Land Use Changes and Raptor Conservation in Steppe Habitats of Eastern Kazakhstan

Landusechangesandraptorconservationinsteppehabitatsof EasternKazakhstan

Jose´ AntonioSa´ nchez-Zapataa,*,MartinaCarreteb,AndreiGravilovc,SergeiSklyarenkoc, OlgaCeballosd,Jose´ AntonioDona´ zard,FernandoHiraldod aA´ readeEcologıá,DepartamentodeBiologıáAplicada,UniversidadMiguelHernańdez,CtraBenielkm3.2,Orihuela,Alicante,Spain bDepartamentodeEcologıáeHidrologıá,UniversidaddeMurcia,30100Espinardo,Murcia,Spain cInstituteofZoology,Akademgorodok,480060Almaty,Kazakhstan dEstaciońBiolo´gicadeDonãna,CSIC,Avd.MarıáLuisas/n,PabellońdelPeru´,41013Sevilla,Spain

Abstract SteppehabitatsincentralAsiahavesufferedimportantlandusechangesduringthiscenturywhicharesimilartothosethathave beenpointedoutasthecausesofthedeclineofsteppebirdsinwesternEurope.DuringJune1999weconductedroadsurveysof raptorsinEasternKazakhstantodetectspecificandcommunityresponsestolandusechanges.Wedetected11speciesofraptors. Kestrels(Falconaumanni and tinnunculus)werethemostcommonspeciesingrasslandsandagriculturallandscapes,harriers(Circus pygargus and macrourus)weredominantinsalinesteppesandsteppeeagles(Aquilanipalensis)weredominantindrysteppes.There werefewerspeciesinagriculturalhabitatsthaningrasslandandsteppehabitats.Ground-nestingraptorswerenegativelyaffectedby landusechangesandfourspecieswereneverdetectedinagriculturalzones.Raptorabundancepatternsdifferedbetweennatural steppehabitatsandhuman-transformedhabitats,whereapatchydistributionwasdetected.Thefutureofraptorcommunitiesin Kazakhstanseemsuncertainalthoughtheprogressiveabandonmentofintensiveagriculturemaybenefitspeciessensitivetohuman presence.Thelong-termconservationofvertebratecommunitiesmaydependuponthemaintenanceofecologicallyandsocially sustainablegrazingsystems.

Keywords: Steppes;Raptors;Habitattransformation;Grazingsystems;Kazakhstan

1. Introduction inextensivefarmingsystemsofwesternEuropethat resemblenaturalgrassyhabitats(Goriup,1988;Sua´ rez The response of birds to habitat changes has etal.,1997;Blancoetal.,1998;Tellaetal.,1998;Tella been a central topic in conservation biology and andForero,2000)whereitisconsideredthat76%of landscape ecology has widely centred on forest avianspecieslinkedtonaturalorsemi-naturalEurope fragmentationintemperatehabitatsofEuropeand steppehabitatsaresufferingpopulationdeclines(Sua´ rez North America (Wilcove, 1985; Saunders et al., etal.,1997). 1991; McGarigal and McComb, 1995). More LargeOldWorldnaturalsteppeareasremain,how- recently,ecologisthavepaidattentiontochangesin ever,incentralAsia(mainlyKazakhstan,Mongoliaand openhabitatssuchasshrubsteppes,grasslandsand Russia)wherethereislittleinformationavailableonthe extensivefarmingsystems(Herkert,1994;Bignaland effects of land-use changes on bird communities McCracken,1996;PainandPienkowski,1997;Sua´ rezet (Bragin,1999)althoughthishabitatisastrongholdfor al.,1997;SutterandBrigham,1998;Ormerodand globallyrareraptorssuchaslesserkestrels(Falconau- Watkinson,2000;Wolffetal.,2001).Researchonavian manni)orimperialeagles(Aquilaheliaca)(delHoyoet responsestosteppetransformationhasbeencarriedout al.,1994;Parretal.,2000).There,largetransformations insteppeandsemi-desertweremadeduringthemiddle *Correspondingauthor. of20thcenturywhentheUSSR.administrationpro- E-mailaddress: [email protected](J.A.Sa´ nchez-Zapata). motedtheso-called‘‘upturnofvirginlands’’.Thus, from1954to1965,andevenlater,mostpartsof 2. Methods northernhumidsteppesweretransformedintoagri- culturallands(E.Bragin,personalcommunication). 2.1.Studysite Inaridsouthernzones,irrigationprogramstrans- formedhugeextensionsofsemi-desertalongtheSyr Thestudywasconductedbetween12and28June DaryaRiverintointensiveagricultureathighecolo- 1999inEasternKazakhstan(CentralAsia,Fig.1).The gicalcost,includingareductionintheextentofthe climateiscontinental,withverycoldwinters(when AralSeaandthecontaminationandsalinizationof temperaturesremainunder0 Cformonths),andwarm soilsandwaterbeds.InEasternandNorthernKazakh- summers.Theconditionscanalsobeconsideredasarid stan,largeextensionsofsteppesanddrysteppeswere orsemiaridwithannualrainfall <300mm.Theextreme transformedintoextensiveagriculturefollowingpro- temperaturesandrainfalllimitthegrowthoftreespe- gramstochangethenomadicwayoflifeofKazakh cies(Walter,1981)andlandscapeisdominatedby shepherdstosedentaryandagriculturalsettlements. steppeandsemidesertplainsandhillswithgrassesand Thesechangesmayhavealsoinfluencedtheecologi- smallbushes,whereastreesareconfinedtofoothillsof calprocessesofsteppes.Inspiteofsuchtransforma- largemountains(TienShaninthesouth,Altaiand tions,thesizeofthecountryandthelowhuman Alatulintheeast)oraroundhumansettlementsand densityhasallowedthepersistenceoflargesteppe rivervalleys. zoneswithlittleornohumaninfluence.Furthermore, Weconsideredfourdifferenthabitattypesinthe humanpopulationhasbeendecreasingoverthelast studyarea: yearsandsomeagriculturalareashaveundergonea recoveryprocessintonaturalsteppes.By1992the 1.Agriculturallandscapes(AGRI).Thelandscape liberalization of theeconomywasunderway,agri- isdominatedbycerealsandirrigatedcrops, culturewasrapidlydecreasingand >50%ofploughed managedpasturelands,citiesandvillages.There landinsteppeshadbeenabandoned.Furthermore,asin arealsonumerousherdsincludingsheep,horses otherInnerAsiacountries,livestockhadbeenpriva- andcows.Smallwoodsandscatteredtreefor- tizedandherdingcollectivesdismantled(Ferna´ ndez- mations are frequent around large villages Gime´ nez,1999a). (>1000inhabitants)andalongsomeroads. Inthispaperweexaminehowland-usetransforma- 2.Seminatural grasslands (GRASS). Here the tionsofdryKazakhsteppeshaveaffectedthebreeding landscapeisdominatedbyextensivelivestock communityofbirdsofprey.Raptorsareusuallycon- andsomeextensiveagriculture,butvillagesare sideredastoppredatorswhosepresenceandrichness smallwith <500inhabitants.Therearemany indicatehealthyecosystems(seeNewton,1979;butalso abandonedfieldsanddegradedsteppeareas. Rodrı´ guez-Estrellaetal.,1998).Inaddition,raptorscan 3.Salinesteppe(SAL).Thishabitatincludeslarge bedetectedwithrelativeeasecomparedtomoreelusive salinesteppeandmarshesinthevicinityofBal- organisms.Thus,indicesofraptorabundancehavebeen jashandAlakolLakes.Vegetationisdominated widelyusedtoassesstheimpactofhuman-induced byvarioussalineshrubs(Sarcocornia spp. Scir- transformationswithinavarietyofhabitats,ecosystems pus spp. Artrocnemum spp)andreedbeds Phrag- andbiomes(Burnhametal.,1990;Ellisetal.,1990; mites spp.withsparsetrees(Populustremula, Rodrı´ guez-Estrellaetal.,1998).Theseindicesallowus Eleagnushortensis). toexamineifraptorabundancefollowsspatialpatterns 4.Drysteppes(STEPPE).Thisincludeslargeareas thatcanberelatedtoapatchyorhomogeneousdis- ofnaturaldrysteppeswithlittleornohuman tributionofresources.Theidentificationofthesepat- presence.Treesarelackingandthevegetationis ternscanbemadethroughspatialautocorrelation dominatedbygrassesandforbs(Artemisia spp., approximations(LegendreandLegendre,1998),asta- Limonium spp., Salsola spp., Ephedra spp., tisticalapproachwidelyemployedinplantecologybut Haloxylon spp.).Therearealsosparserocky stillpoorlyknownamongvertebrateecologists(see outcrops. LegendreandFortin,1989). Summarizing,ourspecificaimswere:(i)todescribe 2.2.Roadcounts thebreedingraptorcommunitiesoftheKazakhsteppe ecosystems;(ii)totestifraptordistributionpatternsare ThesurveyswereconductedinJune1999:habitat1 repeatedalonghabitatswithdifferentdegreesofaltera- (504kmbetweenAlmatyandBeskol(AlakolLake); tion;and(iii)toanalysetheeffectsofsteppetransfor- habitat2(142kmbetweenAqtoghayandAyako¨ zand mationontheabundanceandcompositionofthese closedtoZaysanLake);habitat3(89kminfloodplains communitiesanddiscussfutureconservationscenarios aroundBaljashandAlakollakes)andhabitat4(155 forraptorsinsteppehabitats. aroundBaljashLake). landuses.Asweregisteredthenumberofraptors detectedateachkmoftransect,weusedaPoissondis- tributionaserrorandlinkfunctions(Vincentand Haworth,1983;Nichols,1989;Dona´ zaretal.,1993a,b; Sa´ nchez-ZapataandCalvo,1999).ForGLMsweused theprogramSTATISTIX(AnalyticalSoftware,1992). Finally,becauseoflowdataavailableonhumanset- tlements(e.g.herdsorhumanpopulation)weanalysed spatialpatternsofraptorassemblagebymeansofspa- tialautocorrelogramswithMoran’sIndexasauto- correlationstatistic(Legendre,1993;Garcı´ a-Charton andPe´ rez-Ruzafa,1999),usingtheprogramauto- correlationincludedinthe‘‘R’’v.3package(Legendre andVaudor,1991).Foreachhabitatcategoryweper- formedsuccessivetestscomparingsimilaritybetween pairsofsitesseparatedbyincreasingdistances,from1 km(lowestdistanceclass)tohalfthetotallengthofthe roadcensus,withthatobtainedfrompairsofsitesran- domlyselected.Moran’sindexvariesbetween+1and Fig.1.Studyarea. À1andiscomparabletoaPearsoncorrelationcoeffi- cient.Values >0representsimilaritiesbetweenpoints Roadsidecountsweredonefollowingmethodswidely greaterthanexpectedfromrandom;those <0repre- employedinsimilarstudies(seee.g.FullerandMosher, sentinganoppositetrend.Significanceofthetestswere 1987;Dona´ zaretal.,1993a,b;Travainietal.,1995).The determinedattwolevels.First,weperformedaglobal averagedrivingspeedwas40–50km/h.Werecorded testbycheckingwhethertheautocorrelogramcontained everyraptorseenandthekmintervalwheretheywere atleastonevaluethatwassignificantatthe =0.05/n observed.Inafewcasesitwasnecessarytostopthe Bonferroni’scorrectedlevelformultipletests(Legendre vehicletoidentifythebirds(Ellisetal.,1990).Inthis andLegendre,1998).Second,thestatisticalsignificance case,wedidnotrecordnewbirdsobservedduringthe ofeachMoran’sindexwasexaminedbycomparingthe stop.Inthecaseofsteppeeagles(Aquilanipalensis),we actualdistributionswhiththoseobtainedbyrandom. distinguishedbetweenadultandimmaturebirdsonthe Patchy distributionswouldtakeplacewhenauto- basisofplumage(Forsman,1998).Finally,wealso correlogramsshowasignificantpositiveornegative recordedthenumberofvillages,cattleherdsandsusliks peakofMoran’sIndexatagivendistanceclass.Gra- (Citellus spp.,Rodentia)alongeachsurveyroutefora dientsarerepresentedbysignificantpositivevaluesof furtherdescriptionofthehabitat.Thesurveyswere Moran’sIndexatlowdistanceclassesfollowedby carriedoutinfineweather,withoutcloudsandwith negative values (Garcı´ a-Charton and Pe´ rez-Ruzafa, windvelocity <20km/h. 1999).Duetolimitationsinthenumberofdatathat couldbeenteredintheprogram(LegendreandVaudor, 2.3.Dataanalysis 1991),AGRItransectwasclassifiedintosamplingunits of3and5km(Fig.2aandb,respectively).Fortherest Eachraptorspecieswasassignedtooneofthefol- ofthetransectsthesamplingunitwas1km. lowingnestinghabitats:ground,tree,rockorrock/tree accordingtotheirbreedinglifehistories(delHoyoet al.,1994).Thefirstlevelofanalysiswasdonethrough 3. Results contingencytablesand w2-tests(Siegel,1956)pooling thedatatolookfordifferencesinthefrequencyof Wedetected11speciesofraptorsduringthesurveys groundbreedingraptors(nestingsubstrateaccessibleto (Table1).Thehighestspeciesrichness(10species)was predators)rockortree-nestingspecies.Differencesin reachedinnatural(dryandsaline)steppes;grassland raptorabundance,measuredasbirdsobservedperkm, habitatsheldeightspecieswhereasinagriculturalareas weretestedbymeansofnonparametricKruskal–Wallis itdroppedtoonlyfivespecies.Commonandlesserkes- tests. trels(Falcotinnunculus and F.naumanni,respectively) Second,weusedGeneralizedLinearModels(GLMs) werethemostabundantspeciesinagriculturalareas (Dobson, 1983; McCullagh and Nelder, 1989)to andseminaturalgrassland.PallidandMontagu’shar- describetheabundance(raptors/km)ofthemostcom- riers(Circusmacrourus and C.pygargus,respectively) monspeciesinthefourmainhabitats.Eachkminterval madeupthebulkofraptorsinsalinesteppe.Finally, ofthetransectwasassignedtooneofthefourmajor steppeeagle(Aquilanipalensis)wasthecommonestspe- Fig.2.Serialautocorrelationbetweenpairsofsamplingunitsseparatedbydistancesrangingfrom1(adjacentunits)tohalfthelengthofthe transects,forthetotalabundanceofraptorsineasternKazakhstan.(a)and(b)AGRI:highlyhumanizedagriculturelandscape(Bonferroni’scorrectionat =0.0003and =0.0005,respectively),(c)GRASS:extensiveagricultureandgrasslands(Bonferroni’scorrectionat =0.0006),(d) SALINE:salinesteppeandmarshlands(Bonferroni’scorrectionat =0.0004)and(e)STEPPE:drysteppes(Bonferroni’scorrectionat0.0003). Blackdots:Moran’sIndex P<0.05.**Significantautocorrelograms(P <0.05).

Table1 ciesindrysteppes.Aroundvillagesandmoremanaged Raptorcounts(birds/100km)infourdifferenthabitatsinEastern areaswedidnotfindfourspecies:steppeeagle,saker Kazakhstan(inbrackets,numberofbirdsobservedduringroad falcon(Falcocherrug),short-toedeagle(Circaetusgalli- counts) cus)orpallidharrier. Species AGRI GRASS SALINE STEPPE Globalraptorabundancewasdifferentamonghabi- H P= Milvusmigrans 2.2(11) – 3.4(3) – tats( =10.96, 0.0119).Theabundanceofraptors Buteorufinus 0.8(5) 2.1(3) 5.6(5) 5.0(8) washigherinseminaturalgrasslandsandsalinesteppes Aquilaheliaca – – – 0.6(1) thaninagriculturalanddrysteppehabitats.Asregards Aquilanipalensis – 3.5(5) 1.1(1) 11.3(18) nestinghabitat,wefoundamuchhigherfrequencyof Circaetusgallicus – – – 1.3(2) individualsofground-nestingspecies(steppeeagle,har- Circus spp. – 2.1(3) 3.4(3) – Circuspygargus 0.2(1) 2.1(3) 7.9(7) – riers)innaturalsteppehabitats(dryandsalinesteppes) Circusmacrourus – 1.4(2) 7.9(7) 1.3(2) (53.5%, n=71)thanintransformedagriculturaland Falcocherrug – – – 1.3(2) grasslandzones(5.5%, n=256).Thefrequenciesof Falcosubbuteo 1.2(6) 1.4(2) 1.1(1) – contactswithspeciesbreedinginrocksand/ortrees Falconaumanni 2.8(14) 22.4(32) – – (safeplacesagainstpredators),were46.5%insteppe Falcotinnunculus – 9.1(13) 3.4(3) 1.9(3) Falconau/tin 27.1(136) 13.3(19) 3.4(3) – habitatsand94.5%inagriculturalandgrasslandzones 2 Total 34.5(174) 57.3(82) 39.3(35) 23.1(36) andthedifferencewashighlysignificant(w =95.97, Towns 3.0(15) 4.9(7) – – df=1, P<0.0001).Finally,immaturesteppeeaglesmade Cattleherds 12.7(64) 10.6(15) 16.9(7) – up33–67%oftheobservedindividuals.Theyweremore Citellus spp. 0.8(4) 0.65(1) – 10.3(16) frequentintransformedseminaturalgrasslands(66.6%; AGRI,highlyhumanizedagriculturelandscape(504km);GRASS, n=9)thaninnaturalsteppes(dryandsaline)(34.2%; extensiveagricultureandgrasslands(142km);SALINE,saltysteppe n=38)althoughthesedifferencesdidnotreachsig- andmarshlands(89km);STEPPE,drysteppes(155km). nificance(Fisherexacttest, P=0.129)(Table2). 7

Table2 respectivelywithdryandsalinesteppehabitatsaswell Proportionofraptorsnestingontheground(Circusmacrourus,Circus asgrasslands.Modelsforpallidharrier,lesserkestrel, pygargus,Aquilanipalensis),onrocks(Falcocherrug,Falconaumanni), commonkestrelandlong-leggedbuzzardexplained ontrees(Falcosubbuteo,Milvusmigrans,Circaetusgallicus,Aquila heliaca)orboth(Buteorufinus,Falcotinnunculus)infourdifferent 19.4,16.0,10.8and7.6%oftheinitialdeviance, habitatsinEasternKazakhstan(inbrackets,numberofbirdsobserved respectively.Thevariablesenteringintothemodels duringroadcounts) revealedthatthepresenceofthesespecieswasincreas- inglyindependentofthedegreeofhabitattransforma- AGRI GRASS SALINE STEPPE tion;therewasaslighttrendtofindharriersinsaline Ground 0.6(1) 15.9(13) 51.4(18) 55.5(20) steppes,lesserkestrelsingrasslandandbuzzardsin Rock 8.0(14) 39.0(32) – 5.5(2) naturalsteppeareas. Tree 9.8(17) 2.4(2) 11.4(4) 8.3(3) Rock/tree 81.6(142) 42.7(35) 37.1(13) 30.5(11) Alltheautocorrelograms,exceptthatcorresponding tosteppes,showedsignificanceforMoran’sIndexesof AGRI,highlyhumanizedagriculturallandscape;GRASS,extensive autocorrelationafterapplyingBonferroni’scorrection, agricultureandgrasslands;SALINE,salinesteppeandmarshlands; whichrevealsirregularspatialdistributionofraptor STEPPE,drysteppes. assemblages(Fig.2c).Atlowdistanceclasses,Moran’s Indexes were not significant, except for auto- Table3 correlogramsb(AGRI—5kmsamplingunits)andc GLMsonraptorabundanceinrelationtohabitatcategoriesineastern (GRASS)(Fig.2).Autocorrelogrambshowedsig- Kazakhstan nificantpositiveautocorrelationsatdistanceclasses2,4 Coeff. S.E. P and6km,butnorepeatedspatialpatternwasdetected. Repeatablespatialpatternswereonlyfoundingrass- Aquilanipalensis Constant À6.38439 0.96496 *** landareaswheremaximumnumbersofraptorswere Steppe 4.19959 0.99333 *** regularlyfoundat8kmintervals. Grass 3.03099 1.06356 ** %deviancechange=26.8% 4. Discussion Buteorufinus Constant À4.52766 0.37319 *** Steppe 1.53193 0.51407 ** Kazakhsteppehabitatsseemtobeveryrichinrap- Saline 1.64846 0.58246 ** tors,whichwouldindicateagoodconservationstatus %deviancechange=7.6% fortheselands.Thenumberofdetectedspecieswashigh (11),andtheindexofabundance(23–57/100km)is Circuspygargus Constant À6.49668 0.96086 *** similarorevenhigherthanthosefoundinothersteppe Saline 3.95396 1.03253 *** areasoftheworldsuchastheArgentineanPatagonia Grass 2.63245 1.12098 * (10.0–32.6individuals/100km,Ellisetal.,1990;Dona´ - %deviancechange=25.1% zaretal.,1993a,b).Itcanbearguedthattheresultsof thesurveyscanbebiasedbyseveralfactors.Forexam- Circusmacrourus Constant À5.30823 0.49790 *** ple,roadsmayattractscavengerspecieslookingfor Saline 2.76551 0.62511 *** cadavers,andmanyspeciesperchonpolesandfencesso %deviancechange=19.4% theycanbemoreeasilydetectedinthevicinityofroads. However,weneversawabirdofpreyfeedingonroad- Falconaumanni killedcorpses,althoughsomespeciesliketheblackkite Constant À3.98632 0.26644 *** Grass 2.48921 0.31975 *** (Milvusmigrans)maybenefitlocally.Polesandfences %deviancechange=16.0% may,indeed,facilitatethedetectionofmostspecies recordedinthearea,asinotheraridecosystems Falcotinnunculus (Dona´ zaretal.,1993a,b;Leptich,1994;Rodrı´ guez- Constant À3.90801 0.44657 *** Estrellaetal.,1998),buttheywouldhaveaffectedall Agr 2.60343 0.45467 *** Grass 1.88961 0.50205 *** speciesequally(withtheexceptionofharriers,whichdo %deviancechange=10.8% notusethem)thustendingtoincreasethenumbers recordedinmoredevelopedregions. * P<0.05. Salineanddrysteppesweretheonlyhabitatsthat ** P<0.01. showednosignificantautocorrelograms,suggestinga *** P<0.001. uniformdistribution.Thismayberelatedtotheobvious homogeneousdistributionofresourcessuchasbreeding GLMmodelsforsteppeeagleandmontagu’sharrier placesandprey.Ontheotherhand,analysesdetecteda explained26.8and25.1%oftheinitialdeviancerespec- patchyspatialdistributionofraptorsintheothertwo tively(Table3).Thesetwospecieswereassociated alteredhabitatsandaspatialpatternthatrepeated alonggrasslandtransects.Ingeneral,patchydistribu- seemsuncertain.Progressiveabandonmentofintensive tionmayberelatedtoirregularhabitatfeaturespro- agricultureincentralAsiamaybenefitlargespeciestra- ducedbyecologicalheterogeneity(marshareasinsaline ditionallysensitivetohumanpresence,suchaslarge steppes)andclearlybyhumaninfluenceingrassland eaglesandfalcons.Ontheotherhand,smallerspecies andagriculturalareas.Theapparentlyregulardistribu- suchaskestrelsbenefitfromhumansettlementsand tionofraptorsingrasslandareasisprobablyduetothe couldbenegativelyaffected.Probably,thelong-term spatialdistributionofherdersettlements,whichare conservationofsteppevertebratecommunitiesrequires regularlydistributedasaconsequenceofthecollectivi- themaintenanceofecologicallyandsociallysustainable zationofnomadicpeopleinthemid20thcentury(Fer- grazingsystems(Dona´ zaretal.,1993a,b;Ferna´ ndez- na´ ndez-Gime´ nez,1999a).Theinhabitantsnowmanage Gime´ nez,1999a).Inparallel,itwouldbedesirableto pasturesandmaintainsmallcropswithinasubsistence createandmaintainlargereservezoneswithoutorwith economy(Ferna´ ndez-Gime´ nez,1999b).Thelesserkes- verylowhumanpresence,becausenumbersofcertain trelandMontagu’sharrierhavebeenabletoshift speciessuchassakerfalconmayalsobelimitedbyfal- breedingsitesandadapttolowintensityfarmingsys- conry.Tourismactivitieslinkedtonatureobservation, tems,nestingwherenecessaryonbuildingsandincereal andotherlowintensityusessuchasregulatedhunting, crops,respectively.Bothspeciesremainasbreedersin couldhelptomaintainpopulationsoflargevertebrates pseudo-steppehabitatsofWesternEuropewheremore insuchareas.Furtherresearchonhabitat-speciesrela- sensitiveraptorspeciessuchaspallidharrierorsteppe tionshipsinAsiansteppehabitatsmayalsoprovide eagleareabsent(TuckerandHeath,1994;A.Ladyguin newinsightsformanagementpracticesinpseudo- andV.Gory,personalcommunication) steppehabitatsinwesternEurope,whensocioeconomic Transformationsinsteppehabitatsdeterminestrong and environmental criteria of the Common Agri- variationsinthespeciescompositionofthedifferent culturalPolicy(CAP)havetobeupdatedinthenext raptorcommunities.Fivespecieswereneverdetectedin fewyears. agriculturalzones:imperialeagle,steppeeagle,saker falcon,pallidharrierandshort-toedeagle,andmostof themarerareoutsidetheirbreedingrangeinCentral Acknowledgements Asia(delHoyoetal.,1994).Rarefactionofground- nestingspecies,especiallyeagles,maybeinterpretedasa TheInstituteofZoologyofAlmatyprovidedvaluable resultofincreasingriskofpredation(humans,dogsand helptoorganisethefieldwork.J.A.Garcı´ a-Charton cats)derivedfromhumanactivities.Alternatively,agri- andM.Martı´ nezhelpedwithdataanalysis.Wethank culturalintensificationappearstoreducetheabundance Jose´ L.Tellaforcommentsonthemanuscript. ofmedium-sizedandlargerodentssuchassusliks (Citellus spp.Table1)andgerbils(Rhombomybs spp.), References whichseemtobethemainpreyofsteppeeagle,long- leggedbuzzardandsakerfalcon(personalobservation). 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