10.2307/3546136
ViewArticle
PubMed/NCBI
GoogleScholar
19. PolisGA,MyersCA,HoltRD(1989)TheEcologyandEvolutionofIntraguildPredation:PotentialCompetitorsThatEatEachOther.AnnualReviewof
EcologyandSystematics20:297330.doi:10.1146/annurev.es.20.110189.001501
ViewArticle
PubMed/NCBI
GoogleScholar
20. HoltRD,PolisGA,UrlS(1997)ATheoreticalFrameworkforIntraguildPredation.TheAmericanNaturalist149:745764.doi:10.1086/286018
ViewArticle
PubMed/NCBI
GoogleScholar
21. DonnellyCA,WoodroffeR,CoxDR,BourneFJ,CheesemanCL,etal.(2006)Positiveandnegativeeffectsofwidespreadbadgercullingontuberculosis
incattle.Nature439:843846.doi:10.1038/nature04454
ViewArticle
PubMed/NCBI
GoogleScholar
22. TrewbyID,WilsonGJ,DelahayRJ,WalkerN,YoungR,etal.(2008)Experimentalevidenceofcompetitivereleaseinsympatriccarnivores.Biology
letters4:170172.doi:10.1098/rsbl.2007.0516
ViewArticle
PubMed/NCBI
GoogleScholar
23. BourneJ,DonnellyCA,CoxDR,GettinbyG,MorrisonI,etal..(2007)BovineTB:TheScientificEvidence.AScienceBaseforaSustainablePolicyto
ControlTBinCattle.AnEpidemiologicalInvestigationintoBovineTuberculosis.TheFinalReportoftheIndependentScientificGrouponCattleTB.
London:Defra.
24. BourneF(2007)BovineTB:thescientificevidence.FinalreportoftheIndependentScientificGrouponCattleTB.London:DepartmentforEnvironment,
FoodandRuralAffairs.

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0095477

4/5

6/6/2015

PLOSONE:ImpactsofRemovingBadgersonLocalisedCountsofHedgehogs

25. SmithG,CheesemanC(2007)Efficacyoftrappingduringtheinitialproactivecullsintherandomisedbadgercullingtrial.Veterinaryrecord160:723
726.doi:10.1136/vr.160.21.723
ViewArticle
PubMed/NCBI
GoogleScholar
26. WoodroffeR,DonnellyCa,CoxDR,BourneFJ,CheesemanCL,etal.(2006)EffectsofcullingonbadgerMelesmelesspatialorganization:implications
forthecontrolofbovinetuberculosis.JournalofAppliedEcology43:110.doi:10.1111/j.13652664.2005.01144.x
ViewArticle
PubMed/NCBI
GoogleScholar
27. YoungRP,DavisonJ,TrewbyID,WilsonGJ,DelahayRJ,etal.(2006)Abundanceofhedgehogs(Erinaceuseuropaeus)inrelationtothedensityand
distributionofbadgers(Melesmeles).JournalofZoology269:349356.doi:10.1111/j.14697998.2006.00078.x
ViewArticle
PubMed/NCBI
GoogleScholar
28. SaloP,NordsmM,ThomsonRI,KorpimkiE(2008)Riskinducedbyanativetoppredatorreducesalienminkmovements.JournalofAnimalEcology
77:10921098.doi:10.1111/j.13652656.2008.01430.x
ViewArticle
PubMed/NCBI
GoogleScholar
29. DowdingCV,HarrisS,PoultonS,BakerPJ(2010)Nocturnalrangingbehaviourofurbanhedgehogs,Erinaceuseuropaeus,inrelationtoriskandreward.
AnimalBehaviour80:1321.doi:10.1016/j.anbehav.2010.04.007
ViewArticle
PubMed/NCBI
GoogleScholar
30. HaighAJ(2011)TheecologyoftheEuropeanhedgehog(Erinaceuseuropaeus)inruralIreland:UniversityCollegeCork.
31. YoungRP(2005)Fieldtestofimpactsofintraguildpredationonamammalianpreypopulation:thebadgerMelesmelesandhedgehogEricaceus
europaeus[PhDThesis]:UniverstiyofSouthampton.
32. MorrisDW(2005)Paradoxicalavoidanceofenrichedhabitats:havewefailedtoappreciateomnivores?Ecology86:25682577.doi:10.1890/040909
ViewArticle
PubMed/NCBI
GoogleScholar
33. JacksonD,GreenR(2000)Theimportanceoftheintroducedhedgehog(Erinaceuseuropaeus)asapredatoroftheeggsofwaders(Charadrii)on
machairinSouthUist,Scotland.BiologicalConservation93:333348.doi:10.1016/s00063207(99)001354
ViewArticle
PubMed/NCBI
GoogleScholar

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