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REVIEW PAPER A review on the impacts of feral cats (Felis silvestris catus) in the Canary Islands: implications for the conservation of its endangered fauna Fe ´lix Manuel Medina Manuel Nogales Received: 4 February 2008 / Accepted: 8 October 2008 / Published online: 21 October 2008 Ó Springer Science+Business Media B.V. 2008 Abstract Feral cats have been directly responsible for the extinction of numerous species on islands worldwide, including endemic species of mammals, birds and reptiles. The diet of feral cats in the main habitats of the Canary Islands, as generally occurred on oceanic islands, is mainly composed of introduced mammals, and native species of birds, reptiles and insects. The impact of feral cat upon the endangered species was assessed by evalu- ating their relative abundance in the cats’ diet and by considering their current conservation status. A total of 68 different preys were identified at species level in all studies carried out in the Canary Islands (5 mammals, 16 birds, 15 reptiles and 32 invertebrates). From all the species preyed by feral cats in the Canary Islands, only four of them are considered threatened by the IUCN Red List of Threatened Species: one endemic bird Saxicola dacotiae and three endemic giant lizards, Gallotia simonyi, Gallotia intermedia, and Gallotia gomerana. Although some efforts on management control have been carried out, it is necessary to enforce these conservation activities on those areas of Tenerife, La Gomera and El Hierro where giant lizards are still present. Furthermore some local areas where endangered bird species are highly predated should be protected. Nevertheless, it is important to take into account the presence of other introduced species such as rats, mice or rabbits in order to avoid problems derived from the hyperpredation process and meso- predator release effect. Keywords Canary Islands Á Endangered species Á Felis silvestris catus Á Food habits Á Island conservation F. M. Medina (&) Consejerı ´a de Medio Ambiente, Cabildo Insular de La Palma, Avenida Los Indianos 20, 28, 38700 Santa Cruz de La Palma, Canary Islands, Spain e-mail: [email protected] F. M. Medina Á M. Nogales Island Ecology and Evolution Research Group (IPNA-CSIC), Astrofı ´sico Francisco Sa ´nchez 3, 38206 La Laguna, Tenerife, Canary Islands, Spain 123 Biodivers Conserv (2009) 18:829–846 DOI 10.1007/s10531-008-9503-4

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REVI EWPAPERAreviewontheimpactsofferalcats(Felissilvestriscatus)intheCanaryIslands:implicationsfortheconservationofitsendangeredfaunaFelixManuelMedina ManuelNogalesReceived:4February2008 / Accepted:8October2008 / Publishedonline:21October2008SpringerScience+BusinessMediaB.V.2008Abstract Feral cats have been directly responsible for the extinction of numerous specieson islands worldwide, including endemic species of mammals, birds and reptiles.The dietofferalcatsinthemainhabitatsoftheCanaryIslands,asgenerallyoccurredonoceanicislands,ismainlycomposedofintroducedmammals,andnativespeciesofbirds,reptilesandinsects. Theimpactofferalcatupontheendangeredspecieswasassessedbyevalu-ating their relative abundance in the cats diet and by considering their current conservationstatus. A total of 68 different preys were identied at species level in all studies carried outin the Canary Islands (5 mammals, 16 birds, 15 reptiles and 32 invertebrates). From all thespecies preyedbyferal cats inthe CanaryIslands, onlyfour of themare consideredthreatened by the IUCNRed List of Threatened Species: one endemic bird Saxicoladacotiae and three endemic giant lizards, Gallotia simonyi, Gallotia intermedia, andGallotia gomerana.Althoughsomeeffortsonmanagementcontrolhavebeencarriedout,it is necessarytoenforce these conservationactivities onthose areas of Tenerife, LaGomeraandElHierrowheregiantlizardsarestillpresent.Furthermoresomelocalareaswhere endangered bird species arehighly predatedshould be protected. Nevertheless,it isimportant to take into account the presence of other introduced species such as rats, mice orrabbits inorder toavoidproblems derivedfromthehyperpredationprocess andmeso-predatorreleaseeffect.Keywords CanaryIslands Endangeredspecies Felissilvestriscatus Foodhabits IslandconservationF.M.Medina(&)Consejer adeMedioAmbiente, CabildoInsulardeLaPalma,AvenidaLosIndianos20,28, 38700SantaCruzdeLaPalma,CanaryIslands,Spaine-mail:[email protected] M.NogalesIslandEcologyandEvolutionResearchGroup(IPNA-CSIC), Astrof sicoFranciscoSanchez3,38206LaLaguna,Tenerife,CanaryIslands,Spain1 3BiodiversConserv(2009)18:829846DOI10.1007/s10531-008-9503-4IntroductionSince its domestication from the African wildcat (Felis silvestris lybica) some9,000yearsago(Randi andRagni 1991; Serpell 2000; Driscoll et al. 2007), cats(Felissilvestris catus) has long been associated with humans (Vigne et al. 2004) and hastraveled with themto most parts of the world (Fitzgerald 1988). Their generalistbehaviour converts this introduced predator into a very adaptable species which iscapable of surviving in inhospitable conditions on the most remote oceanic islands(Courchampet al. 2003). Onthese, feral cats preyona highvarietyof prey, mainlyintroducedmammals, althoughtheyincludeintheirdiet anelevateproportionofnativebirds, reptiles andinvertebrate species (Fitzgerald1988; FitzgeraldandTurner 2000).Oceanicislands arereallyrelevant inthecontext of wholeworldbiodiversitybecauseharboranimportant numberofendemicspecies(MacArthurandWilson1967; Gorman1979; Whittaker and Fernandez-Palacios 2007). On its, feral cats have been directlyresponsiblefortheextinctionofnumerousmammals(Mellink1992;Tershyetal. 2002),reptiles(Iverson1978;Mitchelletal.2002)andbirds(JehlandParks1983;Lever1994;Veitch2001). For this reasonthis predator is consideredtobe one of the 100worstinvasivespeciesintheworld(Loweet al. 2000).The Canary Islands is an important hotspot of global biodiversity because 2,985animal and524plant speciesareendemicelements(Mart net al. 2005a). Nevertheless,intheCanaries, at least 1,434specieshavebeenintroducedbymanandapproximately150of themareconsideredsuchasinvasive(Mart net al. 2005a). CatswereprobablyintroducedintheCanaryIslandsinthefteenthCenturyaftertheEuropeanconquestandpresentferalpopulationsinallthemainislandsandhabitatsoftheCanarianArchipelago(Nogalesetal.2006;MedinaandNogales2007).Feralcatsarethepredatoratthetopofthefoodchain,amongthevertebratesintheseislands(Nogalesetal.1992).Thefeedingecologyof feral catsontheCanaryIslandsfollowsthegeneral patternof other islandslocatedat similar latitude, beingcharacterizedbyanimportant biomass supportedbyintroduced mammals (mainly rabbits and mice) when they are present (Nogales andMedina 1996). However, some specic preys, such as lizards, rats or birds, play animportant role in particular habitats in which these prey are available and ratherabundant.Historically, feral cats have been considered co-responsible of the extinction ofseveral vertebrate species in the Canaries such as Pufnus holeae, Pufnus olsoni,Pterodromasp., Carduelistriasi orEmberizaalcoveri althoughotherintroducedspeciessuch as rats and, overall, the effects of humans by direct consumption and habitatdestructionwerethemost probablecauseof their extinction(Rando2003). Theseandother extinct species in the Canary Islands such as Gallotia goliath, Canariomystamarani, C. bravoi or Pufnus olsoni werecommonlyincludedintheaborigines diet(Rando2003).AlthoughtheCanaryIslandsconstitutetheplacewheretheferal cats diet hasbeenmoreextensivelystudiedaroundtheworld(Nogalesetal.2006),theireffectonendemicandendangeredspecies hadnot beenevaluated. For this reasonthe mainaimof thiscontributionis toreviewtheimpact of feral cat uponendemicfaunaof theseislands,evaluating the conservation status of the different prey items consumed by feral cats in allenvironmentspresentinthisarchipelago.830 BiodiversConserv(2009)18:8298461 3MaterialsandmethodsStudyareaThe Canary Islands are of volcanic origin and are located about 100km from the northwestcoast of Africa. It is composed of seven main islands and several islets (Fig.1), the highestof whichis Tenerife(PicoTeide: 3,718ma.s.l.) andthe lowest Lanzarote (Penas delChache: 671ma.s.l.). Tenerife is the most extensive island (2,034.4km2), while thesmallest is El Hierro (268.7km2). The climate in the Canaries varies according to altitudeand orientation. Mean temperature and annual precipitation ranges from &21C and 100300mm, respectively, incoastal zones, toabout 9Cand500800mm, respectively, athigheraltitudes. TheeasternislandsLanzaroteandFuerteventura, nearest totheAfricancontinent,arelowinaltitudeandinuencedgreatlybydrywindsfromthenearbySaharaDesert.Thevegetationischaracterizedbyadryxerophyticshrubthatonlyappearsinthelowlands of the higher central and western islands of the Canaries (Gran Canaria, Tenerife,La Gomera, El Hierro and La Palma) and is characterized by species of the genusEuphorbia(Euphorbiaceae). The vegetationof the Canaries is clearlydistributedas afunction of altitude and orientation, including other more humid forest habitats. Thevegetationalsoincludesatermophyleforest, locatedat300550ma.s.l., composedofamixed wood of Dracaena draco (Agavaceae), Phoenix canariensis (Arecaceae), Juniperusturbinata(Cupressaceae), etc. Between550and1,300ma.s.l. appears themost humidhabitat, thelaurel forest, whichisconstitutedbyabout 20treespecies, several of themendemic. Some of the most relevant species are Laurus novocanariensis and Persea indica(Lauraceae), Myricafaya (Myricaceae) and Erica arborea (Ericaceae), which formadense evergreen forest. Following in altitude is a dryer monospecic pine forest(1,3002,000ma.s.l.) of theendemicPinus canariensis (Pinaceae), a generallysparseplant formationthat showsascarceunderstorycover. Finally, above2,000ma.s.l., thevegetationischaracterizedbysparseleguminousshrubs, suchasSpartocytisussupranu-bius and Adenocarpus viscosus(Fabaceae).This last habitatharborsa great componentofendemicplants.30 km 15 0292818 17 16 15 14LA PALMAELHIERROLA GOMERATENERIFEGRAN CANARIAFUERTEVENTURALANZAROTE AlegranzaLobosMontaa Clara La GraciosaATLANTIC OCEANNFig.1 Map of the Canary Islands showing thelocalities where the diet of feral cats (Felis silvestris catus)hasbeenstudiedBiodiversConserv(2009)18:829846 8311 3ProceduresAlthough dietary studies could be a poor indicator of impacts on populations (Towns et al.2006), thestudyofferal catsdiet couldbeconsideredastherst steptointerpret theirimpact onpreypopulations (Paltridgeet al. 1997), as hadbeenmadeinseveral cases(Seabrook1990;Chapuisetal.1994;Donlanetal.2000;WinterandWallace2006).Forthisreason,theimpactsofferalcatintheCanaryIslandswasreviewedcompilingallthestudiesofferal catsdiet inthearchipelagoandtheinformationfoundinpublishedandgrayliterature, includingscientic articles, books, andinternal reports. Impact of thedifferent prey items captured by feral cats was assessed by evaluating their relativeabundance and by considering their respective status at international (IUCN2007),national(GobiernodeCanarias2003)andregionallevels(Mart netal. 2005b).ResultsanddiscussionCatpreyandthreatenedspeciesAtotalof68differentprey wereidentiedatspecieslevelin allstudiescarriedoutin theCanaryIslands(Table1). Fromthese, veweremammals, 16birds, 15reptilesand32invertebrates. All mammal prey are introduced species while two bird species are endemic(PhylloscopuscanariensisandFringillateydeapolatzeki), andotherveareendemicatsubspecies level (Anthus berthelotii berthelotii, Erithacus rubeculasuperbus, Fringillacoelebs palmae, Parus caeruleus teneriffae,and Sylvia melanocephala leucogastra). Onlyoneofthem,Alectorisrufa,isconsideredsuchasanintroducedspecies. Withregardstoreptiles, allofthemareendemicfromtheCanarianArchipelago,thesebeinglizards(theendemicgenusGallotiaspp.),geckos(Tarentolaspp.)andskinks(Chalcidesspp.).Fromthe total of 32invertebrate species identiedinthe analysis of feral cat scats inthearchipelago,15(47%)areendemic.From a conservation point of view, it is also important to consider some results of severalstudies carried out in the Canaries do not focus on feral cat diet but do on other biologicalaspects of native birds, such as Bulweria bulwerii, Pufnus assimilis, Hydrobates pelagicus,Saxicola dacotiae and Calandrella rufescens rufescens, which demonstrate importanteffectsofferalcatsintheirpopulations(ArduraandCalabuig1993;Lorenzoetal.2002;IlleraandD az2006).Thesestudiesareimportantwhenarealknowledgeoftheeffectofferal cats upon native species want to be understood, because bird remains in their faeces didnot permit the identication of birds at species level in most of the diet studies carried out intheCanaries(seeNogalesandMedina1996).Itismostsignicantinthecaseofseabirdsbecauseoceanicislandshavethemostimportantbreedingcoloniesofseabirds,wherethesmaller species areespeciallyvulnerabletocat predation(FitzgeraldandTurner 2000;Courchampetal.2003).IntheCanaryIslandsthepopulationsofsmallProcellariiformesspecies arerestrictedtouninhabitedislets andinaccessible cliffs due totheferal catspressure (Mart n et al. 1989) and currently probably present a lower population sizes than inthepast(Rando2003).Fromall thespeciespredatedbyferal catsintheCanaryIslands, onlyfour of themareconsideredsuchasthreatenedbytheIUCNRedList of ThreatenedSpecies(IUCN2007). TheCanaryIslandStonechat (Saxicoladacotiae) consideredasendangered, andthethreegiantlizardsfromtheCanariesareincludedascriticallyendangeredbecauseofthe high threat posed by the feral cat predation: Gallotia simonyi fromEl Hierro832 BiodiversConserv(2009)18:8298461 3Table1Listofspeciespreyedbyferalcats(Felissilvestriscatus)intheCanaryIslandsPreysDistributionBGRIUCNSRDBCNEACEACReferencesMammalsAtelerixalgirusT,C,F,LINTLR/lcMedinaetal.(2008)AtlantoxerusgetulusFINTLCMedinaetal.(2008)Rattusspp.P,G,H,T,C,F,LINTLR/lcNogalesandMedina(1996)MusmusculusdomesticusP,G,H,T,C,F,LINTLR/lcNogalesandMedina(1996)OryctolaguscuniculusP,G,H,T,C,F,LINTLR/lcNogalesandMedina(1996)BirdsNon-passerinesBulweriabulweriiP,G,H,T,C,F,LNATLCENLCVUArduraandCalabuig(1993)CalonectrisdiomedeaborealisP,G,H,T,C,F,LNATLCVULCLCNogalesetal.(1988)PufnusassimilisP,G,H,T,C,F,LNATLCENVUVUArduraandCalabuig(1993)HydrobatespelagicusG,H,T,C,F,LNATLCVULCLCArduraandCalabuig(1993)AlectorisrufaCINTLCSantanaetal.(1986)ColumbaliviaP,G,H,T,C,F,LNATLCNogalesetal.1988;MedinaandNogales(1993)PasserinesCalandrellarufescensrufescensTNATLCCRLCLCLorenzoetal.(2002)AnthusberthelotiiberthelotiiP,G,H,T,C,F,LENDLCLCLCNogalesetal.(1988);Medinaetal.(2006)ErithacusrubeculasuperbusT,CENDLCLCLCRodr guezetal.(1996)SaxicoladacotiaeFENDENENVUVUIlleraandD az(2006)TurdusmerulacabreraeP,G,H,T,CNATLCMedinaetal.(2006)SylviaconspicillataorbitalisP,G,H,T,C,F,LNATLCLCLCMedinaetal.(2006)SylviamelanocephalaleucogastraP,G,H,T,C,F,LENDLCLCLCMedinaetal.(2006)PhylloscopuscanariensisP,G,H,T,CENDLCLCLCNogalesetal.(1988);Medinaetal.(2006)BiodiversConserv(2009)18:829846 8331 3Table1continuedPreysDistributionBGRIUCNSRDBCNEACEACReferencesParuscaeruleusteneriffaeG,T,CENDLCLCLCRodr guezandMoreno(1993)PyrrhocoraxpyrrhocoraxbarbarusPNATLCENLCVUMedinaetal.(2006)FringillacoelebspalmaePENDLCENLCLCMedinaetal.(2006)FringillateydeapolatzekiCENDLCCRENENRodr guezandMoreno(1993)SerinuscanariusP,G,H,T,C,F,LNATLCSantanaetal.(1986);Nogalesetal.(1988)ReptilesGekkonidaeTarentolaangustimentalisF,LENDLCLCNogalesetal.(1990);Medinaetal.(2008)TarentolaboettgeriH,CENDLCLCSantanaetal.(1986);Nogalesetal.(1988)TarentoladelalandiiP,TENDLCLCNogalesetal.(1990);Medinaetal.(2006)LacertidaeGallotiaatlanticamahorataeFENDLCLCMedinaetal.(2008)GallotiacaesariscaesarisHENDLCLCNogalesetal.(1988)GallotiacaesarisgomeraeGENDLCLCNogalesandMedina(1996)GallotiagallotigallotiTENDLCLCMedinaandNogales(1993)GallotiagallotipalmaePENDLCLCMedinaetal.(2006)GallotiagomeranaGENDCRCRNogalesetal.(2001)GallotiaintermediaTENDCRCRENENHerna ndezetal.(2000)GallotiasimonyimachadoiHENDCRCRENENRodr guez-Dom nguezetal.(1998);Garc a-Ma rquezetal.(1997)GallotiastehliniCENDLCLCLCLCSantanaetal.(1986)ScincidaeChalcidessexlineatusCENDLCLCLCLCSantanaetal.(1986)834 BiodiversConserv(2009)18:8298461 3Table1continuedPreysDistributionBGRIUCNSRDBCNEACEACReferencesChalcidesviridanuscoeruleopunctatusG,HENDLCLCNogalesetal.(1988)ChalcidesviridanusviridanusTENDLCLCMedinaandNogales(1993)InvertebratesMolluscaGastropodaThebageminataP,G,H,T,C,F,LENDMedinaetal.(2008)RumiadecollataP,G,H,T,C,F,LNATMedinaetal.(2008)MalacostracaIsopodaArmadillidiumvulgareP,G,H,T,C,FINTMedinaandGarc a(2007)DiplopodaJulidaBlaniulusguttulatusP,G,TNATMedinaandGarc a(2007)ChilopodaScolopendravalidaP,G,T,CNATSantanaetal.(1986);MedinaandGarc a(2007)HexapodaOdonataAnaximperatorP,G,H,T,C,F,LNATLCMedinaetal.(2008)OrthopteraAcrididaeDericoryslobataC,F,LNATMedinaetal.(2008)GryllidaeGryllomorphalongicaudaP,H,TNATMedinaandGarc a(2007)GryllusbimaculatusP,G,H,T,C,F,LINTSantanaetal.(1986)BiodiversConserv(2009)18:829846 8351 3Table1continuedPreysDistributionBGRIUCNSRDBCNEACEACReferencesTettigonidaeDecticusalbifromsP,H,G,T,CNATMedinaandGarc a(2007)DermapteraAnisolabididaeAnisolabismaximaP,T,C,FNATNogalesetal.(1988)ColeopteraCarabidaeLaemostenuscomplanatusP,H,T,C,F,LINTMedinaandGarc a(2007)CurculionidaeBrachyderesrugatusP,H,T,C,LENDSantanaetal.(1986)ConiocleonusexocoriatusP,H,T,C,F,LNATMedinaetal.(2008)ConorhynchusconicirostrisP,H,T,C,F,LNATMedinaetal.(2008)HerpisticuscalvusC,F,LENDMedinaetal.(2008)ScarabeidaeOryctesprolixusP,H,G,T,CENDNogalesetal.(1988);MedinaandGarc a(2007)PachydemafuscipennisP,G,T,C,FENDMedinaandGarc a(2007)PhyllognatusexcavatusP,T,C,F,LNATMedinaetal.(2008)TropinotasqualidacanariensisP,G,H,T,C,F,LENDMedinaandGarc a(2007)StaphylinidaeOcypusolensP,G,H,T,C,F,LNATMedinaandGarc a(2007)TenebrionidaeHegeterglaberPENDMedinaandGarc a(2007)HegetertristisP,H,G,T,F,LNATMedinaandGarc a(2007)PaivaeahispidaC,F,LENDMedinaetal.(2008)PimelialaevigatacostipennisHENDNogalesetal.(1988)836 BiodiversConserv(2009)18:8298461 3Table1continuedPreysDistributionBGRIUCNSRDBCNEACEACReferencesPimelialaevigatalaevigataPENDMedinaandGarc a(2007)CerambycidaeArhopaluspinetorumP,H,G,TNATMedinaandGarc a(2007)DeropliaalbidaP,G,T,C,F,LENDMedinaandGarc a(2007)LepidopteraGracillariidaePhyllocnistiscanariensisP,G,TENDMedinaandGarc a(2007)SphingidaeHylestithymalitithymaliP,G,H,T,C,F,LENDMedinaandGarc a(2007)HymenopteraAnthophoridaeAnthophoraalluaudiP,G,H,T,C,F,LENDMedinaandGarc a(2007)MelectacurvispinaP,G,T,CENDMedinaandGarc a(2007)BRG,biogeographicrange;IUCN,IUCNRedListofThreatenedSpeciescategories;SRDB,SpanishRedDataBooks;CNEA,Spanishendangeredspecieslist;CEAC,Canarianendangeredspecieslist;INT,introducedspecies;END,endemicspecies;NAT,nativespecies;P,LaPalma;G,LaGomera;H,ElHierro;T,Tenerife;C,GranCanaria;F,Fuerteventura;L,Lanzarote;LR,lowerrisk;LC,leastconcern;VU,vulnerable;CR,criticallyendangered;EN,endangeredBiodiversConserv(2009)18:829846 8371 3(Garc a-Marquez et al. 1997; Rodr guez-Dom nguez et al. 1998), Gallotia intermediafromTenerife(Hernandezetal.2000)andGallotiagomeranafromLaGomera(Nogaleset al. 2001). Therest of species includedintheIUCNRedList arebeingconsideredspeciesof least concernor theyhadnot beendeemedasthreatened(seeTable1).At national level, 12speciesaffectedbyferal catsappearedintheSpanishRedDataBooks (Pleguezuelos et al. 2004; Madronoet al. 2005): thethree giant lizards, previ-ously commented, and nine bird species (Calonectris diomedea borealis andH. pelagicusasvulnerable, C. r. rufescensandF. t. polatzeki ascriticallyendangered,andB. bulwerii, P. assimilis, S. dacotiae, F. c. palmae andPyrrhocoraxpyrrhocoraxbarbarus as endangered). However, these species are not equally considered in theSpanishandtheCanarianEndangeredSpecies Acts (seeTable1). For example, whiletwoof thethreethreatenedgiant lizards(G. simonyi andG. intermedia) areconsideredendangeredinbothEndangeredSpeciesActs(GobiernodeCanarias2003; Mart net al.2005b), theother one(G. gomerana) isnot currentlyincludedeventhoughthisspeciesprobably is one of the most threatened lizard of the world (Valido et al. 2000).Nevertheless, the CanaryGovernment has recentlyproposedits incorporationintotheSpanishEndangeredSpeciesAct(Mart netal. 2005b). Otherspeciespreyedonbyferalcats in the Canaries, such as B. bulwerii, P. assimilis, C. diomedea, H. pelagicus,C. rufescens, P. pyrrhocorax, andF. c. palmae, arealsoincludedintheSpanishRedDataBookof Birds (Madronoet al. 2005) as endangeredor criticallyendangeredbuttheyare consideredas least concernspecies inbothofcial endangeredspecies lists,except P. assimilis which has been considered as vulnerable in the Canary Islands(Table1). Consequently, we believe that it is important to secure the legal status ofthese species inorder topromote aneffectivelyprotectionof their endangeredpopu-lations (Taylor 2000).Althoughferal catshavebeengloballyassociatedwithdeclinesor extinctionsof thelargest number of native andendemic species onislands (Dickman1996; Courchampet al. 2003), it isverydifcult toconcludethat thereisadirectlyrelationshipbetweenintroducedspeciesanddecliningindigenousspeciesbecauseof other complicatingfac-tors(Townset al. 2006). Someevidenceofferal cat impact onnativespeciesincluded,among others, predation, restoring endangered species after eradication, comparingreproductivesuccessbeforeandaftercontrol,comparisonbetweencat-inhabitedandcat-free islands, and local extinctions (Arnaud et al. 1993; Hodges and Nagata 2001;Mitchell et al. 2002; Edwardset al. 2004; Phillipset al. 2005; Rodr guezet al. 2006).Dietarystudies couldbeapoor indicator of impacts onpopulations, particularlywhenthepreyisrareoraffectedintermittently(Townset al. 2006). Moreover, thenumberofpreyeatenbyapredatorisnotnecessarilythesameasthenumberkilledbyit, whichisimportantinassessingtheecologicalroleofferalcats(Patronek1998). Nevertheless, onislandswhereahighnumber of endemicandendangeredspeciesarefound, studiesondiet or feedingecologyof feral cats hadbeenperformedconsideringthepredationaspotentiallyimportant evidenceoftheirincidenceonnativespecies(Nogaleset al. 1990;Seabrook1990;Arnaudetal. 1993;Chapuisetal. 1994;Donlanetal. 2000;WinterandWallace2006). Moreover, thestudyof feral cats diet couldbeconsideredas therststeptointerpret theimpact of thispredator onpreypopulations(Paltridgeet al. 1997).Accordingwiththis, weconsider that reviewingtheresultsobtainedinstudiesof feralcatsdiet constituteaninterestingtool topredict detrimental effectsintheconservationof the endangered species on island environments, even when the presence of onepredatoronlyhasbeenconsideredsufcienttoinducetheextinctionoftheendemicprey(Courchampet al. 1999).838 BiodiversConserv(2009)18:8298461 3FeralcatcontrolintheCanaryIslandsFeral cats are directly responsible for a large percentageof global extinctions, particularlyon islands worldwide, and for this reason they have been removed from at least 48 islandsaroundtheworld(Nogaleset al. 2004). Takingintoaccount that feral catsconstituteamajorthreattoendemicreptilespeciesonislands(Iverson1978;Arnaudetal.1993)andconsideringthe negative impact of feral cats inthe particular case of the endangeredCanariangiantlizards(Garc a-Marquezetal.1997;Hernandezetal.2000;Nogalesetal.2001), several management controls of feral cat populations hadbeenimplementedinthoseareaswheregiant lizardsarestill extant (Table2). WhileinElHierroIslandferalcats wereremovedfromtheareas where a reintroductionprogramof G. simonyi wasperformed(SilvaandMateo2003), themanagement control offeral catsinLaGomeraIslandprovokedabenecialeffectonthewildpopulationofG.gomerana(Mateo2001).Lastly,inthecaseof G. intermediathecontrolwasmainlycarriedoutinordertopreventtheproliferationofdomesticcatsinthegiant lizarddistributionareas(Rando2005). Inadditiontocat controlapermanent fencewasplacedinordertoprevent theentranceofcatsintothepopulationofthisendangeredspecies(Rando2005).InthecaseoftheCanarianendangeredbirdspecies,severalcontrolprogrammeshavebeenperformedinspecic areas inorder tominimize the effect of feral cat ontheirpopulations(Table2).Forexample,apopulationcontrolwascarriedoutinGranCanariaand Tenerife to minimize their effects on an endangered population of F. t. polatzeki and C.r. rufescens, respectively(Rafael Riera, pers. com.; Lorenzoet al. 2002). Onotherhandtwo successfully feral cat eradications have been carried out in two small islets (AlegranzaandLobos)wheresomeimportant seabirdcolonieshavebeenseriouslyaffectedbythispredator(ArduraandCalabuig1993;Rodr guez-LuengoandCalabuig1993;Mart netal.2002a, b). The most noteworthy programs of feral cats eradication had been carried out onislandswhereahighnumberofendemicbirdspeciesarefound(Nogalesetal. 2004).Althoughtheseshort termconservationactionsareimportant toreducetheimpact offeral cats on these endangered species, we consider that these efforts should be maintainedat longtermtimescale, andregularlyduringeachyear, inordertominimizetheircon-servationconcerns. Asoccur worldwide, intheCanaryIslandspeople(about 2millionpersons) lovecats becausetheyareconsideredas abenecial animal helpinghumanscontrollingplaguessuchasratsandmice. Moreover, insomeislandsofthearchipelago,endemicspeciessuchaslizards(Gallotiaspp.)orpigeons(Columbabollii andC. juno-niae)areconsideredasplaguesduetotheirdamagetofarmandcrops.Probably,forthisreasoncatshadnot beenpreviouslyconsideredasaconservationconcerninthisarchi-pelago. Moreover, acommonpracticeinseveral islandsistoreleasekittensinthewildwhen the owners do not want more cats at home. These facts could be considered as a veryimportantlimitationtodevelopcontrolprogrammesinadditiontothegreatvariabilityofhabitatsandtheabrupttopographyofthemainislands.GeneralconsiderationonhyperpredationandmesopredationreleaseIngeneral, therelationshipbetweenpopulationsizeof feral cats anddamageontheirendangered populations could be overestimated, ignoring temporal and spatial variation inenvironmental conditions andvariationinthedensityof other species (Edwards et al.2004). In the case of the Canary Islands the presence of other introduced mammals, such asrats or mice, together withhumandisturbance(habitat destructionandfragmentation),could compound feral cats effects on endemic and endangered species. So, the presence ofBiodiversConserv(2009)18:829846 8391 3Table2Resumeofthecontrolprogrammesofferalcats(Felissilvestriscatus)carriedoutintheCanaryIslandsIslandIs.surface(km2)Controlarea(km2)HabitatControlperiodMethodsCatsremovedOtherintroducedpredatorsMainspeciesaffectedReferencesAlegranza10.210.2XSMay1998Livetraps1MusdomesticusBulweriabulweriiCalonectrisdiomedeaPufnusassimilisPelagodromamarinaHydrobatespelagicusOceanodromacastroMart netal.(2002a)Lobos4.44.4XSSep19921923May19972129Aug2002Livetraps,poison603MusdomesticusBulweriabulweriiCalonectrisdiomedeaPufnusassimilisPelagodromamarinaHydrobatespelagicusOceanodromacastroArduraandCalabuig(1993)M.Gonza lez,pers.com.Mart netal.(2002b)TenerifeLosRodeos2034.4&1Grassland20012002Livetraps54MusdomesticusRattusrattusRattusnorvegicusAtelerixalgirusCallandrellarufescensLorenzoetal.(2002)TenerifeTenoa2034.40.25XSNov04Feb05Livetraps35MusdomesticusRattusnorvegicusRattusrattusGallotiaintermediaRando(2005)GranCanariaInagua1560.139.2PF1996199719992002Livetraps1751MusdomesticusRattusrattusRattusnorvegicusAtelerixalgirusFringillateydeaRodr guezetal.(1997)Rodr guezetal.(2003)GranCanariaTamadaba1560.1&1PFNovDec2002Livetraps3MusdomesticusRattusrattusRattusnorvegicusAtelerixalgirusFringillateydeaRodr guezetal.(2003)840 BiodiversConserv(2009)18:8298461 3Table2continuedIslandIs.surface(km2)Controlarea(km2)HabitatControlperiodMethodsCatsremovedOtherintroducedpredatorsMainspeciesaffectedReferencesCanariaMaspalomas1560.1&5TF,DunesJulOct2008Livetraps30MusdomesticusRattusrattusRattusnorvegicusAtelerixalgirusCharadriusalexandrinusR.Riera,pers.com.GranCanaria1560.1300PFJulOct2008Livetraps85MusdomesticusRattusrattusRattusnorvegicusAtelerixalgirusFringillateydeaR.Riera,pers.com.ElHierro268.7&40XS,TF20012005Livetraps275MusdomesticusRattusrattusGallotiasimonyiRodr guez-Dom nguezetal.(2007)LaGomeraa369.8&1XS1730Sep199920002001Livetraps758MusdomesticusRattusrattusGallotiagomeranaIllera(1999)Mateo(2001)XS,XerophyticShrub;TF,TermophyleForest;PF,PineForestaThesemanagementcontrolsweremainlycarriedoutondomesticorstraycatpopulationsduetotheirclosedistributiontothegiantlizardpopulationsBiodiversConserv(2009)18:829846 8411 3acontrolledpopulationof cats might be, at least temporarily, morebenecial totheirendemicpreythanitseradication, asoccur inthecasesonmanyislandswhererodentshave also been introduced (Courchamp et al. 1999; Rayner et al. 2007). On islands, whereintroduced mammals constituted the main prey of feral cats (Fitzgerald 1988; Nogales andMedina1996), thispredatorcouldreducetherodentpopulationsatlowlevelsandcouldoutweigh the damage done to the endemic prey species. For example in the laurel forests ofthe Canary Islands there are not evidence of predation by feral cats upon endemic pigeons(Columba bolli and C. junoniae) (Nogales and Medina 1996) but by rats (R. rattus) whichprey oneggs,chicks, juvenilesand adultsofthesetwoendemic pigeons(Hernandez etal.1999; Mart net al. 2000). Ratsconstitutedthemainpreyitemofferal catsinthiseco-system(Nogalesetal.2006),showingtheindirectrolethatcatsmightplayinpreservingnativefaunathroughreductionof rat predationpressureonendemicpigeons. For thisreasonthemanagementofferalcatpopulationsshouldbedevelopedtakingintoaccountthemesopredatorreleaseeffect,wheretheeliminationofferalcatpopulationsfromsuchecosystems could lead to a more severe negative impact on the endemic species, as a resultof expansionof rodent populationoncetheir predators areremoved(Courchampet al.1999). Thisimportant aspect hasbeenconsideredwhentheferal catscontrol campaignshavebeencarriedout inthecases of G. intermedia(Rando2005) andC. r. rufescens(Lorenzo et al. 2002). In both cases other mammals predators such as rats, mice orHedgehogs(Atelerixalgirus)whichcouldbeaffectedtheirendangeredpopulationswerealsoremoved. So, takingintoaccount thepredictionsmadeinthedifferent modelspre-sented (Courchamp et al. 1999, 2000) and the results obtained in the different studies (i.e.,Rayneretal. 2007), weconsiderthatcontrolprogrammesoftheintroducedpredatorsonislands must be performed simultaneously to the mesopredators and introduced preycontrolinordertoavoidundesirablesandirreversibleeffectsonendemicspecies.ConclusionsIntheCanaryIslandsit isnecessarytoenforcemanagement controlsofferal cat popu-lationsonthoseareaswherehighlyendangeredspeciespopulationshavebeenseriouslyaffected by feral cats, such as the distribution areas of giant lizards in Tenerife, La GomeraandEl Hierroislands. Nevertheless, it isimportant totakeintoaccount thepresenceofother introduced predators and prey such as rats, mice or rabbits in order to avoid problemsderivedfromthehyperpredationprocess andmesopredator releaseeffect as havebeenpreviously mentioned. Finally, in the Canary Islands, as well as in other island ecosystems,prioritymanagementrequirementsforendangeredspeciesmustincludethepreventionofthe introducedmammals spread (Nogaleset al. 2004), removing introducedpredator fromendangeredspecies distributionareas, researching their ecological role for eradicationprogrammes (Courchampet al. 2003; Taylor 2000), andthedevelopment of educationcampaignsforpeopleabouttheproblematicofpetsreleaseintonaturalecosystems.Acknowledgments We thankall the co-authors whoparticipatedwithus inthe partial contributionscarriedoutonthestudyofferalcatdietintheCanaryIslands,withouttheirbasicinformationthisreviewhadnot beenpossible. Manuel Gonzalez-Acebes, AurelioMart n, JuanCarlosIllera, JuanCarlosRando,RafaelRieraandMiguelAngelRodr guez-Dom nguezprovidedusefulinformationaboutferalcatcontrolprogrammes performed in the Canary Islands. 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