Biologia,Bratislava,/2:219|231,2004Structureofthebreedingbirdassemblag - PDF document

Biologia,Bratislava,/2:219|231,2004Structureofthebreedingbirdassemblageofaprimaevalbeech-rforestinthe©rámkováNationalNatureReserve,theMaláFatraMtsMartinKSlovakNationalParkService,TheAdministrationoftheMaláFatraNationalPark,Nám.sv.Floriána,SK{Varín,SlovakiaCurrentaddress:CenterforEcologicalStudies,Ústredie,SK{VeµkéRovné,Slovakia;e-mail:mkornan@stonline.skORÒAN,M.Structureofthebreedingbirdassemblageofaprimaevalbeech-rforestinthe©rámkováNationalNatureReserve,theMaláFatraMts.Biologia,Bratislava,219|231,2004;ISSN0006-3088.Thestructureofabreedingbirdassemblageofaprimevalbeech-rforestintheMaláFatraMts,theWesternCarpathianswasstudiedintheperiod1997{2001.A27.5ha(500550m)forestinteriorstudyplotwasestablishedforbirdcensusing.Populationabundanceswereestimatedbymeansofanim-provedversionofthemappingmethodfromApriltomid-July.Altogether,57birdspecies(52breeders)wererecordedwithinthereserveduringthe5-yearperiod.Ofthese,48speciesbredinthestudyplotreachingameandensityof58.173.60pairs/10ha(CV=6.18%).Theyearassemblagedensitiesvariedbetween52.15{61.20BP/10ha.TheShannondiversityindex()variedbe-tween4.10{4.36bites.Theevennessindex()reachedvaluesbetween0.78{0.82.Theyearrarefactionestimates[100pairs)]were22.77{25.16species.Theyearrarefactionestimatesonarea[10ha)]variedbetween19.46{21.30species.BirdspeciesdiversitywascomparabletoothermixedprimevalstandsintheWesternCarpathianse.g.DobroèandBadínreserves.Intotal,sevenspecieswerecharacterizedasdominant(5%):FringillacoelebsErithacusrubeculaSylviaatricapillaParusaterPhylloscopuscollybitaRegulusreg-,andPrunellamodularis.Thespeciesstructureandrelativeabundanceofthebirdassemblageshowedhightemporalstability.Majordierenceswerefoundinassemblagecompositionincomparisontothepastsurveyinthereserve.Aseriousneedforreevaluationandnewrigorouslydesignedquanti-tativeinventorysurveysoftheSlovakreservenetworkisstressed.Keywords:birdcommunity,primevalbeech-rforest,mappingmethod,pop-ulationdensity,rarefaction,the©rámkováNationalNatureReserve,Slovakia.IntroductionNaturaloldgrowthforestsrepresenttheuniqueremainsoforiginalecosystemsandusuallyfunc-tionascoreareasinmostecologicalnetworks.Theyareidentiedasareasofhighnatureconser-vationvaluefromthelandscapeecologyaspect.InSlovakiamostprimevalforestscreateaback-boneoftheexistingnaturereservesystemandinthenearfuturewillbetransformedintospe-cialareasofconservation(SAC)orspecialprotec-tionareas(SPA)meetingthecriteriaofNATURA 2000.Therefore,itiscrucialtodescribeandana-lyzestructureoftheseecosystemsforthemanage-mentofothermanmadeforestecosystemsandnatureconservationaims.Duetothesereasons,theoriginalfragmentsofclosetoprimevalornaturalforestwerethesub-jectoflongtermstudiesintheWesternCarpathi-ansforalongtime.TherstmonitoringplotsweresetupbyProf.Dr.A.ZLATNÍKintheECarpathiansintheperiod1935{1938(VOLO©ÈUK1998).Later,surveysoforiginalrepresentativefor-estecosystemswereconductedinordertoestab-lishanetworkofnaturereserves(ZLATNÍK,1959)thatlaterbecomepartoftheecologicalnetworksysteme.g.territorialsystemsofecologicalstabil-ity,ECONET,etc.ThenaturereservesoftheMaláFatraNationalParkwerealsothesubjectofba-sicinventorysurveysfornatureconservationandalsocomplexquantitativeecologicalstudies(e.g.TOLLMANN,1981;K,1998a).Duringthe1990'srigorousquantitativeinventorysurveyswerecarriedoutinselectedforestreservesinthenationalparktodetectchangesinecosys-temstructureandevaluatethequalityofpreviousstudiesanddevelopmethodologyforquantitativeinventorysurveys.The©rámkováNationalNatureReserveisconsideredtobethemostvaluablereserveinthenationalparkfromaforestryperspective.There-serverepresentsaWCarpathianbeech-rforestclosetotheprimevalstage.In1996,thereservewasselectedasamodelecosystemtostudycom-plexecologicalquestionsrelatedtotheorganiza-tionandfunctioningofbirdassemblagese.g.guildstructure,resourcepartitioning,interspeciccom-petition,ecomorphologicalpatterns,habitatselec-tionandpopulationdynamicsofbirdsundernearprimevalconditions.Theresultswillalsobeuti-lizedfornatureconservationpurposestoimproveforestmanagementpractices.The©rámkováReservehasalreadybeenthesubjectofaclusterofornithologicalandecologicalstudies(e.g.J,1988;K,1998b,2000,2001,2002;KDAMÍK,1999,inpress;DAMÍKetal.,2003).Mostofthecitedstud-iesdealtwithguildstructureandresourceparti-tioningexcepttheolderfaunisticpaperofJ(1988)withsimplequantitativeanalyses.Thisistherstpaperbasedontheunpublishedbirdin-ventorysurveys(K,2001,2002)dealingwiththeanalysesofbirdcommunitystructure.Theprimaryobjectivesofthepresentpaperareasfollows:(i)toanalyzethequantitativestructureofbreedingbirdassemblagecensusedbythemap-pingmethodovertheveyearperiod;(ii)toana-lyzepopulationdensity,dominance,speciesdiver-sity,andrelationshipbetweenanumberofspeciesagainstthetotalnumberofbreedingpairs(rar-efactioncurves)ofthebreedingbirdassemblage;(iii)toaddnewdataandcontributetothecurrentknowledgeofthebreedingbirdassemblagestruc-tureofmixedclosetoprimevalforestsintheWCarpathians;(iv)toreviewandevaluatepreviousbirdinventorysurveysinthereserve.StudyareaThestudywasconductedinthe©rámkováNationalNatureReserve,theMaláFatraMts(49E).TheMaláFatraMtslieinNWSlovakia.Thecorezoneofthereserveisspreadover243.65ha.Theinvestigationwascarriedoutina27.5ha(500550m)forestinteriorstudyplotrepresentingthecli-maxstageofaWCarpathianbeech-rforestgrowingonhillsidegranitewaste.Theforesthasallthechar-acteristicfeaturesofprimevalstands,however,someplantsspeciese.g.silverrarenegativelyaectedbyairpollutants.Thestudyplotissituatedattheele-vationof825{1123ma.s.l.(measuredbytheMagel-lanGPSProMARKX).ThereservebelongstoacoldmountainclimaticzonewiththemeanJulyairtem-peratures10{12C.Totalmeanannualprecipitationvariesbetween900{1200mm(VOLO©ÈUK,1986).Theslopeinclinationis20{60Theforestisunevenagedwithconsiderablever-ticalandhorizontalheterogeneity.Alldevelopmentalstagesofaprimevalbeech-rforestoccurinthestudyplot.Theageoftreesisapproximately200{250years.SpatialstructureofthedevelopmentalstageststheWatt's`gapphase'model,whichexplainsthemecha-nismsbehindtheregenerationofclimaxforests.Theoriginalplantspeciescompositionhasbeenpreserved.Thetnessofthesilverrpopulationissignicantlydecreasedbytheemissionofairpollutants.ThestudysiteisdominatedbybeechFagussylvaticaL.(44.8%),silverrAbiesalbaMill.(20.2%),NorwaysprucePiceaabies(L.)Karst.(4.8%),sycamoreAcerpseudopla-tanusL.(4.3%),wychelmUlmusglabraHuds.em.Moos(2.9%),androwanSorbusaucupariaL.emend.Hedl.(2.4%)withamixtureofothertreespeciessuchas,silverbirchBetulapendulaRoth,EuropeanlarchLarixdeciduaMill.,NorwaymapleAcerplatanoidesL.,andsmall-leavedlimeTiliacordataMiller.Stand-ingdeadtreesreachdominanceof14.7%.Thedomi-nanttreespecieshavestrongregeneration.Thecanopyheightisupto45m.ThescrublayermainlyconsistsofCorylusavellanaL.,red-berriedelderSambucusracemosaL.,currantRibesspp.,andsamplingsofthedominanttreespecies.Theherblayerismainlycom-posedofschottmalefernsDryopterislix-mas(L.)Schott,lady-fernsAthyriumlix-femina(L.)Roth,forbsberriesRubusspp.,IndianbalsamsImpatiensglanduliferaRoyle,ragworthsSenecionemorensisL.,wood-sorrelsOxalisacetosellaL.,sweetwoodrusiumodoratum(L.)Scop.,coral-wortsDentariabulbi- feraL.,perrenialhonestiesLunariaredivivaL.,purplecolt'sfeetHomogynealpinaCass.,wood-rushesLuzulanemorosa(Poll.)E.Mey.,small-reedsCalamagrostisarundinacea(L.)Roth.,andsmallscrubsbilberriesVacciniummyrtillusL.AdetaileddescriptionofthetreelayerisgivenbyKORÒAN(2000,2002).TheselectedstudyplotrepresentsmainlythegroupofforesttypesFageto-AbietumFageto-AceretumsensuZlatníkforestphytosociologicalap-proach(VOLO©ÈUK,1986)thatcorrespondstotheLuzulo-Fagionalliance,associationAbieti-Fagetafol-lowingtheBraun-Blanquetclassicationsystem.MaterialandmethodsBirdcensusingPopulationdensitieswereestimatedbythecombinedversionofthemappingmethod(TOMIA,1980;ORÒAN,1996).Inordertoconstructaneectiveori-entationsystemwithinthestudyplot,a5050mgridsystembasedoncolorplastictapemarkingontreetrunkswasestablishedinthe27.5harectangu-larstudyplot.Breedingbirdcensuseswerecarriedoutintheyears1997{2001fromApriltomid-July.Into-tal,10{11validcensusvisitsperbreedingseasonwereperformedinthetimeperiodbeginningat04:30andendingusuallyby9:00CET(sometimesby10:00CET)formorningvisits,from16:00to19:30CETforeveningvisits,andfrom19:00to22:00CETfornightvisits.Theproportionofeveningvisitswasalwaystwooutofthetotalnumberofvisits.AtthebeginningofAprilonenightvisitfocusedonowlregistrationswascarriedout.Thestudydesigncontrolledtheeectsofweatherandseasonondensityestimates.Eachvisitinvolvedwalkingandmapping100malternategridlinesbeginningatoneplotedgeandend-ingontheopposite.Thestartingpointanddirectionofobservermovementwereregularlychangedsothatcen-sustimingduringtheseasonwassimilarbetweendif-ferentpartsoftheplot.ThespeciesnameabbreviationsystemandmappingsymbolsystemforbirdactivityfollowedKROPIL(1992).Whilecensusing,allacous-ticandvisualobservations,nestsfoundorotherim-portantdatarelatingtobirdoccurrenceanddispersalpatternswererecordedontothevisitmapsat1:1667scale.Specialattentionwaspaidduringcensusvisitstocontemporarycontactsofterritorialsingingmalessothattheneighboringterritoriescouldbecorrectlydis-tinguished.Somebirdspeciese.g.TurdusphilomelosMuscicapastriataCerthiafamiliarisColumbassp.,Pyrrhulapyrhulla,andCoccothraustescoccothraustescausedconsiderableproblemsduringthecensusandinterpretationalprocedure.Censusingandtheidenti-cationofterritoriesonthespeciesmapswasbasedontherecommendationsofNILSSON(1977),S(1978,1980),TOMIA(1980,1994),TOMIAONTKOWSKI(1989),MOROZOV(1994),andK(1996).Inordertocorrectlyidentifythepropor-tionsofedgeterritoriesinthestudyplots,birdobserva-tionswererecorded100mbeyondtheplotedgelines.Overlapofedgeterritoriesinsidethestudyplotwasestimatedat1/4(25%),1/3(33%)or1/2(50%);how-ever,onlyspecieswithabundanceequaltoorhigherthen0.5pairperstudyplotwereincludedintheto-talcountofbreedingpairs(territories).FurtherdetailsregardingthemappingprocedureandtheprinciplesofspeciesmapanalysesaregiveninKORÒANObservationsofindividualspeciesfromvisitmapsweretransferredtospeciesmapsbeginningwiththerstspeciesrecordintheplot.Thecriteriaforterri-toryinterpretationwereprincipallybasedontheIBCC(1969)recommendations.However,inthecaseofse-cretivespeciesorspecieswithpoorlyevolvedterrito-rialbehaviour(discussedabove),speciesspecicmini-mumnumberofregistrations(acceptancelevelofter-ritory),andothercriteriarequiredtoacceptaclusterofregistrationsasaterritorymayhavebeenmodi-ed(S,1978).Especiallyusefulinformationondispersalpatternsofthesespeciesintheplotweregainedduringindependentplotvisitswhenforagingbirdobservationswererecordedthatwerepartofan-otherstudy(KORÒAN,1998b;ADAMÍKetal.,2003).Inthecaseofspecieswithabundancelessthan0.5pairsperplot,onlybreedingpresence"+"denotingthesta-tionaryoccurrenceofapartofbirdterritorywithintheboundariesoftheplotwasrecorded.Thissymbolwasprimarilyusedforspecieswithterritorysizesmuchlargerthanthestudyplotsizesuchassomewoodpeck-ers,owls,birdsofprey,andcorvids(TOMIA,1980;ORÒAN,1996).StatisticalanalysesBirdcommunitystructurewasanalyzedonpopula-tionabundance,density,speciesdiversity,evenness,andspecies{arearelationship(rarefaction).Standarddeviation(SD)andcoecientofvariation(CV)wereappliedtoestimatevariationbetweenyears.Standarddeviationwasappliedtomeasurethevariabilitybe-tweenyears.Theuseof1inthedenominatorwasappliedinsteadof.Coecientofvariationwasap-pliedtomeasuretherelativedispersioninthesample.Itisthestandarddeviationdividedbythemean(CV=SD/100).Specieswithabundancelowerthat0.5pairsperstudyplothada0valueforthecalculationsofvariationmeasures.Speciesdiversityandevennessweremeasuredbythreecommonformulae{Shannondiversityindex)asaninformationtheorymeasure,Simpsonin-dex()asameasureofconcentration,andtheBril-louinindex()(MAGURRAN,1991).Inaddition,rarefactionasanalternativetotraditionaldiversityindiceswasalsoapplied(HURLBERT,1971;HECKal.,1975).Toincludespecieswithverylowpopulationdensities(\+")inthecomputationofspeciesdiver-sityandevenness,constantnumbersofdensities(seeTab.1fortheconstantvalues)wereaddedtothesespecies.Similarityinstructureofassemblageswasmea-suredbythequalitativeSörensenindexandquantita-tiveCzekanowski-Sörensenindex(MAGURRAN,1991).SpeciesdiversityindicesandevennesswerecalculatedusingthePCstatisticalpackageNuCoSa1.0(TÓTHMÉRÉSZ,1993).Variabilitymeasures,assemblagesim- ilarityindices,rarefactionvaluesandcurveswerede-terminedusingMSExcel.Onlythemathematicalformulaeforthecalcula-tionofgeneralevennessandrarefactionarepresentedfurther:Evenness(equitability):DIV DIVmaxwhere:DIV{speciesdiversitymeasuredaccordingtoShannon,Simpson,orBrillouinformulas,DIVmaxmaximaltheoreticalvalueoftheseindices.Rarefaction(JAMESATHBUN,1981): where:)isanexpectednumberofspeciesinarandomsampleofindividualsdrawnwithoutre-placementfromindividuals,isthetotalnumberofspeciesfoundinthestudyplotinayear,isthenumberofindividualsinspeciesResultsSpeciesstructureIntotal,57birdspecieswererecordedwithinthe©rámkováNationalNatureReserveduringthepe-riod1997{2001(Tabs1,3).Themeannumberofspeciesoccurringinthestudysitewas47.60.Themaximumnumber,50species,wasfoundin1998,whiletheminimumnumber,45species,wasfoundin2000.Altogether,52breedingspecieswerede-tectedduringthestudiedperiodinthereserve.Themeannumberofbreedingspeciesinthere-servewas42.Thehighestnumber,45species,wasdetectedin1998,whereasthelowestnumber,40species,wasdetectedin2000.57recordedspeciesbelongedto9ordersaccordingtothemorpho-logicalsystematicsasfollows:Falconiformes{4species(7.02%),Galliformes{2species(3.51%),Charadriiformes{1species(1.75%),Columbi-formes{2species(3.51%),Cuculiformes{1species(1.75%),Strigiformes{2species(3.51%),Apodiformes{1species(1.75%),Piciformes{5species(8.77%),andPasseriformes{39species(68.42%).Inthe27.5hastudy,therewereatotalof48breedingspecies.Themeannumberofspecieswas38.80.Themaximumnumber,42species,ofbreederswasfoundin1999,whiletheminimumnumber,37species,weredetectedin1997and2000(Tabs1,3).Thirty(62.50%)breederswerepresentineveryyear.Other6species(intotal,36species:75.00%)bredinatleastthreeyearsduringtheve-yearperiod.Carduelisspinuswasdetectedduringthebreedingseason,usuallyfromtheendofMay,howeverbreedingwasassumedonlyin1999whenastationarysingingmalewasrecordedseveraltimesinonelocation.However,thepossibilityofbreedingcannotbeexcludedinotheryears.Also,recordinginconspicuouscifragacaryocatactescausedseriousproblemsduetopoorlyevolvedterritorialbehavior.ThesimilarityofspeciesstructurebetweenindividualyearswascomparedbythequalitativeSörensenindexandquantitativeCzekanowski{Sörensenindex.ThequalitativeSörensenindexreachedvalues0.84{0.95.Thelowestspeciessim-ilaritywasfoundbetweenyears1997and1999,andthehighestsimilaritylevelwasfoundbetweenyears2000and2001(Tab.2).TheCzekanowki{Sörensenindexreachedvalues0.79{0.88.Thelow-estvaluewasfoundbetween1997and2001,whereastheassemblagesin1999{2000and2000{2001reachedthehighestlevelsofsimilarity(Tab.2).Similaritymeasurementsindicatedverylowbetweenyearvariabilityoftheassemblagefromqualitativeandquantitativespeciesstruc-ture.Thus,thestudiedcommunityseemstobeverystableregardingspeciesstructure.DensityanddominanceThemeantotaldensitywas58.17BP/10ha.Stan-darddeviation(3.60)andcoecientofvariation(6.18%)valueswererelativelysmallindicatingverylowbetweenyearvariabilityintotalden-sityoftheassemblage.Theassemblagereachedthehighesttotaldensity,61.20BP/10ha,intheyear2000,thelowesttotaldensity,52.15BP/10ha,wasdetectedin1997(Tab.1).Dominantspecies(5%)contributedbe-tween51.33{63.46%tothetotalassemblagedomi-nance.Onaverage,thesesevenspeciescontributed59.44%tototaldominanceusingpooleddata.Allofthesespeciesreachedahigherpopulationden-sitythan2.5BP/10ha.ThehighestvaluesofdensityanddominancewerereachedbyFringillacoelebsineveryyear.Itsdominanceslightlyex-ceeded20%ofthetotalassemblagedominanceduringthestudyperiod.Basedonthepooleddata,ErithacusrubeculaParusaterSylviaatr-icapillaRegulusregulusPhylloscopuscollybitaandPrunellamodularisweredominantspecies.Fromthisgroup,onlyErithacusrubeculawasdominanteveryyear.Theremainingspecieswerenotdominantinatleastoneyear.In1997,Ficedulaalbicolliswasalsoadominantspecies.Thisisgenerallyaneectofpopulationdynamics Table1.Yearandmeanabundance,density,anddominanceofthebreedingbirdassemblageofaprimaevalbeech-rforestinthe©rámkováNationalNatureReserve. AbundanceDensity(pairs/10ha)Dominance(%)SDCVSpecies 199719981999200020011997199819992000200119971998199920002001(%) FringillacoelebsL.,175825.725.634.335.734.59.359.3112.4712.9812.5511.3317.9215.7921.5021.2120.7219.481.8416.23Erithacusrubecula(L.,1758)10.714.515.616.211.23.895.275.675.894.074.967.468.959.789.636.738.530.9218.59Sylviaatricapilla(L.,1758)8.77.612.415.914.03.162.764.515.785.094.266.074.697.779.458.417.331.2829.93ParusaterL.,175813.515.37.08.810.04.915.562.553.203.643.979.419.444.395.236.016.831.2431.24Phylloscopuscollybita(Vieillost,1817)7.19.810.311.510.62.583.563.754.183.853.594.956.056.466.836.376.160.6016.86Regulusregulus(L.,1758)8.011.19.39.79.32.914.043.383.533.383.455.586.855.835.765.595.930.4011.71Prunellamodularis(L.,1758)6.78.87.59.09.52.443.202.733.273.453.024.675.434.705.355.715.190.4213.97Ficedulaalbicollis(Temminck,1815)7.56.95.36.36.02.732.511.932.292.182.335.234.263.323.743.604.000.3113.17CerthiafamiliarisL.,17586.44.95.77.87.02.331.782.072.842.552.314.463.023.574.634.203.980.4117.67ColumbapalumbusL.,17586.97.07.36.52.02.512.552.652.360.732.164.814.324.583.861.203.710.8137.39ColumbaoenasL.,17585.33.03.55.011.01.931.091.271.824.002.023.701.852.192.976.613.481.1657.42Troglodytestroglodytes(L.,1758)4.55.95.06.06.41.642.151.822.182.332.023.143.643.133.573.843.480.2814.08TurdusmerulaL.,17584.54.53.53.66.21.641.641.271.312.251.623.142.782.192.143.722.790.3924.28TurdusphilomelosBrehm,18311.85.53.24.84.70.652.001.161.751.711.451.263.392.012.852.822.500.5437.21Phylloscopussibilatrix4.74.04.31.04.51.711.451.560.361.641.353.282.472.700.592.702.310.5641.39(Bechstein,1793)SittaeuropaeaL.,17581.84.02.84.05.60.651.451.021.452.041.321.262.471.752.383.362.280.5239.32Pyrrhulapyrrhula(L.,1758)3.34.52.01.01.01.201.640.730.360.360.862.302.781.250.590.601.480.5564.58Muscicapastriata(Pallas,1764)3.03.03.8o+0.51.091.091.380.10.180.752.091.852.380.000.301.290.6282.21Ficedulaparva(Bechstein,1794)3.51.01.52.51.51.270.360.550.910.550.732.440.620.941.490.901.250.3650.00Coccothraustescoccothraustes2.01.03.02.00.50.730.361.090.730.180.621.390.621.881.190.301.060.3557.33(L.,1758)TurdusviscivorusL.,17581.82.01.02.01.00.650.730.360.730.360.571.261.230.631.190.600.980.1933.19TurdustorquatusL.,1758+2.41.51.01.30.10.870.550.360.470.450.001.480.940.590.780.780.3269.98Phylloscopustrochilus(L.,1758)+1.82.20.51.30.10.650.800.180.470.420.001.111.380.300.780.730.3378.21ParuspalustrisL.,17581.01.01.01.51.30.360.360.360.550.470.420.700.620.630.890.780.730.0819.85Dendrocoposleucotos1.00.50.81.50.80.360.180.290.550.290.330.700.310.500.890.480.580.1340.23(Bechstein,1803)MotacillacinereaTunstall,17711.01.01.01.00.50.360.360.360.360.180.330.700.620.630.590.300.560.0824.85Regulusignicapillus(Temminck,1820)|1.01.51.01.00.000.360.550.360.360.330.000.620.940.590.600.560.2060.86Picoidestridactylus(L.,1758)0.52.0+1.01.00.180.730.10.360.360.330.351.230.000.590.600.560.2782.40ScolopaxrusticolaL.,17581.01.0ppp0.360.360.000.000.000.150.700.620.000.000.000.250.20136.93LoxiacurvirostraL.,1758|p1.0p1.00.000.000.360.000.360.150.000.000.630.000.600.250.20136.93Aegithaloscaudatus(L.,1758)0.5|0.25+0.80.180.000.090.10.290.110.350.000.160.000.480.190.12110.81Bonasabonasia(L.,1758)|1.0++0.50.000.360.10.10.180.110.000.620.000.000.300.190.16149.07ParusmajorL.,1758||0.51.0|0.000.000.180.360.000.110.000.000.310.590.000.190.16149.07Dendrocoposmajor(L.,1758)1.0+||o+0.360.10.000.000.050.070.700.000.000.000.000.130.16223.61 Table1.(continued) AbundanceDensity(pairs/10ha)Dominance(%)SDCVSpecies 199719981999200020011997199819992000200119971998199920002001(%) Anthustrivialis(L.,1758)|+0.50.5+0.000.10.180.180.10.070.000.000.310.300.000.130.10136.93CuculuscanorusL.,1758++0.5++0.10.10.180.10.10.040.000.000.310.000.000.060.08223.61ParuscaeruleusL.,1758|0.5||o+0.000.180.000.000.050.040.000.310.000.000.000.060.08223.61Carduelisspinus(L.,1758)pp0.5pp0.000.000.180.000.000.040.000.000.310.000.000.060.08223.61Accipitergentilis(L.,1758)+++++0.0040.0040.0040.0040.004|||||||||Dryocopusmartius(L.,1758)+++++0.10.10.10.10.1|||||||||Garrulusglandarius(L.,1758)+++++0.10.10.10.10.1|||||||||Aquilachrysaetos(L.,1758)p++++0.000.0010.0010.0010.001|||||||||Cincluscinclus(L.,1758)++p++0.10.10.000.10.1|||||||||Glaucidiumpasserinum+p+o+|0.050.000.050.050.00|||||||||(L.,1758)Buteobuteo(L.,1758)+|||p0.0040.000.000.000.00|||||||||Pernisapivorus(L.,1758)||+||0.000.000.0040.000.00|||||||||StrixalucoL.,1758|o++o+|0.000.010.010.010.00|||||||||Nucifragacaryocatacteso+o+o+|+0.10.10.10.000.1|||||||||(L.,1758)Corvuscoronecornixo+o+|||0.050.050.000.000.00|||||||||L.,1758ParuscristatusL.,1758|o+|||0.000.050.000.000.00|||||||||PicuscanusGmelin,1788o+o+||p0.10.10.000.000.00|||||||||Pheonicuruspheonicuruso+o+|||0.10.10.000.000.00|||||||||(L.,1758)Delichonurbica(L.,1758)ppppp0.000.000.000.000.00|||||||||TetraourogallusL.,1758ppppp0.000.000.000.000.00|||||||||CorvuscoraxL.,1758p|||p0.000.000.000.000.00|||||||||Parusmontanusp||||0.000.000.000.000.00|||||||||Baldenstein,1827Apusapus(L.,1758)||p||0.000.000.000.000.00||||||||| Total143.4162.1159.55168.3166.552.1558.9558.0261.2060.5558.17100.00100.00100.00100.00100.00100.003.606.18 Key:SD{standarddeviation,CV{coecientofvariation.Markplussing(\+")indicatesbreedingabundancelessthe0.5territory(pair)perstudyplot;mark\o+"indicatesbreedingpresenceinthereserve,butthespecieswasnotdetectedasbreederinthestudyplot;mark\p"isusedforspeciesdetectedinthestudyplotasnonebreedersorrarevisitors;mark\|"indicatesabsence.Inthedensitycolumns,densityestimatesfor\+"and\o+"speciesaregivenbyqualiedguessbasedonobservationsinthereserveandthenationalpark.Thedensityestimatesinparenthesiswereonlyroughlyestimatedforcalculationofdiversityindicesandrarefactionoftheassemblage.Polygamouspair,threeadultbirds. { uctuationsandpopulationdeclinesinindivid-ualpopulations.Intotal,ninespeciesweresubdominant(2%)andrepresented28.52%ofthetotalas-semblageusingthepooleddata.Theirmeanpop-ulationdensitywashigherthan1.5BP/10haandlessthan2.5BP/10ha.Recendentspecies(1%)constituted5.08%oftheassemblage.Intotal,fourspeciesrepresentedthisgroup.Pop-ulationdensityusuallyvariedbetween0.60{1.50BP/10ha.Thehighestnumberofspecies,28,wasfoundforthegroupofsubrecedents(1%),yetrepresentedonly6.97%ofthetotaldominance.Allspeciesreachedmeanpopulationdensitieslowerthan0.60BP/10ha.Anotherfourspecies,PicusParuscristatusPheonicuruspheonicurusandCorvuscoronecornix,thatwouldbelongtothisgroupbredoutsidethestudyplot.Thehighestlevelofbetweenyearvariationinspeciesbreedingpresenceinthestudyplotoccurredinthisgroup.Tenspeciesbredonlyinthreeorfeweryearsinthestudyplotduringthetimeperiod(Tab.1).DiversityandevennessStandarddiversityindicessuchasShannon,Simp-son,andBrillouinwereappliedtoestimatespeciesdiversityandevenness.Theseindiceswereselectedforcomparativepurposeswithpreviousstudieswheretheyhavebeenwidelyusedincommunityecologystudiesformanydecades.However,severalauthors(e.g.HURLBERT,1971;JAMESATHTable2.Betweenyearcomparisonofthebirdas-semblagesimilarityinspeciesstructurebyqualitativeSörensenindexanddensitystructurebyCzekanowski-Sörensenindicex.Onlybreedingspeciesrecordedinthestudyplotwereincludedtotheanalyses. SimilarityCzekanowski-Sörensenindices Sörensen19971998199920002001 1997|0.860.820.800.7919980.89|0.840.820.8119990.840.86|0.880.8620000.860.920.91|0.8820010.870.920.930.95| BUN,1981)concludedthatmostofthesecom-monlyusedmeasuresofbiologicaldiversity,in-cludinginformationtheorybasedindices,areinmanysituationsinappropriatelyusedasindicatorsofbiologicaldiversity.Moreover,theirapplicationinvolvesasignicantlossofinformationbecauseindicesconfoundseveralcommunityparameterse.g.numberofspecies,theirrelativeabundance,andareasampledintoonenon-metricnumber.Insummary,dierencesattributabletotheaccumu-lationofspecieswithincreasingareaareignored,andmanycombinationofspeciesrichnessandrel-ativeabundancecanproducethesamevalueoftheindex(JAMESATHBUN,1981).OneoftheTable3.Estimatesofbirdspeciesdiversityandevennessofthebeech-rprimevalforestinthe©rámkováNationalNatureReservebystandardindicesandrarefaction.Onlybreedingspeciesrecordedinthestudyplotwereincludedtodiversitycalculations. 19971998199920002001MeanSDCV DiversitymeasuresTotalnumberofspecies475048454847.601.934.06Totalnumberofbreeders414543404142.002.135.07Numberofbreedersinplot373942373938.802.185.62Shannon()4.274.364.254.104.154.230.112.58Brillouin()3.853.953.833.733.783.830.092.30Simpson()0.930.930.920.910.920.920.010.97Rarefaction50pairs)19.3920.0019.4818.0818.2719.040.884.63Rarefaction100pairs)24.0225.1624.9322.7722.9023.961.184.93Rarefaction5ha)14.4415.6315.0314.4714.7114.860.523.53Rarefaction10ha)19.6721.3020.7019.4619.6220.150.864.27 Evennessmeasures EvennessShannon()0.820.820.780.790.790.800.022.49EvennessBrillouin()0.830.830.790.800.800.810.022.46EvennessSimpson()0.950.880.940.940.940.930.033.24 ([SHFWHGVSHFLHV(6Q ([SHFWHGVSHFLHV(6QFig.1.Rarefactioncurvesforveyearsamplesofthe27.5habeech-rinteriorplotinthe©rámkováNationalNatureReservepredicttherateofspeciesaccumulationwithincreasingnumberofterritorialpairs(grapha)andarea(graphb).Thecalculationsarebasedontheyearabundancedata.alternativestoovercomethisproblemistheappli-cationofrarefaction(HURLBERT,1971;HECKal.,1975).Inthisstudy,rarefactionwasappliedinordertocomparebetweenseasonalspeciesrich-nessbystandardizingsamplestoanequalnumberofindividualsandequal-sizedplots(Tab.3).Therelationshipbetweenexpectednumberofspeciesandincreasingnumberofpairsandareawasan-alyzed(Figs1a,b).Evennesswasexpressedbyagraphofspeciesdensitiesinadecreasingway(Fig.2).Valuesofalldiversityindiceswereverysta-ble(Tab.3).Standarddeviationvalueswereclosetozero.TheShannondiversityindexreachedameanvalue4.23.TheShannonindexvaluesvariedbetween4.10{4.36bites.Nosignicantdierenceswerefoundbetweenyearcomparisons.ThemeanvalueoftheBrillouindiversityindexwas3.83.Thevaluesvariedbetween3.73{3.95.ThisindexhadasimilartrendastheShannonindex(Tab.3).ThevaluesofSimpsondiversityindexwerethemostregular,mean0.92,variation0.91{0.93.Evennessindiceswerealsoverystable.Themeanvalueswereasfollows:Shannon{0.80,Simpson{0.93,Brillouin{0.81.EvennessvaluesbasedontheShannonandBrillouindiversityindiceshadsimi-lartrends.TherewereonlyveryslightdierencesbetweenyearsinSimpsonevennessnumbers.Thehightemporalstabilityinthevaluesofdiversitymeasuresindicatesahighstabilityofspeciesstruc-tureoftheecosystemaswellastherelativeabun-danceofspecies.SimilarvalueswereestimatedinotherstructurallysimilarWCarpathianclosetoprimevalmixedforests(seeDiscussion).Rarefactionspecies-individualandspecies-areacurvesareshownintheFigs1a,b.Themeanexpectednumberofspeciesper50pairswas19.040.88species(Tab.3).Onarandomsampleof100pairs,itwas23.961.18species.Rarefaction100pairs)valuesvariedbetween22.77{25.16.Valueshaddierenttrendsforeachstandardizednumberofpairs.Themeannumberofexpectedspeciesonthestandardizedareaof5hawas14.860.52species,onanareaof10ha,itwas20.150.86species.Thenumberofexpectedspeciesontheareaof5havariedbetween14.44{15.63species,thevariationontheareaof10hawas19.46{21.30species.Thehighestspeciesdiversityestimatedbyalldiversitymeasureswasfoundin1998,thelowestdiversitywasdetectedintheyear2000alsobyalldiversitymeasuresexcepttherar-efactiononastandardizedareaof5ha.Alleven-nessmeasuresdetectedthehighestvaluesin1997,whereasthelowestvalueswerein1999excepttheSimpsonevenness.DiscussionAssemblagestructureanddiversityTherstquantitativestudiesofforestbirdassem-blagesinSlovakiawerepublishedinthersthalfof1950's.Themainimpetustoquantitativecom-munityecologycamefromtheworkoftheSlovakecologistF.J.Turèekandhiscolleagues.TURÈEK(1952)developedhisowntime-quadratmethodtostudybirdassemblageinnaturalgroupsoffor-esttypesFagetumtypicumandPiceetumtypicum(GFTsensuZLATNÍK,1959)inthePoµanaMts. 63(&,(625'(51 'HQVLW\SDLUVKD Fig.2.Communitycurveofthebreedingbirdassemblageoftheprimevalbeech-rforestinthe©rámkováNationalNa-tureReserve.Pooleddatafromthepe-riod1997{2001wereused.Speciesarerankedindescendingorder.Theauthoranalyzedthedensity,biomass,forag-inggroups(guilds),anddistributionofspeciesinverticalstrataintheforest.Thestudyhadaveryprogressivedesignandwasaheadofitstimeinmanyaspects.Theauthor,morethan14yearsbeforetheocialprinciplesoftheconceptofeco-logicalguilds(R,1967)wereformulated,hadthoughtaboutfunctionalgroupsofspecieswithsimilarforagingrequirements,feedinghabitsandbehavior,andpreferredstratumoffoodconsump-tion.Todate,moststudieshavedealtwithvari-ousanalysesofbroadleafforestassemblages.Rel-ativelyfewstudieshavebeenfocusedonanalysesofbirdassemblagesofmixedforests.Moststud-iesarestillnotcomparablewiththepresentstudyduetodierencesinsurveymethodology,habitatclassication,andstudyobjectives.FERIANCOVÁASÁROVÁ(1968,1971,1978)describedthequal-itativestructureofbirdassemblagesofallmainhabitatsonaregionalscaleinLiptov.Theau-thoralsostudiedassemblagesofmixedforest;however,duetodierencesinthehabitatclas-sicationschemeitisnotpossibletocomparetheresultsbecausemixedforestsweregroupedwithconiferousforestsintoonehabitatcategory.ERIANCOVÁASÁROVÁetal.(1987)analyzedbirdassemblagesoftheGFTFageto-quercetumwithamixtureofspruceandrintheSitnoNa-tionalNatureReserve,the©tiavnickévrchyPro-tectedLandscapeArea.Unfortunately,onlycom-binedresultsforallhabitattypesinthereservewerepresented.Consequently,itisnotpossibletoextractdataforindividualhabitatsandcomparethemwiththepresentstudy.TOPERCER(1989)studiedthequantitativestructureofbreedingbirdassemblagesoffourhabitatsintheSkalnáalpaNa-tionalNatureReserve,theVeµkáFatraMts,usingabelttransectmethod.Hefoundatotalof14speciesinthetotalbreedingbirddensityof62.86ex./10ha(31.43BP/10ha)ina200{240yearoldmixedforest(GFTAbieti-fagetasuperioraSpeciesstructureaswellaspopulationdensitiesofindividualspeciesseemtobeunderestimatedforallspeciesexceptthechichaPhylloscopuscolly-bita.Thisspeciesisconspicuous,singsthroughouttheday,andcanbeeasilymappedwithhighaccu-racy(TASTIAN,1983).Underestima-tionmayhavebeenprimarilycausedbyonlytwovisitstothestudysiteinoneyearandthedif-ferencesincensusmethodology.Theauthorcom-binedthreecensustechniquesforestimatingpop-ulationdensities(mappingmethod,pointcount,andbelttransect).Breedingbirdassemblagesinthe©rámkováNationalNatureReservewerestudiedusingatransectmethodbyJ(1988)between1982-84.Theauthorstudiedbirdassemblagesinthreehabitattypes:beech-randbeech-sycamorefo-rests,andmeadowgaps.Inthebeech-rstand,theauthorfoundatotalof34speciesatthedensityof489ex./100ha(24.45BP/10ha).Thebeech-sycamorestandhadonly26speciesattheto-taldensityof520ex./100ha(26.00BP/10ha).Thedataappearstosignicantlyunderestimatespeciesrichness,andalsothemeanassemblageaswellasindividualpopulationdensities.Themeanassemblagedensitiesareapproximatelyhalfofthevaluespresentedinthispaper.Inaddi-tion,commonbreederssuchasthecollared y-catcherFicedulaalbicollis,mistlethrushTurdusviscivorus,recrestRegulusingicapileus,greywag-tailMotacillacinerea,dipperCincluscinclus,werenotdetectedinanyhabitatorwereconsidered\rarebreeders"e.g.thestockdoveColumbaoenasIncontrast,othersspecies,forinstancetheeld-TurduspilarisandwhitethroatSylviacom-muniswererecordedascommonbreedersinforeststands.ThegardenwarblerSylviaborinwascon- cludedtobeararebreederinforesthabitats.Themostsurprising\discovery"istheassessmentofPrunellacollarisasararebreederinthemeadowgaps.Theseconclusions,theresults,andmethod-ologicaldesignofthestudyaremisleading,andtheinterpretationofspeciesoccurrencepatternsisquestionable.Theerrorsmayhavebeencausedbyalownumberofvisitstothestudysiteormoreprobablypoorskillsinidenticationofbirds(e.g.T.pilarisversusT.viscivorus).TheoccurrenceofthegreywagtailMotacillacinereaanddipperCincluscinclusinthe©rámkováforestwaseco-logicallydependentonastreamhabitat.Streamsdidnotrunthroughtheplotsanalyzedbelow,thustheoccurrenceofthesetwospecieswillnotbedis-cussedinfurthercomparisons.ANIGA(1994,1995)studiedbreedingbirdassemblagesofvegetationbeltsintheMaláFatraandVeµkáFatraMtsin1989{1991byastriptran-sectmethod.Sanigaconductedcensusesintran-sectswithatotallengthof3,300minther-beechbeltlocatedintworeserves.Thetotallengthoftransectsusedfordescriptionofthespruce-beech-rvegetationtierwas10,400mlocatedinsixre-serves.Hefoundatotalof57breedingspeciesatthemeandensityof75.7BP/10hainr-beechand59speciesatthemeandensityof60.2BP/10hainspruce-beech-rvegetationtiers.Themeanto-talassemblagedensityofspruce-beech-rtierwasnotsignicantlydierenttothe©rámkováforest.TheShannondiversityindexwas4.46forther-beechtierand4.33forthespruce-beech-rtier.Surprisingly,populationdensitiesanddominanceofthemostabundantspeciesareverysimilartotheresultsfromthe©rámkováforest,eventhoughthestudywascarriedoutbytherelativequantita-tivemethod.TheoccurrenceofthegardenwarblerSylviaborinandlesserwhitethroatSylviacurrucainther-beechbelt,andthebramlingFringillamontifringillainthespruce-beech-rbeltdeviatesfromresultsinotheroldgrowthforeststandsofthistypeinSlovakia(KROPIL,1996a,b).BothsilviidwarblerswerefoundbyG LOWACINSKIROFUS(1992)inanold-growthbeech-rstandintheTatryMtsinPoland.Slightlyhigherspeciesrichnessanddiversityisprobablyduetotheverylargesamplesizes.Duetoahighvariationinpopulationesti-matesproducedbydierentquantitativemeth-ods,onlymappingmethodstudiesaretakenintoconsiderationforfurthercomparisons.MixedprimevalforestswerestudiedontheSlovaksideoftheWCarpathiansbyKROPIL(1993,1996a,b).ROPILdescribedthestructureofbreedingbirdassemblagesintheDobroèandBadínprimevalforestNationalNatureReservesoverathree-yearperiod.Intotal,44specieswerefoundina24haplotintheDobroèforest,and42speciesina16haplotintheBadínforest.ThemeannumberofspeciesperyearintheDobroèforestwas36,witharange35{37.InBadínforestthemeannumberofspecieswas36,witharange34{38.TherewerenosignicantdierencesindiversityestimatedbytheShannonformulabetweenyearcomparisonofthesetwostands(K,2001).However,allBadínsampleshadsignicantlyhigherdiversityincomparisontothe©rámkovásamples.Theto-talmeandensityoftheassemblageintheDobroèstand(62.6BP/10ha)wasnotdierentfromthe©rámkováforestorthespruce-beech-rtierde-scribedbySaniga(seeabove).TheBadínstandhadhighertotalassemblagedensities{71.0BP/10ha.ThespeciesstructureoftheDobroèreservewasverysimilartothe©rámkováreserve.Elimi-natingraptorsandlargecorvids,tracespecieswithlessthan0.5territoryperstudyplot,theonlydif-ferencewasinregularbreedingofthewillowtitParusmontanusandcrestedtitParuscristatusthatcouldberelatedtothepresenceofconifers.ThecrestedtitParuscristatusbredin©rámkováforestonlyin1998ontheborderofthereservewithaspruceplantation.Thisdierencecanbearesultofthehigherrelativeproportionofconifer-oustreesintheDobroèforest(KROPIL,1996a)incomparisonto©rámkováforest(K,2000),53%to25%.ThewillowwarblerPhylloscopustrochilus,long-tailedtitAegithaloscaudatus,andtreepipitAnthustrivialiswereabsent,butwereregularnestersinthe©rámkováforest.Theoccur-renceofthelattertwospecieswasaresultoflargegapscreatedbyatornadointheearlyspringof1998.Astheresultofincreasedheterogeneityoftheforestinterior,thetreepipitA.trivialisbe-ganbreedingregularlyandthewillowwarblertrochilushaveincreasedpopulationdensitiessince1998.Bothspeciespositionedtheirterritoriesinthegapsorontheiredges.Densityanddomi-nancevaluesofmostspeciesweresimilarinbothforests.Thehighestdierenceinpopulationdensi-tieswasfoundforthecollared ycatcherFicedulaalbicollis.Itspopulationdensity(0.2versus2.33BP/10ha)in©rámkováwasapproximately11timeshigherthanintheDobroèforest.Incon-trast,thepopulationdensityofthisspecieswasal-mosttwicethesizeintheBadínforestcomparedtothe©rámkováReserve.TheblackcapSylviaatri-capillaandchichaPhylloscopuscollybitahadhigherdensitiesinthe©rámkováplot.SpeciesstructureoftheBadínforestbirdassemblagewasevenmoresimilar.Thelesserspottedwoodpecker DendrocopusminorandwillowtitP.montanuswereabsentinthe©rámkováforest,butthewil-lowwarblerPh.trochilus,ringouzelTurdustorqua-,andrecrestRegulusignicapilluswereabsentintheBadínforest,andthegoldcrestR.regulusreachedonlyverylowpopulationdensities.BadínforestistypicalforhighbreedingdensitiesofthestockdoveColumbaoenas.Ithadalmostthreetimeshigherpopulationdensitiesincomparisontothe©rámkováorDobroèforest.Thesedierencescanalsobeattributedtoalowerproportionofconiferoustrees,approximately18%,andaloweraltitude(720{760m)ofthestudysiteincompar-isontothe©rámkováorDobroèforests(720{1000m).Inaddition,furthervariationinspeciesstruc-tureandpopulationdensitiescanbeattributedtodierentverticalandhorizontalheterogeneityofthestudiedareas(ADAMÍKetal.,2003).The©ramkováReserverepresentsanecosysteminamosaicstateofseveraldevelopmentalstages,fromthestageofearlygrowingtothelatedevelopmen-talstagesthatcreateaverywideresourcespacesupportingawiderangeofspeciesfrom oris-ticspecialiststogeneralistsaswellasstructuralspecialisttogeneralists.Presumably,thespecicquantitativestructureof oristicparametersincombinationwithstructuralparametersre ectedineachdevelopmentalstagecreatingacomplexmosaicspatialstructureonalocalscaleeectsthenalstructureofthecommunitycomposition.AVELKAAVELKA(1990)studiedthreeprimevalbeech-rstands(Razula,Salajka,andKutavý),theMoravskoslezskéBeskydyMts,foraperiod6{7years.Thesizeofthestudyplotsvariedfrom14.9{23.2ha.Thetotalnumberofspecieswas29{33,andwasgenerallylowerincomparisontotheSlovakplots.Speciesstructurewasmostsimi-lartothe©rámkováforest,onlyfourbreeders,thewillowtitParusmontanus,swiftApusapus,pied ycatcherFicedulahypoleuca,andstarlingnusvulgarisdidnotoccurinthe©rámkováforest.TheredstartPhoenicurusphoenicuruswasasub-recendentspeciesintwoplots.Meandensityfromthethreeplotsvariedbetween45.5{51.6BP/10ha,however,betweenyearvariationwasstronger41{61BP/10ha.LOWACINSKIROFUS(1992)studied100{200yearoldbeech-rforestintheStrazyskaValley,thePolishTatryNationalPark,during1981{1982.Theyfoundatotalof38speciesina10hastudyplotwiththeoverallassemblagedensityof68.2BP/10ha.Numberofspeciesvar-iedbetween29{35.SpeciescompositionwasthemostdistinctiveincomparisontothethreeSlo-vakplots,butthepopulationdensitiesofmostofthespeciesweresimilarapartfromtherobinrubeculaandwoodwarblerPh.sibilatrixhavinghighernumbersandblackcapS.atricapillalowernumber.ThegardenwarblerSylviaborin,lesserwhitethroatS.curruca,greennchCarduelischlo-ris,pied ycatcherFicedulahypoleuca,andeld-Turduspilariswerelisted,butsometypicalspeciessuchasthewoodpigeonColumbapalum-,white-backedwoodpeckerDendrocopusleu-,collared ycatcherFicedulaalbicollisandmarshtitParuspalustisfoundinthistypeofhabi-tatinSlovakiawerenotrecorded.Theoccurrenceofsilviidwarblersandgreennchcouldhavebeenaneectoftheforestedge,afragmentedforestpatch,alsocausingincreasedspeciesdiversityinarelativelysmallplot.CHAFFNER(1990)foundonly28speciesina70haplotlocatedinAbieti-Fagetumnaturalforestreserve,theSwissJuraMts,duringaoneyearstudy.Thetotalmeanbreedingassemblagedensitywasslightlyhigherincomparisontothe©rámkováandDobroèReserves.TORCHOTECKÝ(1999)meta-analysed133breedingbirdassemblagesofallhabitattypes(woods,clearings,steppe,openmosaic,reed,andurbanhabitats)frompublishedquantitativedataintheCzechRepublic.Theyappliedcanonicalcor-respondenceanalysistostudytheeectsofhabi-tat,censustechniques,andareaonthegeneralpatternsofbirdassemblages.Assemblagestruc-turewasmeasuredintermsofspeciesrichness,indexofdominance,andevenness.Insummary,lowerspeciesrichnesswasdetectedformixedforests(=14,25.575.12)incomparisontodeciduousforests(=35,26.609.10),butwashigherincomparisontoconiferousforests(=16,14.58).AsimilarpatternhasbeenfoundinNorthAmericanstudies(JAMESATHBUN1981;JAMESAMER,1982).Thispatternis,however,questionablebecauseprimarydatawerenotrarefactioned,sothestrongeectsofplotsizeonspeciesrichnesscouldhavemajoreectsonthenalpatternsandconclusions(seeResults:Diver-sityandevenness).Incontrast,KROPIL(1993)detectedthehighestspeciesrichnessinmixedprimevalforestsincomparisontodeciduousorconiferousprimevalstands.K(1996)an-alyzedhisownandKropil'sdatabyrarefactionanddidnotndaclearpatternofspeciesrichnessbetweenmixedanddeciduousforests.AcknowledgementsIwishmanythankstotheAdministrationoftheMaláFatraNationalParkandtheStateNatureConservancy 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