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animals Article Moles and Mole Control on British Farms, Amenities and Gardens after Strychnine Withdrawal SandraE.Baker*,StephenA.Ellwood,PaulJ.JohnsonandDavidW.Macdonald TheWildlifeConservationResearchUnit,DepartmentofZoology,TheRecanatiKaplanCentre, UniversityofOxford,TubneyHouse,AbingdonRoad,Tubney,AbingdonOX135QL,UK; [email protected](S.A.E.);[email protected](P.J.J.); [email protected](D.W.M.) * Correspondence:[email protected];Tel.:+44-1865-611-114 AcademicEditors:MarinavonKeyserlingk,KateLittin,TrudySharpandNgaioBeausoleil Received:26February2016;Accepted:9May2016;Published:8June2016 SimpleSummary: MolesareburrowingmammalsthatareregardedaspestsinBritain,anduntil 2006theycouldlegallybekilledusingstrychninepoison. Whenstrychninewaswithdrawnthere were fears that mole populations would increase. We surveyed farmers, amenity managers and householders about moles and mole control on their land in 2007, post strychnine withdrawal. Kill-trappingwasbyfarthepreferredcontrolmethodusedandcontrolmaybeusedmorethancan bejustifiedbydamagelevelsortheeffectofcontrolondamage. Moletrapsareunregulated,unlike mostotherspringtraps,andsomemightnotmeetcurrentwelfarestandards. Wefoundnoevidence thatmoleactivityhadincreasedsincea1992surveyoffarms. Abstract: MolesareconsideredpestsinBritain,butthisissuehasbeenlittlestudied. Lowerwelfare standardshavebeentoleratedformolesthanformostothermanagedwildmammalspecies,asuse ofboththecontroversialpoison,strychnine,andunregulatedtrapshavebeenpermitted. Strychnine waswithdrawnin2006andtherewerefearsthatmolepopulationswouldincreaseasaresult. In 2007,weconductedacomprehensive,nationwidesurveyoflandmanagerperceptions,opinionsand behaviourregardingmolesandmolecontrolonfarms,amenitiesanddomesticgardensinBritain. Wesurveyed2150landmanagers(achievinga59%responserate)andground-truthed29responses. Moles were reported to be present on most farms and amenities, and 13% of gardens, and were more common in lighter soils. Where present, moles were usually considered pests, this being morelikelyinWales,ScotlandandnorthernEngland,onlivestockandmixedfarms,andonlarge, high-valueamenities,e.g.,racecoursesandgolfcourses. Molecontrolfollowedsimilarpatternsto mole presence. More control may occur than is economically, and therefore potentially ethically, justified.Controlshouldbemorecarefullyconsideredand,wherenecessary,moreeffectivelytargeted. Kill-trappingwasthefavouredrecentandfuturemethodonfarmsandamenities,evenifstrychnine wastobereintroduced; however, becausemoletrapsarecurrentlyunregulated, somemightnot meet current welfare standards if tested. We found no evidence for an increase in moles since a farmquestionnairesurveyconductedin1992;thiscouldhavewiderimplicationsforfuturewildlife managementpolicychanges. Keywords: European mole; Talpa europaea; farms; amenities; gardens; mole control; strychnine; trapping;silage;questionnaire 1. Introduction European moles (Talpa europaea) are solitary, fossorial insectivores. Each individual inhabits its own system of underground feeding tunnels, in which it hunts soil invertebrates, principally Animals2016,6,39;doi:10.3390/ani6060039 www.mdpi.com/journal/animals Animals2016,6,39 2of26 earthworms[1,2].MolesaredistributedacrossmuchofEuropeandintoRussia[3],andarewidespread throughoutBritainbutdonotoccurinIrelandoronmanyoftheBritishislands[1,2]. Molesproduce conspicuousabove-groundsignsintheformofspoilheapsknownasmolehills. Moletunnellingand molehillproductionareacauseofcomplaintamongfarmers,amenitymanagersandgardeners,and molesarewidelyconsideredpests[4]. Reported mole/molehill damage on farms includes pollution of silage, coverage of pasture, encouraging weed invasion and causing damage to plants, machinery and drainage systems/watercourses[4,5]. Damagereportedonamenitiesandgardensisoftenmerelyaesthetic, butmoleactivityonracecourses,grassairstripsandsportsfieldsmaypresentriskofinjurytopeople andanimals[4,6]. Someaspectsoffarmers’experiencesandopinionsofmolesaspests,andofmole managementconductedbyfarmers,havebeenstudiedinBritain[5]andScotland[7],butnosuch researchhasbeencarriedoutwithamenitymanagersorhouseholders(domesticgardeners). Inaquestionnairesurvey,conductedin1992,Atkinsonetal. (1994)[5]foundthat64%offarmers perceivedmolesaspestsbutthatonly50%ofsurveyrespondentsmadeeffortstomanagethem;the authorsconcludedthatthedamageattributedtomoleswasslightonmostfarms. Nodetaileddataare availablefromtheScottishstudy[7]. Thecostsassociatedwithmoledamageandcontrolaredifficult toassess,butestimatesforthetotalannualcosttofarminginBritainrangebetween£2.5millioninthe late1980sandupto£5millionin1994[2,4]. Thesefiguresequatetoarangeof£5.6–7.8millionat2015 prices[8]. Toputthisinperspective,theannualcostofEuropeanbadger(Melesmeles)damagecaused tocrops,orthroughburrowingorpredation,inEnglandandWaleswasestimatedtobesomewhere between£21.5–41.5millionperannumin1997[9],equatingto£35–68millionin2015. VariousmethodsareusedformanagingmolesandmolehillsinBritain,includingkill-trapping (spring-trapping),gassingwithphosphine,live-trapping,shootingandultra-sonicdeterrents,aswell asharrowingandrollingtoflattenmolehills. Nopoisonsorchemicalrepellentsarecurrentlyapproved forusewithmoles. RenardineTM,aboneoil-basedrepellent[10],wasapproveduntil2005,whenthe licensewaswithdrawnacrossEuropefollowingconcernsoverthetransmissionofBovineSpongiform Encephalopathy(BSE)[11]. Strychnineisanalkaloidpoison. Actingthroughcompetitiveinhibition of glycine (a neurotransmitter), strychnine allows unimpeded stimulation of motor neurons thus causing the striated muscles to produce generalised rigidity and tetanic seizures [12]. Ultimately thisleadstodeaththroughexhaustionorthroughasphyxiaarisingfromparalysisoftherespiratory muscles. Strychninehasbeendescribedasdangerousandinhumane[5]andwasbannedforwildlife managementundertheAnimals(Cruelpoisons)Regulationsof1963,andyetitscontinuedusewas allowedonlywithmoles. Atkinsonetal.[5]suggestedthisexemptionwastoleratedbecause: (1)mole damage was perceived to demand action; (2) strychnine was the only poison cost-effective on an agriculturalscale(andnosuitablealternativewasavailable);(3)molesdiedundergroundwheretheir painfuldeathscouldnotbeseen. Forapesticidesuchasstrychninetobeusedlegally,notonlymustthepesticidenotbeillegalfor theparticularpurpose,butthepesticidemustalsobeapprovedforthatuse(nowbytheChemicals RegulationDirectorate[13]. Bytheearly1990s,thenumberofpermitsforstrychnineusewasbeing reduced[5],andin2006strychnineapprovalforusewithmoleswaswithdrawn,aspartofanEU pesticide review, when manufacturers failed to produce the necessary health and safety data [14]. Thisfollowedyearsofconcernaboutthehumanenessofstrychninepoisoning,thepotentialforits misuse,andthesignificantcostsinvolvedinadministeringandpolicingitsuse[4,5,15]. Strychnine alsopresentedthepotentialthreatofaccidentalprimaryorsecondarynon-targetpoisoningofwildlife andcompanionanimals(see[15,16]). However,strychninewasconsideredtobethemostcost-effective methodformanagingmolesonalargescale,althoughkill-trappingbyskilledoperatorsmaybebetter forremovingsmallnumbersofmoles[4,5]. Strychnineisreportedtohaveoncebeenthemainmethod of managing moles on British farms [7]; see also [4,5]. It has been suggested that, because of the relativelylowcostofstrychninetreatment,moremolesmayhavebeenkilledthancouldhavebeen Animals2016,6,39 3of26 justified[4]. Anditispossiblethat,ifthesehadnotbeenpoisoned,somemayhavediedanywayas partofthenaturally“doomedsurplus”(seeErrington[17]). Althoughstrychnineuseinmolecontrolwasalreadydecliningbeforeitwasbanned,in2004/2005 sufficient strychnine was sold in Britain to kill almost 800,000 moles [1] and 3000 users were licensed[14]. Itwaspredictedthatthelossofstrychninewouldforceuserstoreconsiderthecosts and benefits of managing moles using alternative methods, and that trapping would become the primarymethodofmolemanagement,whileforsomemanagers,takingnoactionmaybecomethebest choice[4,5]. Becauseofconcernthatnosuitablealternativemolemanagementmethodwasavailable, itwassuggestedthatmolepopulationswouldincrease[4,18],anditwassubsequentlyclaimedthat theyhad[19–21]. Althoughstrychnineapprovalwaswithdrawnin2006,thelegalexemptiontouseit formolecontrolremains,soifstrychninewastobere-approvedinfuture,itwouldbecomelegalto useitagain. We conducted a comprehensive, nationwide questionnaire survey to examine land manager perceptions, opinions and behaviour regarding moles and mole control on farms, amenities and domesticgardensthroughoutBritain,andfactorsaffectingthese. Wesetouttocomparesomefarmer responseswiththoseobtainedinAtkinsonetal.’s1992survey[5].Wealsoaimedtoestimatetheimpact, ifany,ofstrychninewithdrawalonmolemanagement. Weconductedfieldsurveystoground-truth a sample of questionnaire respondents for validation purposes. We found that where moles were present,theywerecommonlyregardedaspests,andoftencontrolled. Ourresultsalsosuggestthat morecontrolmaybeconductedthanisnecessaryandthatitmaybepoorlytargeted.Toourknowledge, thisisthefirstpublishedsurveyofmolemanagementissuesonamenitiesanddomesticgardens,and thelargestsurveyconductedtodateregardingmolesonfarmland. 2. MaterialsandMethods ThisstudywasgrantedethicalapprovalbytheCentralUniversityResearchEthicsCommitteeat theUniversityofOxford(approvalreferencewasSSD/CUREC1/07-003). 2.1. QuestionnaireSurvey We designed a postal survey using Dillman’s Tailored Design Method [22] to optimise surveyresponserate, includingmultiplecontactswithpotentialrespondents: (a)pre-noticeletter; (b)questionnaireandcoveringletter;(c)thankyou/reminderpostcard;(d)replacementquestionnaire tonon-responders. Weobtainedendorsementforthesurveyfrom:TheNationalFarmers’UnionWales, TheNationalFarmers’UnionofScotland,TheBritishAssociationforShootingandConservation,The GameConservancyTrust(nowtheGameandWildlifeConservationTrust),TheNationalTrust,The RoyalHorticulturalSociety, TheRacecourseAssociation, TheAirportOperators’Association, and TheBritishandInternationalGolfGreenkeepers’Association. Thesupportoftheseorganisationswas mentionedinthecoveringletteraccompanyingquestionnairemailings. Wedevisedseparatequestionnairesforfarmers,amenitymanagersandhouseholders,wherever possible using similar-style questions across the three target groups to allow comparison of responses. Questions covered: (a) Your farm/amenity/garden or allotment; (b) Moles on your farm/amenity/garden or allotment; (c) Mole control on your farm/amenity/garden or allotment; (d)Youropinionsaboutmolecontrol;(e)Strychninepoison;(f)Interestinparticipatinginafollow-up study(ground-truthing). Questionnairesweremarkedwithindividualidentitynumbersandprinted as A4 double-sided booklets. Questionnaires were piloted, and contacts and questionnaires were translatedfromEnglishintoWelsh(recipientsinWalesreceivedversionsinbothlanguages). Weused asystemdetailedunder“Recipientdatabaseabstraction”,inTextS1,torandomlyselect1204farmers, 551 amenity managers and 504 householders from across England, Scotland and Wales as survey recipients. Contactswerepostedtothe2259recipientsasfollows: (1)21June2007pre-noticeletter (allrecipients);(2)28June2007questionnaireandcoveringletter(allrecipients);(3)4July2007thank you/reminderpostcard(allrecipients);(4)18July2007replacementquestionnaire(non-responders Animals2016,6,39 4of26 only). FulldetailsofthequestionnairesurveymethodaregiveninTextS1andexamplesofthecontacts andofthequestionnairearegiveninTextS2–S6. 2.2. QuestionnaireAnalysis QuestionnairedatawerestoredinaMicrosoftAccessdatabaseandanalysedusingSASstatistical software(TheSASInstitute,Cary,NC,USA)[23]. Weusedχ2and(whereexpectedcountswerelow) Fisher’sExactteststoexaminerelationshipsamongcategoricalvariablesincontingencytables. We usedFriedmanTeststoanalyserankedmonthlymoleactivityandrankedfeaturesofamolecontrol method. Kruskal-Wallisχ2testswereusedtocomparecostsdataacrosscategoriesofrespondenttype. 2.3. Ground-TruthingSurvey Weground-truthedaproportionofquestionnairerespondents, toassesstheaccuracyoftheir reportsregardingthepresenceofmolesandthescaleofmoleactivityontheirland. Ground-truthing tookplaceinspringandearlysummer2008and2009tocoincidewithperiodsofpeakmoleactivity andcontrolasreportedinourquestionnairestudy,andbyAtkinsonetal.[5]. Weselectedpotential participantsfromquestionnairerespondentswhoindicatedaninterestinparticipatinginfollow-up work. Weattemptedtoidentifyarepresentativesampleof29questionnairerespondents,consistingof 17farmers(59%),eightamenitymanagers(28%)andfourhouseholders(14%),closelyreflectingthe proportionsinwhichtheyrespondedtothequestionnaire(57%,24%,and19%). Thefarmsincluded sevenmixed,sixlivestock,twoarable,andtwootherfarms,andtheamenitieswerethreeornamental gardens,twogolfcourses,oneracecourse,oneancientmonumentandonepark. Ground-truthingwasatwo-stageprocessconsistingofa30–60minsemi-structuredface-to-face interview,followedbyafieldsurvey. Interviewswereintendedlargelytohelpusidentifysuitablesites forground-truthingusingfieldsurvey. Duringtheinterviewparticipantswereaskedtoidentify,on mapsandaerialphotographs,sixsites(fields/areas),ontheirland,whichhadatthattimethegreatest (threesites)andleast(threesites)moleactivity. Wherearespondentreportednocurrentmoleactivity onhisland,hewasaskedtonominatejustthree“noactivity”sites. Werecordedfurtherinformationfor eachsiteincludingcurrentland-use(e.g.,pasture,cereal,mowngrass)andsiteaccesspoints.Following eachinterview,weselectedtwositesforsurvey—oneeachidentifiedbytheparticipantashavinghigh moleactivityandlowmoleactivity;weattemptedtobalancetherepresentationofdifferentfield/area types,thenprioritisingsiteswiththemostandtheleastreportedmoleactivity. Whereaparticipant nominatedonly“noactivity”sitesweselectedoneforsurvey. Ofthe29participants,23reportedboth highandlowactivitysitesandsix(oneamenitymanager,onefarmerandfourhouseholders)reported nocurrentmoleactivity;so,altogetherwesurveyed52sites. Fieldsurveyswereconductedwithina fewdaysoftheinterview. Ateachsiteweimplementedasimplesurveystrategythatcouldbeadapted todifferentfieldcircumstances;thisinvolvedconductingapproximately30full-pointsurveysateach sitetorecordmolehillactivity. Fulldetailsoftheground-truthingsurveymethodaregiveninTextS7. Duringtheground-truthinginterviewwealsoaskedparticipantswhethertheyfeltthatmole activityhadincreased,decreasedorstayedthesameontheirlandovertheprevious5years. 2.4. Ground-TruthingAnalysis Firstwecheckedwhetherlandmanagershadsuccessfullypredictedmolehillpresenceand/or absence on their land. Then, using our survey data, we calculated molehill density per site and comparedtheseestimateswithlandmanagers’reportedperceptionofrelativemolehillactivityon theirnominatedsites. FurtherdetailsofdatamanipulationandanalysesaregiveninTextS8. Animals2016,6,39 5of26 3. Results 3.1. QuestionnaireResponseRates We included responses received up to the 11 November 2007. After excluding ineligible recipients,therewere2150potentialrespondentscomprising1143farmers,526amenitymanagersand 481householders.Weachievedanoverallresponserateof59%(n=1265),withresponseratesdiffering amongrespondenttypes(farmers63%(n=720),amenitymanagers58%(n=303),householders50% (n=242)(χ2 =22.9,p<0.001,TableS1)),andregions,withmoreinScotlandandinthenorthand (2) south-west of England (χ2 = 12.3, p = 0.015, Figure S1). There was no evidence that respondent (4) typevariedwithregion(χ2 =6.7,p=0.574). Moreinformationonquestionnaireresponseratesis (8) availableinTextS9. 3.2. QuestionnaireResponses 3.2.1. FactorsAffectingMolePresenceandPestStatus Weexaminedfactorsaffectingreportedhistoricmolepresence(whethertheyhadeverhadmoles on their land) and mole pest status. Where “mole presence” and “pest status” are used here we refer respectively to mole presence at any time in the past and perceived mole pest status. In the questionnaires, respondents were asked first whether moles were present on their land and then whethertheyconsideredmolestobepests. Thereisatleastsomepossibilitythataskingwhethera particularspeciesisapestmight“lead”somerespondentstosaythatitiswhenthatspeciesmightnot haveappearedonanunpromptedlistofpests. However,thisapproachistransparentandavoidsbias whenmakingcomparisonsacrosscategoriesof,forexample,respondenttype,farmenterprisetypeetc. Reportedmolepresencedifferedsignificantlyamongrespondenttypeswithmostfarmersand amenities, but only 13% of householders, reporting that they had ever had moles on their land (Figure1a). Themajorityofrespondentsreportingmolepresenceconsideredthemtobepestsandthe likelihoodofthemdoingsodifferedsignificantlyamongrespondenttypes,withamenitymanagers themostlikelytoconsidermolesontheirlandaspests(Figure1b). Reportedmolepresenceandpest statuswererelatedtobothfarmenterpriseandamenitytype(Figure1c–f). Livestockfarmsweremost likelytoreportmolepresencefollowedbymixedandthenarablefarmers,andofthesefarmerswith moles,livestockfarmerswerealsomostlikelytoconsiderthempests(Figure1c,d). Molepresence wasreportedbyaminorityofbowlinggreensandsportsfields,butbymostgolfcourses,ornamental gardens,parks,racecoursesandotheramenities(Figure1e). Ofthosewithmoles,over96%ofgolf coursesandracecoursesconsideredmolespeststogetherwiththemajorityofotheramenities(bowling greenswereexcludedbecausesofewreportedmolepresence)(Figure1f). Therewasnoevidencethatmolepresencevariedregionally(Figure1g),althoughrespondents fromWales,ScotlandandnorthernEnglandweremorelikelytoconsidermolesontheirlandaspests (Figure1h). Soiltypewasrelatedtomolepresence,withmolesoccurringmorefrequentlyonloamy soilsandleastonclay(Figure1i),butwheremoleswerepresenttherewasnoeffectofsoiltypeonpest status(Figure1j). Soiltypealsovariedwithregion(χ2 =93.5,p<0.001),withchalkyandclaysoils (24) moreprevalentincentralandeastern,andsouth-western,Englandandloamy-peaty-sandysoilsmore commoninScotland,WalesandnorthernEngland(FigureS2),thussuggestingthatregionalvariation inthepresenceofmolescouldbeatleastpartlyrelatedtosoiltype. 3.2.2. FactorsAffectingRecentMoleActivityandDamage Weexaminedtheimpactofthesamefactorsonrecentmoleactivityandwhetherrespondents consideredthisactivitytobedamaging. Where“recentmoleactivity”and“damage”areusedhere wereferrespectivelytomoleactivityintheprevious12monthsandperceivedmoledamage. Data on recent activity indicated that respondents from farms (particularly livestock and mixed farms) andamenities(particularlygolfcourses),andfromWalesandnorthernEngland,reportedagreater Animals2016,6,39 6of26 incidenceofrecentactivitythandidhouseholdersandrespondentsfromScotland,centralandeastern England and south-western England (Figure S3). When those respondents reporting recent mole activitywereaskediftheyconsideredthistoconstitutedamage,amenitymanagers,livestockfarmers andrespondentsfromWalesandnorthernEngland,weremorelikelythanotherstosaythattheydid. Animals 2016, 6, 39 5 of 26 MoredetailsareavailableinTextS10. FiFgiugruer e1. 1M. oleM porleesepnrcees eanncde manodle mpeoslte stpaetusts, srteastpuesc,tirveeslpye, cbtiyv: e(lay,, bb)y r:esp(ao,ndbe)ntr etsyppoen; d(ce,n dt) tyfaprem; en(ct,erdp)rfiasrem tyepnet; e(rep, rfi)s eamtyepneit;y (tey,pfe), a(gm, ehn) itryegtiyonp;e ,(i,( gj), sho)il rteygpioe.n ;Am(i,enj)itys otiylpteysp aer.e: ABomweln =it ybotwypliensg graereen:s, BGoowlfl == gboolfw cloinugrsgerse, eOnrsn, =G oorlnfa=megnotlafl gcaorudresness,, POarrkn = =paorkrns,a Rmaecnet =a lragcaercdoeunrsse,s,P Saprokrt=s =p saprokrst,s fieRladcse a=ndra Ocetcho =u rostehse,rS apmoretnsit=iess.p Roertgsiofinesl dasrea: nNd =O ntohrt=heortnh Eerngalmanedn,i tCieEs .= Rceengtiroanl sanadre e:aNste=rnn Eonrtghlaenrnd, SWEn =g lsaonudth,-CwEes=tecrnen Etnragllaanndd, Secaostt e=r nScEotnlagnladn adn,dS WWa=less o=u Wtha-lwese.s StetartnistEicnsg slahnodw,nS acroet r=esSucltost loafn d2 taensdts. Wales=Wales.Statisticsshownareresultsofχ2tests. 3.2.3. Factors Affecting Mole Control We examined factors affecting whether historic and recent mole control had been conducted. Where “historic control” and “recent control” are used here we refer, respectively, to control conducted at any time in the past and control conducted in the previous 12 months. Historic control was affected by respondent type, farm enterprise, amenity type, region and soil type (Figure S4a,c,e,g,i). Patterns of control were generally similar to those reported for mole presence but occurring at a lower level (Figure S4a–j, e.g., while 93% of farmers reported mole presence, 65% reported control). One key difference, however, was that historic mole control was related to region Animals2016,6,39 7of26 3.2.3. FactorsAffectingMoleControl We examined factors affecting whether historic and recent mole control had been conducted. Where“historiccontrol”and“recentcontrol”areusedherewerefer,respectively,tocontrolconducted atanytimeinthepastandcontrolconductedintheprevious12months. Historiccontrolwasaffectedbyrespondenttype,farmenterprise,amenitytype,regionandsoil type(FigureS4a,c,e,g,i). Patternsofcontrolweregenerallysimilartothosereportedformolepresence butoccurringatalowerlevel(FigureS4a–j,e.g.,while93%offarmersreportedmolepresence,65% reportedcontrol). Onekeydifference,however,wasthathistoricmolecontrolwasrelatedtoregion (whereashistoricpresencewasnot), withcontrolmoreprevalentinWales, Scotlandandnorthern England(χ2 =33.1,p<0.001,FigureS4g). (4) Of those respondents that had ever controlled moles, there was no evidence for difference amongfarmenterprises,amenitytypesorregions,intheproportionthatcontrolledthemrecently (FigureS4d,f,h). Thissuggeststhatwhetherahistoriccontrollertendedtocontrolmolesfromyearto yearwasnotaffectedbythesefactors. Howeverthelikelihoodthatahistoricmolecontrollercontrolled molesrecentlywasaffectedbyrespondenttypeandsoiltype,withamenitymanagersandfarmers, and respondents on peaty or sandy soil, being more likely to have controlled moles recently and perhapsthereforeregularly(FigureS4b,j). 3.2.4. CostsofMoleDamageandControl Meancostsofdamageandcontrolwerecalculatedforrespondentsreportingdamageorcontrol respectivelyinthepreviousyear. Totalcontrolcostscomprisedprofessionalcosts(asreported)and non-professionalcosts(whichwerecalculatedbyaddingthereportedcostofmaterialstotheestimated cost of labour—produced by multiplying the reported number of hours spent on mole control by theUKhourlyminimumwagerateforpeopleaged22yearsorabove(£5.73,theequivalentof£7.10 in2014[8])). There was no evidence that raw damage costs per holding differed among respondent types (KruskalWallis(KW)χ2 =1.58,p=0.45). Incontrast,rawtotalcontrolcostsdifferedsignificantly (2) amongrespondenttypes(Figure2a,KWχ2 =21.70,p<0.001),withthoseforamenitiesbeinggreatest, (2) followedbyfarmsandthenhouseholds;underlyingprofessionalandnon-professionalcontrolcosts followedasimilarpatternbutdidnotdiffersignificantlyamongrespondenttypes(KWχ2 ď5.56, (2) pě0.06). Whencostswereexpressedperha,damage,totalcontrolandprofessionalcontrolcostsall differedsignificantlyamongrespondenttypes,withthoseforhouseholdsbeinggreatest,followedby amenitiesandthenfarms(Figure2b,KWχ2 ě5.82,pď0.05),butprofessionalcontrolcostsdidnot (2) (KWχ2 =4.71,p=0.10). (2) Morerespondentsreportedcontrolcoststhanreporteddamagecosts(seesamplesizesinFigure2b legend: farms150/24,amenities89/19,householders4/2),andamenitiesspentmoreoncontrolthan theysustainedindamage,whilefarmersandaverysmallsampleofhouseholderssustainedagreater valueofdamagethantheyspentoncontrol(basedonmedianvalues). Damage and control costs¨ha´1 for farmers reporting damage or control, respectively, were greatestforlivestockfarmers,followedbymixedandarablefarmers. Damagecostsandtotalcontrol costsvariedmarginallysignificantly(KWχ2 =5.54,p=0.063)andsignificantly(KWχ2 =15.07, (2) (2) p<0.001),respectively,amongfarmenterprises,butprofessionalandnon-professionalcontrolcosts didnot(Figure3a,KWχ2 ď2.69,pě0.260). (2) Samplesizesweresmallforsomeamenitytypesbutneverthelesstherewasasignificantdifference amongamenitytypesinbothtotalandprofessionalcontrolcosts¨ha´1 (KWχ2 ě11.5,pď0.02), (4) withracecoursessufferingthegreatesttotalcosts¨ha´1(NBtherelativelylargecostsforsportsfields were reported by very few respondents). There was no significant difference in damage costs or non-professionalcontrolcosts(Figure3b,KWχ2 ď7.03,pě0.13). (4) Animals 2016, 6, 39 7 of 26 Animals2016,6,39 8of26 FFigiguurer e22. .DDamamaaggee aanndd ccoonntrtrool lccoossttss ffoorr ffaarrmmss,, aammeenniittiieess aanndd hhoouusseehhoollddss ((rreessppoonnddeennttssw witithhd daammaaggeeo orr coconndduuctcitnign gcocnotnrtorlo)l: )(:a()a R)Rawaw cocsotsst spepre hrohlodlidnign g(d(admamagaeg-efa-framrms ns n= 2=62, 6a,maemneitnieitsi ens =n 2=1,2 h1,ohuoseuhsoehldosl dns = 2;n t=ot2a;l tocotanltrcooln-ftarorml-fsa rnm s= n1=571, 5a7m, aemnietineist iens n= =989,8 ,hhoouusseehhooldlds snn == 55;; pprroofefessssioionnaall cocnontrtorol-lf-aframrmss nn == 226699,, aammeenniittiieessn n= =1 3103,0h,o uhsoeuhsoelhdoslnds= n6; n= o6n;- pnroonfe-spsrioofneaslscioonnatrlo lc-ofanrtmrosl-nfa=rm16s8 ,na m= e1n6i8ti,e sanm=en1i0ti8e,s nh =o 1u0s8e,h hooldussenho=ld5s); n( b=) 5C);o (sbt)s ¨Choas´ts1·h(da−a1m (daagme-afgarem-fasrnm=s n2 4=, 2a4m, aemnietineistines= n 1=9 1,9h,o huosuesheohldolsdns n= =2 ;2t; ototatall coconntrtorol-lf-afarmrmss nn == 115500, ,aammeenniittiieess nn == 8899,, hhoouusseehhoollddss nn == 44;; pprrooffeessssiioonnaall ccoonnttrrooll--ffaarrmmssn n= =2 25555,,a ammeennitiiteises n=119,householdsn=5;non-professionalcontrol-farmsn=161,amenitiesn=99,householdsn=4). n = 119, households n = 5; non-professional control-farms n = 161, amenities n = 99, households n = 4). Barsaremeanswithstandarderrors,anddotsindicatemedianvalues. Bars are means with standard errors, and dots indicate median values. Animals 2016, 6, 39 8 of 26 Animals2016,6,39 9of26 FFigiguurree 33. .DDamamagaeg eanandd ccoonntrtorol lccoosststs·h¨ah−a1´, 1a,nadn dtottaolt aflinfiannacniacli alloslosessse sdudeu etot ommoloel eacatcitviivtiyt·yh¨ah−1a,´ b1y,: (ab)y f:ar(ma) efanrtmerpernistee r(pdraimseag(dea-amraabglee- nar =a 5b,l elivnes=to5c,k lniv =e 7st, omcikxend =n =7 ,11m; tioxetadl cnon=tr1o1l-;atroatballe cno =n 2tr8o, ll-iavreasbtoleck nn == 2783,, livmeisxteodc knn == 7432,; mpirxoefdesnsi=on4a2l; pcoronfterossl-iaornaabllec onnt r=o l-4a8ra, bllievenst=oc4k8 ,nli v=e st1o2c0k, nm=ix1e2d0 ,nm i=x ed78; nnon=-p7r8o;fensosnio-pnraol fceosnstiroonla-alrcaobnletr no l=- a3r0a, blilveensto=ck3 0n, =l i7v8e,s mtoicxkedn n= = 7485,);m (bi)x aemdenn=ity4 t5y)p;e(b (d)aammaegneit-ygatrydpeens n( d=a 3m, gagoelf- gcoarudrseenss nn == 73, ,pgaorklfsc no u=r 6se, srance=co7u,rpsaersk ns =n 2=, s6p,orartcse cfioeuldrsse ns =n 1=; t2o,tsapl ocrotnstfiroell-dgsarnd=en1s; nt o=t a2l8, gcoolnf tcrooul-rgseasr dne =n s27n, p=a2rk8s, gn o=l f14co, ruarcseecsonur=se2s 7n, =p a1r5k, sspnor=ts1 f4ie,lrdasc enc =o u3r; spersofnes=sio15n,als pcoonrttsrofil-egldarsdnen=s n3; = 4p4r, ogfoeslfs icoonuarlsceos nnt r=o l3-g5a, rpdaernkss nn == 4146,,g roalcfeccoouurrsseess nn= =3 159,,p saprkosrtns =fie1l6d,sr anc e=c 3o;u nrsoens-pnr=of1e9s,sisopnoarlt scfioneltdrosl- gnar=de3n;ns onn =- p2r8o, fgeosslfi ocnoaulrsceosn tnr o=l -2g9a, rpdaernkss nn == 2185,, rgaoclefccoouurrsseess nn == 2209, ,sppaorrktss fnie=ld1s5 n, r=a 5ce).c Bouarrss easren m=e2a0n,s wspitohr ststafinedldarsdn e=rr5o)r.sB, aanrsda dreotms einadniscwatiet hmsetdaniadna vrdaleurerso.r s,anddotsindicatemedianvalues. 33.2.2.5.5. .SSeeaassoonnaal lPPaattteterrnnss inin MMoolele AAcctitvivitiyty aanndd CCoonnttrrool l WWee eexxaammiinneedd sseeaassoonnaal lppaattteterrnnss inin rreecceennt tmmoolele aacctitvivitiyty aanndd ccoonntrtorol.l .FFirisrts twwe easakskeded rersepspoonnddenentst sto rtaonrka tnhke tthhereteh rmeeonmthosn tinh sthine tphreevpiroeuvsi oyuesary iena rwihniwchh micohlem aoclteivaicttyi vwitays wgraesagterseta. tWeset. tWheent halelnocaaltleodca at e“d3” toa t“h3e” mtoonththe rmanonkethd rhaingkheedst,h “i2g”h etos tt,h“e2 m”otontthh eramnkoendth sercaonnkde dansdec “o1n”d toa nthde “m1”ontotht rhaenmkeodn tthhirrda.n Fkreodm ththeisred .nFurmombetrhs ewsee npurmodbuecresdw meeparnod ruelcaetdivme eraannkrienlagtsi vfoerr amnokninthglsyf oacrtmivoitnyt;h olyvearcatlilv, iatyct;iovviteyr arlelp,aocrttievdit byy arlel preosrpteodnbdyenatlsl rveasrpieodn daecnrotsssv tahriee dmaocnrtohsss (tFhreiemdmonatnh’ss (χF2r(i1e1)d =m 1a3n3’1s.0χ,2 p(1 1<) =0.010313)1. .0P,aptt<er0n.s0 0o1n) .fParamttes rannsd aomnefnairtmiess awnderaem bernoaitdielsy wsiemreilbarro waditlhy sai mlairlagrerw sitphrianlga rpgeearks parnindg ap semakaallnedr oansem ianl lethreo naeuitnumthne,a wutiuthm tnh,e swpriitnhgt hpeeaskp rbienigngp emakorbee eixnaggmgeorraeteedx afogrg fearramteedrsfo thrafanr ammeersnitthieasn aanmde tnhiet iaeustaunmdnt hpeeaaku tmuomrne ppreoankomunocreed fporr oanmoeunnictieeds fthorana mfaernmiteiress (tFhiganurfea r4ma)e. rTsh(eF ivgeuryre s4maa).ll Tnhuemvbeerry osfm hoalulsneuhomldbeerrso tfhhato uressephoonlddeerds ttoh athte qrueespstoionnd erdeptoortthede qau seinstgiolen preeapko rinte dacativsiintyg lien ptheaek suinmamcteivr.i t yinthesummer. Animals 2016, 6, 39 9 of 26 We asked respondents to report the months in which they conducted mole control during the previous year and plotted the proportion of responses for each month (Figure 4b). Control patterns broadly followed activity patterns. Overall, control reported by all respondents varied over the year (χ2(11) = 259.7, p < 0.001). Patterns in control differed between farmers and amenities (χ2(11) = 67.0, p < 0.001), but data for householders were too few for testing and care should be taken in interpreting householder figures as a result. Seasonal patterns in mole control did not differ among farm enterprise tyApniemsa l(sχ220(3136) ,=6 ,2379.3, p = 0.745). Data for some categories of amenity were too few for testing. 10of26 FFigiguurere 44. .SSeeaassoonnaal lppaattteterrnnss iinn mmoollee aaccttiivviittyy aanndd ccoonnttrrooll iinn tthhee pprreevviioouuss yyeeaarr,,b byyr reessppoonnddeennttt ytyppe,eb, absaesded oonn: :(a(a) )mmeeaann rraannkkeedd mmoonntthhllyy aaccttiivviittyy ((ffaarrmmss nn == 449966;; aammeenniittiieess nn= = 116688;;h hoouusseehhooldldssn n= =1 111),)s, tsatnadndaradrd eerrrororsrs aarree sshhoowwnn; ;((bb)) pprrooppoorrttiioonn ooff rreessppoonnddeennttss ccoonndduuccttiinngg ccoonnttrrooll eeaacchh mmoonntthh iinn tthheep prreevvioiouussy yeeaar,r, bbyy reressppoonnddeennt ttytyppee (f(afarrmmss nn == 227722; ;aammeennitiiteies snn == 13123;2 h;hoouuseshehooldlds snn == 6)6. ). 3.2.6. Mole Habitats Weaskedrespondentstoreportthemonthsinwhichtheyconductedmolecontrolduringthe prevTiohuersey weaars annod epvliodtetendcet hfoerp ar orpeloarttiioonnshoifpr ebseptwoneseens tfhoer ehaacbhitmatso nptrhes(eFnigt uorne r4ebs)p.oCnodnetnrotsl’ plaanttde rannsd ebitrhoeard mlyofloel lpoewset dstaactutisv iotyr rpeactetenrtn ms.oOlev aecratilvl,itcyo,n ftorro lfarermposr t(eχd2(8b) y≤ a1l2l.r2e, spp o≥n 0d.1e4n4ts), vaamrieenditoievse r(χth2(e4) y≤e 2a.r2, p( χ≥2 (01.16)9=5)2 o5r9 .h7,opus<eh0o.0ld01s )(.χP2(a5)t t≤e r2n.6s, inp ≥co 0n.4tr5o9l).d Hiffoewreedvebre,t whaebeintaftas rwmietrhs raenpdoratmede nreitcieenst( χm2o(1l1e) =ac6ti7v.i0t,y p<0.001),butdataforhouseholdersweretoofewfortestingandcareshouldbetakenininterpreting householderfiguresasaresult. Seasonalpatternsinmolecontroldidnotdifferamongfarmenterprise types(χ2 =27.3,p=0.745). Dataforsomecategoriesofamenityweretoofewfortesting. (33) 3.2.6. MoleHabitats There was no evidence for a relationship between the habitats present on respondents’ land and either mole pest status or recent mole activity, for farms (χ2 ď 12.2, p ě 0.144), amenities (8)

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The Wildlife Conservation Research Unit, Department of Zoology, The householders about moles and mole control on their land in 2007, post
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