111 ) e c Oryctolagus cuniculus In the UK, rabbit damage was recorded 5 ; pest species; small mammals Carlos Rouco, c,d ) and environmental and anthropogenic variables Juan Mario Vargas f Correspondence to: M Delibes-Mateos,Ecología, Departamento Facultad de de Biología, Biología Universidad Vegetal deSevilla, Sevilla, . y E-mail: Apartado [email protected] 1095, 41080, Departamento de Biología Vegetal ydad Ecología, de Facultad Sevilla, de Sevilla, Biología, Spain Universi- Instituto de Estudios Sociales Avanzados (CSIC), Córdoba, Spain Departamento de BiologíaMálaga, Málaga, Animal, Spain Facultad de Ciencias, UniversidadBiogea Consultores, de Málaga, Spain Departamento de Zoología, Campus deCórdoba, Rabanales, Spain Universidad de Córdoba, Division of Biology and Conservation Ecology,ronment, School Manchester of Metropolitan Science University, Manchester, and UK the Envi- Oryctolagus cuniculus f ∗ c a e d b by large numbers of thecan weaken European or rabbit kill ( even persistentburrowing leafy can crops, and degrade scratching and establishment still of further weeds. by encouraging the 3,4 Suc- John E Fa, 1 c Oryctolagus cuniculus The latter 2 Close grazing Miguel Ángel Farfán, wildlife conflict 4 − a,b* b wildlife conflict; landscape change; − wildlife conflict. − Ana Luz Márquez, c animal conflict involving large mammals is wildlife conflict can be both direct, includ- : 111–119 www.soci.org © 2017 Society of Chemical Industry − − 74 2018; Google search; human wildlife conflict are increasingly being developed.

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Although human ing disease, injury andanimals, death and from indirect, encountersdamaged such with infrastructure. as dangerous loss Measureshuman of for crops preventing and or reducing and Pest Manag Sci These species oftenwell cause as ground substantial and damage treeexample, outbreaks crops of to rodents through in -growing grazing areas or incan as Germany browsing. cause For up to €25 million per year of damage. Negative impacts of wildlifeoccurred on humans throughout or prehistory theirquences resources and of have human recorded history. Conse- 1 INTRODUCTION Abstract BACKGROUND: Numerous small andto medium-sized crops mammal all over pestsare the cause fewer globe. widespread However, studies and mostanalysis concentrating economically research of the on on significant ecological and pest damage the anthropogenicbetween species factors agricultural description correlated has damage with focused of by these observed the on patterns. crop European We accounts rabbit investigated damage ( of the relationship the caused level by of pests damage. There at large geographical scales, or on Jesús Olivero, and Rafael Villafuerte damage to agriculture in Spain Miguel Delibes-Mateos, (wileyonlinelibrary.com) DOI 10.1002/ps.4658 A large-scale assessment of European rabbit Research Article Received: 28 December 2016 Revised: 31 May 2017 Accepted article published: 19 July 2017 Published online in Wiley Online Library: 6 September 2017 cessful conflict resolution plans are oftenlation the of result proven of methods the of assimi- controladaptation or to deterrence and the their clear nature of the problem at hand. requires an accurate assessment ofsituation the investigated, human and a clearcost-effectiveness understanding and of social the acceptability efficacy, applied. of the methods to be often highlighted in themuch media, more human livelihoods impacted are by arguably small and medium-sized mammals. Keywords: Supporting information may be found in the online version of this article. throughout Spain. RESULTS: Rabbit damage was mainly concentratedincreased within significantly the between central-southern the regions early of 2000sand Spain. and natural We 2013. vegetation found Greater was that losses scarce, rabbit where were damage main typicalgenerally railways of favourable and for those highways rabbit areas were populations where present, to and farming proliferate. where dominated environmental conditions were CONCLUSION: From our analysis, wespread. The suggest recent that increase roads inapproach and is Spain valuable railway as of a lines such method for act infrastructure assessingmethods drivers as to may of reduce wildlife potential explain human pest the damage corridors at rise along large© spatial in 2017 which scales, Society and rabbit rabbits can of be damage Chemical can used reported Industry to propose in this study. Our et al. The first Searches : 111–119 25 24 74 ha, respectively). 2018; 6 2014. The first 100 10 − × We selected this word 5 24 ∼ (nearly 85% of the Iberian 2 Pest Manag Sci ha versus 6 Mountains border the northern and 10 × 22 12 ∼ 23 The natural vegetation in the mountains is pine and oak 2.2 Temporal andcaused spatial by rabbits characterisation of damage We used thesearch Google terms “pest” and search “rabbit” (in enginerespectively) Spanish: to “plaga” (www.google.es) identify and websites “conejo”, containing and informationaffected on the areas by European rabbits in Spain. 22.1 MATERIALS AND METHODS Study area Peninsular Spain covers 493 518 km 600 consecutive sites resulting fromited our and their Google content search evaluated by were M. vis- Delibes-Mateos. crops and trees represent justand over woody 50% of crops, arable respectively. land Fodderarea crops of cover arable an important land,woody while crops. fruit trees and are common combination based on apermutation, previous but study also which “rabbit” and showed “damage”,“rabbit” that worked and this better “crop”, than and “rabbit” and “overpopulation”. Mediterranean coasts (maximum altitudealso 3478 east–west m) facing but ranges there in the are centre of the Peninsula. and scrubland, butceous lowlands crops are and mostly riversuch dominated as terraces, olive. and by According to hill herba- theture slopes 2010 Survey European by Union (FSS), woody Farm the Struc- of crops area the under whole agriculturenomically in territory active Spain population and was of involved 47% eurostat/web/agriculture/farm-structure). the Arable country around land (http://ec.europa.eu/ 9.8% dominates over of woody crops the ( eco- Peninsula). The country is divided into 8125 administrativepalities munici- (http://www.ine.es), the territorial units used in theanalyses. current Climate varies widelytation throughout decreasing Spain, eastwards withincreasing and precipi- southwards, southwards. and temperature were performed between 18 May and 11 Juneunequivocally 2015. containing Only websites information on damageEuropean rabbit caused in by Spain were the considered valid. Theseincluded sites media mostly reports on rabbithunters’ damage, and discussion farmers’ websites, forums and on relation blogs. to Websites rabbit that damage bore in no ruled Spain out were for discarded. Among analyses those wereregions websites where dealing the with species pest is rabbitsfor in an pet invasive rabbits. alien We or gatheredtural advertisements information crop on damaged by the rabbits. typedetermine As of areas one agricul- where of rabbit ourmented damage main those goals occurred, municipalities was we where to To also such assess damage docu- the was temporal reported. tion evolution on of rabbit the damage annual inwebsite was online Spain, uploaded. we informa- recorded Because the internetincreased year in usage recent that has years, each substantially thusreports affecting on the rabbit availability damage, we of usedGoogle media the Advanced date Search function to availablewords in conduct filtering a by search year ofsites for the yielded the by target each period key- search (i.e. 2004 forand each specific those year) were that visited were validised index were of selected available information to on calculate rabbiti.e. damage a for in standard- each each year; year, theited. number The of temporal evolution valid of websites/100 suchthe websites an evolution index vis- of was compared the with reports ongeneral search rabbit (see damage above). obtained from the 21 )in 18–20 www.soci.org M Delibes-Mateos Orgyia pseu- )intheUSA, © 2017 Society of Chemical Industry The control of the 16 Myocastor coypus There are few studies 13 By contrast, rabbits cause and around €130 million per 17 6 Halyomorpha halys Rabbits in Iberian Mediterranean 16 and by coypu ( 8,9 Given the ubiquity of transport infrastruc- 14 15 7 150 million in 1986, 11 − However, for some species, particularly insect pests, ), eruptions correlated primarily with type and 12 18 have been undertaken. However, analyses of the spatial 10 Knowing where and when rabbit damage occurs in Spain and Assessing crop damage caused by indigenous small mammal Pest outbreaks can be linked to specific environmental condi- Country-wide assessments of the damage caused by rabbits ture in many partsreach of remote the areas world, as itrailway they is lines likely establish where that themselves they pest along canthey species roads become can proliferate. will abundant, or and from which which factors may drive it can helpvarious mitigate stakeholders. In conflicts this between study, the we first describethroughout rabbit damage Spain, untilareas now (i.e. municipalities) unrecorded, where rabbit by: damageto has (1) occur; been (2) reported documenting recording which crops arethe affected; and temporal (3) evaluating evolution ofmultiple rabbit sites, we damage then assess in the probability the ofdamage occurrence country. caused of Across crop by rabbits, andtions and its environmental relationship factors, such with as localtion land or condi- use, infrastructure. natural To vegeta- do this,of we environmental employ and large-scale biotic databases variables, and use spatial modelling. pests, as well as increasing our understandingthis, of is what determines important whenrole the in species a in native ecosystem. questionbit This also in is the plays the Iberian a case Peninsula, key for its the native European range. rab- dotsugata tions. In the case of the Douglas-firclimate. tussock moth ( in the UK and Australia on the rolewhich of pest roads species and maybrown disperse. railways An marmorated as exception stink is potential bugwhich a corridors study showed ( along of that the urbanthe development species’ and spread. railroads facilitated wileyonlinelibrary.com/journal/ps Our results are usedto to determine rabbit whichwith areas damage this. are and more prone assess which factors are correlated and are regardedMoreover, as rabbits a arespecies, one severe hunted of pest in the speciesThese their multiple roles most often in lead millions to important frequent some conflicts in betweeners, small-game farm- the regions. hunters and two conservationists Iberianspecies. regarding how countries. to manage the as being €115 year in Australia. (where damage happens) andfactors determining temporal the (when crop damage damage inflicted byspecies occurs) a have particular pest been largelySuch missing from approaches large-scale allow assessments. forwhat the variables may application correlate of withdamage understanding a and species’ of thus propensity can to permitmeasures cause managers of to control, devise more especiallysmaller effective demonstration as scale what can fail maylandscape when be level. applied at successful the at broader a ecosystems act as abecause they multifunctional are keystone prey species, fortheir over not role 30 as only predators, an but ecosystem also because engineer. of transport infrastructure (roadsmit and their railway dispersal lines) and can establishment. also per- rabbit here needs to be effected inits such unrestrained a way decline that or does extinction; noting cause this effects could in produce cascad- the ecosystem. substantial damage to agriculture in Spain and ,

112 113 180). In this = n Stepwise 30 31 180). Rabbit = n 2006, but cases − (which indicates global 301), or human properties R = n 301). Most rabbit damage was = n wileyonlinelibrary.com/journal/ps ) of each variable with the favourabil- R The regional relevance of every variable was, 32 0.001) between the number of websites containing < P 180), although it was also relatively frequent in olive = n 0.971; 301), with only a few documenting damage to infrastructure = The significance of the influence of all variables in the model = R such as gardens or parks (9.6%; reported for vineyards and cereal crops (66.1%tively; and 53.3%, respec- groves and fruit (28.3% and 21.7%,Green respectively; and other crops were lesson frequently mentioned the websites (15.0%damage and was 9.4%, reported respectively; inland 437 Spain. (5.5%) municipalities These acrossEbro were valleys main- in mostly southern found andCastilla-La northeastern in: Mancha Spain, respectively; (1) and (2) Castilla Guadalquivirand y and León (3) plateaus the in coastalValenciana central eastern region Spain; (Fig. regions, 2). particularly the Comunidad 3.2 Modelling favourabilityThe for favourability European rabbit model damage for the presence ofage European included rabbit 18 dam- different variables relatedture, to transport land infrastruc- use,(Table natural 1). vegetation The and modelpresence environmental showed of variables that European rabbit municipalitieseach damage favouring other tended the (Fig. to 3), be located found mainly near along the and 3RESULTS 3.1 Temporal andcaused spatial by rabbits characterisation of damage The general Google searchwebsites containing yielded information a on damage total causedrabbits by of in European 306 Spain. unique, Thein first valid cases 2004. of There rabbitincreased damage were sharply were few reported thereafter until casessharp 2013, decline during (Fig. after 1). 2004 In which 2014, therewebsites we containing recorded was the information a highest on number rabbit of damage,els reaching higher lev- than in 2012. There( was a strong significant correlation information on rabbit damage and year when weresults standardised our using the Googleysed search websites by reported date crop option.n damage Nearly caused all by anal- rabbitsaround railways (95.7%; or highways (10.3%; way, we specified how muchaccounted of the for variation in by favourabilitythat the was is not pure affected effect byproportion covariation of with was another clearly each factor), attributable and factorshared to what effect). more (i.e. than variation one factor (i.e. was assessed using the univariate Wald test statistic. thus, analysed using twocorrelation approaches. (Spearman Firstly, we measured the methods select variables acting on aand larger at scale subsequent in the steps first onlyrelated steps, add to variables the that residuals are notrated significantly accounted for variables. by previously incorpo- infrastructure, natural vegetationexplanation and land of uses) favourability in for the damage model occurrence. ity output, and comparedrelationship the within sign the of studycoefficient area) in with the model the equation sign (whichvariable indicates of the contribution the sign to variable of the visualised explaining the favourability). regional contribution Secondly, of each we by variable mapping to the the difference model between favourability valuesin obtained successive steps, along the stepwise variable selection. 26 27 1and n ]} ) P − 1 was calculated with ( P ∕ P is the probability of rab- [ P 0.05 were accepted as part + ) < 0 q n ∕ Atlas of Terrestrial Spanish Mam- : 1 n ( { 26,27 ∕ ] ) P − : 111–119 © 2017 Society of Chemical Industry We considered 35 predictor variables (Table 1 Thus, using the presence/absence of European ( 74 28 . ∕ 29 P [ 2018; = F is the favourability value in each municipality; 0.8. Only then did we implement the favourability model F > r 0.05) variables under an FDR of < Overall, our model of European rabbit damage included 136 pre- We excluded nonlinear and interaction effects from the model Firstly, a model of environmental favourability for the presence of 0 are the number of municipalities with and without damage P using the following equation of a multivariate environmental model.minimised Multicollinearity by was excluding variables also that were correlatedman with Spear- S1). The ecological factorsfavourability that of could the explain European thetopography. rabbit environmental Alongside were these, based we on includedland climate descriptors use and of and human humanimpacts. activity to assess potential anthropogenic dictor variables (Table S2): the favourability model for theof presence European rabbits, asmental a conditions variable that representative determine the of presenceecosystem the type of descriptors environ- the based species; on natural 79 vegetationand structure, 56 indicators of anthropogenicinfrastructure activity (i.e. in main terms of railwayslivestock. transport and Spatial highways), analyses agriculture toinfrastructure and calculate were the performed presence using of ArcGISCA, transport 10.0 USA). (ESRI, Redlands, to keep its mathematicalexplanatory purposes. formulation To as account for simplelarge Type number I as of errors possible variables caused considered, by for wecovery the controlled rate (FDR). the false dis- Pest Manag Sci Thus, compared to logisticity function regression, we by cancel using outabsences uneven the in proportions favourabil- the of presences modelledmodel and data. the Here, relationship weenvironmental between variables. use European this rabbit approach damage to and the European rabbit in each10 Universal Transverse x Mercator 10 (UTM) km squaredistribution of map mainland available Spain in was the constructed using the The favourability function assessesmental the extent conditions to change whichorganism environ- the with respect probability to its of overall prevalence in occurrence the study of area. an mals and Red Book Agricultural damage by rabbits in Spain2.3 Predictive model To model the potentialthroughout distribution mainland of Spain, we European used rabbit the favourability damage function. www.soci.org rabbit damage as the dependent variable, wesion ran a on logistic each regres- of( the 136 predictor variables, and only significant n reported in websites, respectively; and where 2.4 Explanatory analysisWe of variables employed in a the variation model partitioningrelative procedure participation to of measure the the S2 four main (favourability factors for described the in presence Table of European rabbit, transport bit damage presence in eacha municipality. multiple logistic regression, usingdamage the in presence each and municipality absence according of as to the the dependent variable. analysed A websites forward stepwiseformed procedure was to per- avoid the inclusionThe of parameters in redundant the models variables were fitted in by athe iteratively maximising model. log-likelihood (using IBM statisticsMachines, New 22, York, USA). International Business et al. hunted : 111–119 74 2018; Pest Manag Sci ). 46 The greater proportion of the spatial variation in favourabil- et al. correlated with the occurrencethey of showed European identical rabbit signsthe damage, variable within i.e. correlations the with the modelmodel model, equation produced and their and an entry increase in intoforests in the (especially favourability (Fig. mature 4). oak Inrepresented environmental forests), contrast, constraints to scrublands the and occurrence ofpean pastures rabbit Euro- damage, i.e. both signs within thevariable model equation correlations and within theentry model into the were model negative, produced a and decrease in their favourability (Fig. 4). ity was explained by habitat descriptors, represented by natural www.soci.org M Delibes-Mateos © 2017 Society of Chemical Industry 25) were primarily transport infrastructure and > Municipalities reporting European rabbit damage within the study area (in white) according to the analysed websites (see text for details). Evolution of the number of websites containing information about European rabbit damage according to a Google search (solid black line), the Variables with the highest explanatory power within the model wileyonlinelibrary.com/journal/ps valleys, several areassouthern of plateaus the and some Guadiana coastal valley, areassular of northern Spain. southeastern and penin- (Wald statistic Figure 2. Figure 1. total length (km) of high-speed railwayin in Spain Spain (dotted (grey line; line; source: source: http://www.magrama.gob.es http://www.spanishrailwaysnews.com), and and the Delibes-Mateos number (in millions) of rabbits land use descriptors (Table 1).lands Humanised and habitats, infrastructure, such and environmentally as favourable crop- areasthe for presence of European rabbits wereage. In positively contrast, related natural to vegetation was dam- negatively associatedrabbit with damage. More specifically, highways and mainand railways, irrigated dry olive crops, and dry and irrigated vineyards positively

114 115 % and its, rest 18,24 SE,E,NE Despite this, media reports have 33 wileyonlinelibrary.com/journal/ps -SW + Thus, we argue that, even if there is some error 34 ++ ++ ++ ++ ++ ++ ++ ++ ++ ++ + Our study demonstrates that the modelling approach we use can According to the analysed websites in Spain, most rabbit dam- likely to be missed using other meansour of model verification. For clearly example, showedregion that areas (northwestern such as Spain)values the have even Castilla y relatively though León thesearchwaslow(Figs2and3). high number favourability of cases found in ourbe Google useful in determining thetial role scales of multiple in factors explaining at theaddition, large damage we spa- show caused that by use a ofage pest online can species. media be In reports an on effectivethe and pest distribution dam- relatively of rapid pest damage option in to a landscape understand country. as However, large a as potential an drawback entire of thisreports information in is the that media onholders’ pest perceptions, but damage perceived may and only actual record damagewildlife caused the may by stake- not be correlated. involved, the information obtained in ourareas of study potential can stakeholder conflict help regarding identify theof management pest species (farmers and hunters in our study model), been successfully used previouslymammal to pests. assess the impact of small can be used in the design of more realistic management plans. age occurred in crops. This is inwhich line with favourable the areas result of for our rabbitlated model with in damage several were land positively uses corre- relatedcrop to types agriculture; were many among different thedamage most (Table 1). important Our predictors results ofa suggest rabbit that, generalist in agricultural Spain, pest the rabbit as is a variety of crops are damaged area); F_WILDRABBIT, favourability model for the presence of European rabb : 111–119 © 2017 Society of Chemical Industry 74 0.05); CfS, sign of the variable coefficient in the model; CrS, sign of the correlation (Spearman) between the variable and favourability < 2018; P Descriptor variables of the environmentally favourable areas for European rabbit damage according to the favourability model based on climate, topography and humanolive activity tree (Table forest S1); (% ORG_CER, area); organic P_UAA,of proportion cereal oaks (% of useful area); (height); WOLIV_COP, agricultural PAST, beech pastures area forest (% (% (% area); area); area); DIV_FOR, NOAK_CLEFOR, WOLIV_FOR, carob diversity wild forest of (% forest area); uses DEV_OAK, and development vegetation phase structure (Simpson diversity index); PINE_FOR, birch fo (% area); GALL_FOR, alignment (% area); OAK_CLEFOR, boxwood forest (% area); PAST_SHRUB, pastures and shrubs (% area). Step, order of entrance insignificant at the model; W, univariate Wald test statistic quantifying variable significance in the model (all the variables shown were HIGHW_RAILWIRR_OLIVDRY_VINIRR_VINF_WILDRABBITORG_CER 1WOLIV_COPWOLIV_FORP_UAA 7NOAK_CLEFOR 8 11Variables describing unfavourable areas for European 4 rabbitDEV_OAK damage 119.444 PAST 10 9DIV_FOR 13PINE_FOR 51.344 6GALL_FOR 15.961 43.624 14OAK_CLEFOR 26.387 PAST_SHRUB 13.372 5 9.238 7.006 12 2 9.686 17 4.157 16 18 15 32.730 7.016 6.276 5.619 5.294 4.075 9.360 ------VariableVariables describing favourable areas for European rabbitDRY_OLIV damage Step 3 W 124.834 CfS CrS AGI Table 1. area); DRY_VIN, dry vineyards (% area); IRR_VIN, irrigated vineyards (% values; AGI, area of geographicalDRY_OLIV, dry influence olive of groves (% the area); HIGHW_RAILW, length variable of in highway and the high-speed railway model per within municipality (m); the IRR_OLIV, irrigated study olive groves area ( (SW, southwest; NE, northeast; E, east; SE, southeast). Pest Manag Sci We showcentral-southern that regions of rabbit Spain,(Fig. 2). and We damage in also somerabbits demonstrate northern is in that an areas the area more likelihoodour depends of model on common damage has a by occurs combination in clearly in municipalities of where identified factors, farming the natural dominates (Fig. which 5). (and vegetation therefore is Most scarce), whereare rabbit main present damage and railways where and environmental highways ulations conditions for to rabbit succeed pop- approach, are we generally were able goodgreater risk to (Table of highlight rabbit S2). those damage Using at areas a potentially our large at national scale, which are 4 DISCUSSION vegetation and land use factors.effects The of sum of these the factors pure53.7%, on and respectively favourability shared (Fig. variation 5). was Transport 81.8%mental infrastructure and favourability and environ- for theexplained presence 8.4% of and Europeanshared 10.6% effects rabbits between of the only different variation, factors (meaning respectively.or either indistinguishable cross explanatory We power). Land found uses andture infrastruc- shared 90.3% ofvegetation shared their 40.8%. influence, The andtors other land were shared considerably use effects smaller. between As and aural fac- natural result, vegetation the explained pure effect 39%of of of nat- damage, favourability whilst for thefavourability the for presence pure the presence effect of Europeanture of rabbits, explained and only land infrastruc- 10.0%, 3.0% uses, and 2.3%, environmental respectively. Agricultural damage by rabbits in Spain www.soci.org 40 and et al. 18 It also 18 : 111–119 74 Similar reasons or the intensive The increase in 2018; 44 46 42,43 Verges and embankments In addition, these verges and Pest Manag Sci 41 39,41 particularly in areas within farmland 39,40 This may also explain the high importance of particularly in some agricultural areas. 36 47,48 In farmland areas, rabbits usually establish dense 45 Our Google searches uncovered very few sites containing infor- explanation is that rabbitsthese can become infrastructures, more abundant close to populations along the vergesnearby of crops roads are their and onlyhas railway feeding lines, almost source where disappeared as natural astural vegetation a practices. consequence of intensive agricul- regions where the availabilitysuccessful of other establishment suitable is habitat for low. their human infrastructure in explaining rabbit damage observedmodel. in the mation related to rabbit pests2000s. in Spain This during was the a first timetheir half when lowest of Spanish the numbers rabbit in populations recordedhad were history not at recovered because from most the ofdisease huge them impact (RHD), of first rabbit detected haemorrhagic in Spain in 1988. embankments offer ideal conditions for rabbit warren building, essential to establish healthy social groups. use of areas adjacentspecies. to roads by juveniles of two lagomorph have been proposed to explainmammals the found in high areas numbers with of high road some density small matches the increase inand the the number number of of harvested rabbits rabbit recorded control in requests Portugal the number of websites2012 matches reporting the damage known betweenthat recovery period, 2006 of and some populations during along roads or railwayschain-mesh (for security high-speed fenced) railways provide thesepredators rabbits are and with often hunters, refuge as predator against abundanceing is often lower banned and in hunt- these areas. in Spain (Fig. 1). These findings closely resemble the situation in 38 98% of > www.soci.org M Delibes-Mateos © 2017 Society of Chemical Industry 0.001). According Using the analysed < ). This contrasts with 35 P 20 et al. 0.904; = R thus decreasing the likelihood of damage to 37 Given that a vast proportion of agricultural land in some Favourability model for European rabbit damage in each municipality of mainland Spain. The small map shows the environmental favourability – losses caused by rabbits to and producers 21 36 Although the number of hunters in Spain has remained rel- Natural vegetation explained a large proportion of favourability wileyonlinelibrary.com/journal/ps typically damaged by rabbits. atively constant (http://www.magrama.gob.es),have rabbits increased hunted alongsideduring the the last spread decade of (Fig.1; high-speed railways to our model,was one the presence of of the railways or main motorways predictors (Table 1; of Fig. 4). rabbit The damage for the presence of European rabbit in Spain, on UTM 10 x 10 km squares. The scaleby ranges this from 0 species (black) (see to 1 also (white). Ríos-Saldaña Figure 3. can be economically crippling. for rabbit damage in our model (Table 1). Thevegetation fact that types most natural were negativelyoccurrence of associated rabbit damage with is not the surprising,the because potential proportion the of higher land occupiedthe by natural availability vegetation, the of lower rabbits crops are found to in low be numbers indense grazed areas scrublands, dominated by by forests rabbits. or Furthermore, other Spanish smallcrop mammal types; e.g. pests voles and that herbaceous crops. are linked to specific regions of Spain is devoted to these woody crops – e.g. websites, we showed thatrecorded crop vineyards type affected were by thegroves rabbits, had and most both significant frequently these effects andthe on olive model (Table rabbit 1). Notable damage reductions favourability inrabbit woody in browsing crop yields have from been reported inregions. some Spanish agricultural adjacent crops. Similarly, we observed thatlikely rabbit in damage areas is environmentally most favourable for theIn species (Table these 1). areas, highbeing rabbit positively densities associated with are the possible presence in of addition woody to crops, the area in the Montilla-Moriles wine-growingSpain region in southern

116 117 r est rea); groves ). This may be 52 suggest that rab- et al. 18,20 wileyonlinelibrary.com/journal/ps 35 0.001). An alternative explanation is that the < P The role of humans in the expansion of small 36 0.869; = R Economic damage caused by mammal pest species can be high, The presence of transport infrastructure was one of the main the observed rise infor rabbit sampling damage effort reports per when yearhunted we and and rabbit the controlled control growing permits requested numbers of rabbits reduction of other foodcrop sources for damage. rabbits has led to increased particularly in farmland areas (e.g. Pimentel mammal pests and damage by these pests hasin been other also observed farmland systems. a source ofaffected conflict by between wildlife, such stakeholders as farmers, whose and other livelihood parties is that make bit damage has in factcase, we do increased recommend that in online data recent be years interpreted cautiously. in Spain. In any explanatory factors ofextensive rabbit development of damage high-speed railways in andhas highways our that model. occurred Given inlandscape the Spain modification may in havein recent been the one years, of rise we the into main suggest rabbit be drivers damage that supported reported such the by in increase the in our the significant study. numberand This overlap of websites the seems observed dealing increase with between in rabbit(Fig. 1; pests the length of high-speed railways in Spain It is very likely that 10-fold between 2003 − 48,50,51 Also, the considerable increase in the use 18 : 111–119 © 2017 Society of Chemical Industry 74 2018; Our findings also show that in 2013 the available 49 Mapped contribution of variables to the model of favourability for European rabbit damage along the stepwise variable selection. Step, order of 51 Our findings indicate that the evolution of online reports on rab- bit damage might be a good indicatorulation of and trends rabbit in damage at the a rabbit large pop- data scale have some in potential Spain, limitations. For although example, such the relationship between rabbit abundance andto damage crops is by complex, rabbits as hasate damage been rabbit also abundance. observed in areas with moder- Pest Manag Sci IRR_VIN, irrigated vineyards (% area); DEV_OAK, development(% phase of area); oaks DRY_VIN, (height); NOAK_CLEFOR, dry carob forest vineyards (%the (% area); presence IRR_OLIV, area); of irrigated European olive WOLIV_COP, beech rabbits, forest based on(% (% climate, area); area); topography P_UAA, ORG_CER, and human proportion organic activity cereal ofbirch (Table (% useful S1); forest area); PAST, (% agricultural pastures F_WILDRABBIT, area); (% area favourability OAK_CLEFOR, area); (% boxwood WOLIV_FOR, model area); wild forest fo olive (% PAST_SHRUB, tree pastures area). for and shrubs (% area); GALL_FOR, alignment (% area); PINE_FOR, Australia, where rabbit numbers were heavilybetween suppressed 1995 by and RHD 2002, but increased 5 Figure 4. entrance in the model; green, negativerailway contribution to per damage; municipality red, positive (m); contribution DIV_FOR, to diversity damage; HIGHW_RAILW, of length of forest highway uses and and high-speed vegetation structure (Simpson diversity index); DRY_OLIV, dry olive groves (% a and 2010. Agricultural damage by rabbits in Spain www.soci.org online information related toseems rabbit to damage fell beby sharply, only which a explicable new by variant the of population the crash RHD caused virus. the decline in rabbitsreduced observed rabbit in damage, agricultural whichber areas was of reflected resulted websites in in addressing theof this lower issue. num- websites Interestingly, reporting thebetween number rabbit 2013 damage andpopulations might 2014 increased have (Fig. considerably recovered 1),areas, at although which least in in suggests naturallow. some areas that agricultural rabbit rabbit numbers were still very of the internet over recent yearsability might of explain online the information regarding growing almost avail- any topic. However, Pest et al. ,ed.by :50–54 ion and : 111–119 109 74 2018; :70–80 (2016). :2365–2376 (2016). 5 New Scientist The economic impacts of 18 :159–171 (2007). 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Values show the proportions of how much of the variation in environmental favourability for European or Co-existence? Kingdom (2005). 1 Woodroffe R, Thirgood S, Rabinowitz A, wildlife damage at large spatialwhere scales potential conflicts as may well emerge. as The approach tothis presented paper identify in can areas be reproduced in otherwildlife systems, pest not damage, only to but reduce alsobetween to stakeholders regarding work the towards management reducing of tension wildlife. wileyonlinelibrary.com/journal/ps REFERENCES SUPPORTING INFORMATION Supporting information may be found inarticle. the online version of this land use factors, and which proportion was attributable to their shared effects (intersections). ACKNOWLEDGEMENTS We wish to thankcomments. three M. anonymous Delibes-Mateos reviewersby and for the constructive Talentia C. Postdoclaunched Rouco by and the were Talent Andalusian Hub supported by Knowledge Program, Agency, the and respectively, co-funded EuropeanSkłodowska-Curie actions Union’s (COFUND – Seventh Grant Agreements Framework n267226 and Program, 291780, Marie respectively)vation, and Science the and Ministry EmploymentDelibes-Mateos of of is the currently Economy, Junta funded Inno- gación de by of the Andalucía. V Universidad de M. Plan Sevilla. This Propiothe study projects is de CGL2013-43197-R a (I Investi- contribution of Spanish Ministry of Science and Innovationand and 1098/2014 (Organismo EU Autónomo Parques Nacionales, Spain). rabbit damage occurrence is explained exclusively by infrastructure, environmental favourability for European rabbit presence, natural vegetat other uses of these species, suchOur as case hunters or study environmentalists. isthe world. a good example, but there are others across Figure 5.

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