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Chapter10Predation,2002byPrenticeHall,Inc.UpperSaddleRiver,NJ07458,Outline,Thereareavarietyofantipredatoradaptations,whichsuggeststhatpredationisimportantinnaturePredator-preymodelscanexplainmanyoutcomesFielddatasuggeststhatpredatorshavealargeimpactonpreypopulations,Outline,ExperimentsinvolvingtheremovalorintroductionofexoticpredatorsprovidegooddataontheeffectsofpredatorsontheirpreyFieldexperimentsinvolvingthemanipulationsofnativepopulationsshowpredationtobeastrongforce,Equilibriumtheoriesofpopulationregulation,A.Extrinsicbioticschool1.Foodsupplyandpopulationregulation2.Predationandpopulationregulation3.DiseaseandpopulationB.Intrinsicschool1.Stressandterritoriality2.Geneticpolymorphismhypothesis3.Dispersal,Thecausesofpopulationchange,keyfactoranalysis主導因子分析(一)Density-dependentfactor密度制約因子:(種內、種間因素)作用強度隨種群密度而變。Afactoraffectingpopulationsizewhoseintensityofactionvarieswithdensity.(二)Densityindependentfactor非密度制約因素(外界環境因素):havinganinfluenceonindividualsthatdoesnotvarywiththenumberofindividualsperunitareainthepopulation.,Density-dependentfactor密度制約因子:1.種間因素,.食物、空間資源種內、種間競爭.病蟲害傳播速度.個體成熟速度.體質和繁殖力、生長發育、自相殘殺、外遷.植物結實數量.抗逆性在橡樹蛾的生活史裡,有不同的生活環境,不同的掠食者,寄生、競爭、環境壓力,在不同時期裡會有不同的死亡率。,2.種間因素,.競爭.掠食、寄生.遺傳反饋機制(抗病種的培育)澳洲野兔粘液病毒抗病種,Densityindependentfactor,.氣候因素.土壤因素.營養.理化.空間.汙染,Extrinsicfactors:,Externalfactorsactingonpopulations.Predation,parasitism.Competitionforfooddensitydepended.Competitionforspacedensitydepended.Randomstochasticchangedensityindependent.Weather,1.種內因素,種群是一個具有自我調節(selfregulation)機制的生活系統,可以按照自身的性質及環境狀況調節它們的數量。植物的自疏現象禾本科植物的分的產生和生長遺傳特性(抗逆性)內分泌調節(旅鼠)Crowdingstress腎上腺髓質(adrenocorticotropin)腦下腺(Epinephrine)腎上腺皮質(Corticoids)危急反應Alarmresponse,Introduction,WolvesinYellowstonePark(Figure10.1)U.S.FishandWildlifeService,1980sReintroduceinYellowstoneParkandstabilizewolfpopulationsinMinnesotaandMontanaConcernsCattleranchersconcerned:Decimateherd?Arepredatorstiedtothehealthofthemainprey?Canpredatorsswitchprey?RamificationstoreestablishmentResults:Nomajoreffects,Introduction,PredationTraditionalview:carnivoryDifferencesfromherbivoryHerbivoryisnon-lethalDifferencesfromparasitismInparasitism,oneindividualisutilizedforthedevelopmentofmorethanoneparasite,Introduction,Predation(cont.)Predator-preyassociationsFigure10.2,AntipredatorAdaptations,AposematicorwarningcolorationAdvertisesanunpalatabletasteEx.BluejaysandmonarchbutterfliesCaterpillarobtainspoisonfrommilkweed,AntipredatorAdaptations,Ex.Bluejaysandmonarchbutterflies(cont.)BluejayssufferviolentvomitingfromingestingcaterpillarEx.TropicalfrogsToxicskinpoisonsFigure10.3a,AntipredatorAdaptations,CamouflageBlendingoforganismintobackgroundcolorGrasshoppers(Figure10.3b),AntipredatorAdaptations,Camouflage(cont.)StickinsectsmimictwigsandbranchesZebrastripes:blendintograssybackgroundMimicry,AntipredatorAdaptations,Mimicry(cont.)AnimalsthatmimicotheranimalsEx.SomehoverfliesmimicwaspsMimicryTypesofmimicryMllerianmimicryFritzMller,1879Unpalatablespeciesconvergetolookthesame,AntipredatorAdaptations,Unpalatablespeciesconvergetolookthesame(cont.)ReinforcebasicdistastefuldesignEx.WaspsandsomebutterfliesMimicryring:agroupofsympatricspecies,oftendifferenttaxa,shareacommonwarningpatternBatesianmimicryHenryBates,1862Mimicryofunpalatablespeciesbypalatablespecies,AntipredatorAdaptations,Batesianmimicry(cont.)Ex.hoverfliesresemblestingingbeesandwasps(Figure10.3d),AntipredatorAdaptations,DifficultydistinguishingtypeofmimicryMonarchbutterfliesandviceroybutterflies(Figures10.3d,e),AntipredatorAdaptations,DisplaysofintimidationEx.ToadsswallowairtomakethemselvesappearlargerEx.Frilledlizardsextendtheircollarstoproducethesameeffect(Figure10.3f),AntipredatorAdaptations,PolymorphismTwoormorediscreteformsinthesamepopulationColorpolymorphismPredatorhasapreference(usuallythemoreabundantform)Preycanproliferateintherarerform,AntipredatorAdaptations,Colorpolymorphism(cont.)Ex.leafhoppernymphs(orangeandblack)Ex.Peaaphids(redandgreen)ReflexiveselectionEveryindividualisslightlydifferentExamples:brittlestars,butterflies,moths,echinoderms,andgastropods,AntipredatorAdaptations,Reflexiveselection(cont.)ThwartpredatorslearningprocessesPreyphenologicallyseparatedfrompredatorEx.FruitbatsEitherdiurnalornocturnalOnlynocturnalinthepresenceofpredatorydiurnaleagles,AntipredatorAdaptations,ChemicaldefenseUsedtowardoffpredatorsEx.bombardierbeetlesPossessareservoirofhydroquinoneandhydrogenperioxideWhenthreatened,ejectchemicalsinto“explosionchamber”,AntipredatorAdaptations,Ex.bombardierbeetles(cont.)MixwithperoxidaseenzymeMixtureisviolentlysprayedatattackerMastingSynchronousproductionofmanyprogenybyallindividualsinpopulation,AntipredatorAdaptations,Masting(cont.)SatiatepredatorsAllowsforsomeprogenytosurviveCommontoseedherbivoryEx.17-yearand13-yearperiodicalcicadas,AntipredatorAdaptations,ComparisonofdefensemechanismsTable10.1,chemicaldefenseismostcommon,Predator-PreyModels,EffectsofpredatorsonpreyDependonsuchthingsaspreyandpredatordensities,andpredatorefficiencyGraphicalmethodtomonitorrelationship,Predator-PreyModels,Graphicalmethodtomonitorrelationship(cont.)PreyisoclineshavecharacteristichumpshapeFigure10.4,Preyincrease,i)Preyiscoline,K,N,N,N,N,K,1,1,1,2,2,2,ii)Predatoriscoline,Preydensity,Predatorincreases,Predatordecreases,Predatordensity,Predatordensity,Predator-PreyModels,Preyisoclineshavecharacteristichumpshape(cont.)Intheabsenceofpredators,preydensitywouldbeequaltothecarryingcapacity,K1Lowerlimit,individualsbecometooraretomeetforreproduction,Predator-PreyModels,Preyisoclineshavecharacteristichumpshape(cont.)Betweenthesetwovalues,preypopulationcaneitherincreaseordecreasedependingonpredatordensityAbovetheisocline,preypopulationsdecline,Predator-PreyModels,Preyisoclineshavecharacteristichumpshape(cont.)Belowtheisocline,preypopulationsincreasePredatorisoclinesThresholddensity,wherepredatorpopulationwillincreasePredatorpopulationcanincreasetocarryingcapacity,Predator-PreyModels,Predatorisoclines(cont.)MutualinterferenceorcompetitionbetweenpredatorsMorepreyrequiredforagivendensitypredatorPredatorisoclinesslopestowardtherightSuperimposepreyandpredatorisoclinesFigure10.5,Predator-PreyModels,Superimposepreyandpredatorisoclines(cont.)Onestablepointemerges:theintersectionofthelinesThreegeneralcasesInefficientpredatorsrequirehighdensitiesofprey(Figure10.5a),Dampedoscillations,Preyisocline,Predatorisocline,a),Predator-PreyModels,Threegeneralcases(cont.)Amoderatelyefficientpredatorleadstostableoscillationsofpredatorandpreypopulations(Figure10.5b),Stableoscillations,Populationdensity,Predatorequilibriumdensity,b),Predator-PreyModels,Threegeneralcases(cont.)Ahighlyefficientpredatorcanexploitapreynearlydowntoitslimitingrareness(Figure10.5c),Increasingoscillations,Predatordensity,c),Predator-PreyModels,AllbasedonhowefficientpredatorisShiftinisoclinesPreystarvation(shifttoleft)Foodenrichment(shifttoright)(Figure10.5d),K1,increasestoK1*withenrichment,Prey,Predator,Predatorisoclineremainsunchanged,“Theparadoxofenrichment”,Preyisoclinechanges,K1,K1*,d),Predator-PreyModels,Foodenrichment(shifttoright)(cont.)CarryingcapacitychangesPredatorisoclinechanges“paradoxenrichment”:Increasesinnutrientsorfooddestabilizesthesystem,Predator-PreyModels,FunctionalresponseHowanindividualpredatorrespondstopreydensitycanaffecthowpredatorsinteractwithprey(Figure10.6),I,II,III,Numberofpreyeatenperpredator,Preydensity,Predator-PreyModels,Functionalresponse(cont.)ThreetypesTypeI:IndividualsconsumemorepreyaspreydensityincreasesTypeII:Predatorscanbecomesatiatedandstopfeeding,orlimitedbyhandlingtime.,Predator-PreyModels,Threetypes(cont.)TypeIII:Feedingrateissimilartologisticcurve;lowatlowpreydensities,butincreasesquicklyathighdensitiesChangesinpreyconsumptionFunctionalresponsechanges(Figure10.7),Predator-PreyModels,Functionalresponsechanges(cont.)DictateshowindividualpredatorsrespondtopreypopulationNumericalresponsechangesGovernshowapredatorpopulationmigratesintoandoutofareasinresponsetopreydensities,FieldStudiesofPredator-PreyInteractions,FieldcomparisonstomodelsDopredatorscontrolpreypopulations?ImportanceofpredatorsincontrollingpreydensityKaibabdeerherdKaibabPlateau(NorthernArizona),FieldStudiesofPredator-PreyInteractions,Kaibabdeerherd(cont.)Declaredanationalparkaround1900AllbigpredatorswereremovedanddeerhuntingwasprohibitedEstimatesof10foldincreaseindeerpopulationReevaluatedbyGrahamCaughley(1970),FieldStudiesofPredator-PreyInteractions,ReevaluatedbyGrahamCaughley(1970)(cont.)Predatorcontrolhadsomeimpact;cessationofhuntingandremovalofcompetingsheepandcattlealsohadanimpactSerengetiplainsofeasternAfricaLargepredatorshavelittleeffectonlargemammalprey,FieldStudiesofPredator-PreyInteractions,SerengetiplainsofeasternAfrica(cont.)MostpreytakenareeitherinjuredorsenileContributelittletofuturegenerationsPreyaremigratoryMoosepopulationonMichigansIsleRoyale,FieldStudiesofPredator-PreyInteractions,MoosepopulationonMichigansIsleRoyale(cont.)Wolf-freeexistenceuntil1949.DurwoodAllen(1958)begantotrackwolfandmoosepopulationsTrendsinpopulations(Figure10.8),60,0,10,20,30,40,50,Wolves,1955,1960,1965,1970,1975,1980,1985,1990,1995,1997,Moose,Wolves,Year,200,400,600,800,1000,1200,1400,1600,1800,2000,2200,2400,2600,Moose,0,FieldStudiesofPredator-PreyInteractions,Trendsinpopulations(cont.)WolfpopulationPeakedat50in1980Severenosedivein1981Smallrecoveryinthelate1990sMoosepopulationIncreasedsteadilyinthe1960sand1970sDeclinedasthewolfpopulationincreaseduntil1981,FieldStudiesofPredator-PreyInteractions,Moosepopulation(cont.)Arecordpopulationof2500wasreachedin1995,whenthewolfpopulationwaslowGoodevidenceofpreypopulationcontrolbypredatorsConfoundedin1996whenthemoosepopulationcrashed-starvation,FieldStudiesofPredator-PreyInteractions,CanadalynxandsnowshoeharePopulationsshowdramaticcyclicoscillationsevery9to11years(Figure10.9),20,40,60,80,100,120,20,40,60,80,100,120,140,160,180,200,Abundanceoflynx,Abundanceofhares,Abundanceoflynx(x1000),Abundanceofhares(x1000),1850,1860,1870,1880,1890,1900,1910,1920,1930,1940,FieldStudiesofPredator-PreyInteractions,Canadalynxandsnowshoehare(cont.)Cyclehasexistedaslongasrecordshaveexisted(over200years)Anexampleofintrinsicallystablepredator-preyrelationship,IntroducedPredators,MethodfordeterminingtheeffectsofpredatorsDingopredationsonkangaroosinAustraliaDingoIntroducedspeciesLargestAustraliancarnivore,IntroducedPredators,Dingo(cont.)PredatorofimportedsheepEliminatedfromcertainareasSpectacularincreasesinnativespecies160foldincreaseinredkangaroosOver20foldincreaseinemus,IntroducedPredators,Dingo(cont.)EffectsonferalpigsShortageofyoungpigsConsiderableimpactonrecruitmentofpigs(Figure10.10),0,60,40,20,20,40,60,Ageclass(years),Males(%),Females(%),0,60,40,20,20,40,60,Males(%),Females(%),6+,5-6,4-5,3-4,2-3,1-2,0.5-1,.05,6+,5-6,4-5,3-4,2-3,1-2,0.5-1,.05,Ageclass(years),(a)Dingoespresent,(a)Dingoespresent,IntroducedPredators,EuropeanfoxesandferalcatsinAustraliaDamagedomesticlivestockEffectswhenremoved(Figure10.11),0,20,40,60,Predatorsshot,Noshooting,1981,1982,Meanno.ofrabbitsperkmoftransect,IntroducedPredators,LampreyandtheGreatLakesConstructionofWetlandCanalallowedlampreytoentertheGreatLakesDramaticreductioninlaketrout(Figure10.12),LakeHuron,Meanproduction,7,6,5,4,3,2,1,0,7,6,5,4,3,2,1,0,7,6,5,4,3,2,1,0,LakeMichigan,LakeSuperior,Meanproduction,Meanproduction,1930,1935,1940,1945,1950,1955,1960,Laketroutproduction(millionsperpound),IntroducedPredators,LampreyandtheGreatLakes(cont.)Troutrecoveredafterlampreypopulationwasreduced,FieldExperimentswithNaturalSystems,LionsinSouthAfricaKrugerNationalPark,1903LionsShotNumberoflargepreyincreasedShootingoflionsends,1960Wildebeastincreasesomuchthattheirnumbershadtobeculledfrom1965to1972,FieldExperimentswithNaturalSystems,Graypartridge,EuropeangamebirdFigure10.13,FieldExperimentswithNaturalSystems,Graypartridge,Europeangamebird(cont.)Over20millionshotinGreatBritaininthe1930sOnly3.8millionshotinthemid-1980sHighchickmortalityduetostarvation,FieldExperimentswithNaturalSystems,Only3.8millionshotinthemid-1980s(cont.)Reducedinsectsduetointroductionofherbicidesinthe1950swassuspectedHowever,smallerpopulationsinareaswheretherewasnocontrolofpredatorsbygamekeepers,FieldExperimentswithNaturalSystems,Only3.8millionshotinthemid-1980s(cont.)PredationcontrolincreasedThenumberofpartridgesthatbredsuccessfullyTheaveragesizeofthebroodsPartridgepopulationsby75%,FieldExperimentswithNaturalSystems,PredatorsandrodentsinFinlandLargescaleremovalofpredators,April1992and1995over2-3km2LargeincreaseinrodentpopulationbyJune(comparedtocontrolplots)(Figure10.14),April,June,April,June,3.5,3,2.5,2,1.5,1,0.5,0,3.5,3,2.5,2,1.5,1,0.5,0,Withoutpredators,Withpredators,Meannumberofrodentspersample,Meannumberofrodentspersample,AppliedEcology,Humansaspredators-whalingExploitationnecessaryIsharvestingatanylevelsustainable?HistoryofAntarcticwhalingFigure1,AppliedEcology,HistoryofAntarcticwhaling(cont.)1930s,bluewhalesprimarilyharvested1950s,bluewhalepopulationdepleted,replacedwithfinwhale1960s,finwhalepopulationcollapsed,AppliedEcology,HistoryofAntarcticwhaling(cont.)1960s,humpbackwhalepopulationcollapsedPriorto1958,SeiwhaleshardlyeverharvestedReductioninotherwhalesmadeSeiwhaleattractive,AppliedEcology,Priorto1958,Seiwhaleshardlyeverharvested(cont.)Peakharvestofabout20,000by1964-65CatchesdeclinedthereafterduetolimitationsTherelativelysmallminkewhaleWasignoredinthesouthernoceansuntil1971-72,AppliedEcology,Therelativelysmallminkewhale(cont.)Begantobetaken,andisnowthelargestcomponentofthesouthernbaleenwhalecatchWhalebanproposedin1985-86,tookeffectin1988,AppliedEcology,Iceland,Norway,andJapan,1994ArguedforresumptionoflimitedcommercialwhalingShouldwebancommercialwhaling?Whalepopulationsarerecovering,AppliedEcology,Whalepopulationsarerecovering(cont.)Ex.BluewhalepopulationshaveincreasedfourfoldEx.Californiagreywhaleshaverecoveredtoprewhalinglevels,Summary,PredationisastrongselectiveforceinnatureAposematiccolorationCamouflageBatesianandMullerianmimicryIntimidationdisplaysPolymorphisms,Summary,Predationisastrongselectiveforceinnature(cont.)ChemicaldefensesModelingpredator-preyinteractionsEvensimplepredator-preymodelsshow,Summary,Evensimplepredator-preymodelsshow(cont.)StablecyclesWildlyincreasingandunstableoscillationsDifficultyinpredictingormodelinghowpredatorsandpreyinteractMutualinterferencebetweenpredators,Summary,Difficultyinpredictingormodelinghowpredatorsandpreyinteract(cont.)ExistenceofspecificpredatorterritorysizesAbilityofpredatorstofeedonmorethanonetypeofprey,Summary,Large-scaleobservationssupportPredatorsonlytakeweakandsicklyindividualsPreypopulationsinfluencepredatornumbers,notviceversa,Summary,AccidentalordeliberateintroductionsofexoticpredatorsProfoundeffectsonnativepreypopulationsPredatorshaveimportantregulatoryeffectsonprey,Summary,Accidentalordeliberateintroductionsofexoticpredators(cont.)Maynotbeindicativeof“naturalsystems”,Summary,EvidencefromnaturalsystemsMoststudieshaveconcludedthatpredatorshaveasignificanteffectonprey,DiscussionQuestion#1,Shouldranchersbeconcernedaboutthereintroductionintotheirvicinityoflargepredators,likewolvesandpanthers?,DiscussionQuestion#2,Dosealions,otters,ordolphinsdecreasethestockoffishavailableforpeoplethatfish?,DiscussionQuestion#3,Wouldthenumberofdeeravailableforhuntersbethesameinthepresenceoflargepredators?,DiscussionQuestion#4,Whatdatawouldyouneedtocollecttoanswertheabove3questions?,DiscussionQuestion#5,Whatcantheeffectsofexoticpredatorstellusaboutthestrengthofpredation?Whatcanttheytellus?,DiscussionQuestion#6,Whichdoyouthinkmorelikely:thatpredatorscontrolpreypopulationsorthatpreycontrolpredatorpopulations?Wouldtheanswervaryaccordingtotheparticularsystem?Giveanexample.,DiscussionQuestion#7,WhatshortcomingsdoyouthinkRosenzweigandMacArthurspredatorandpreyisoclineshave?Whatwouldtheseshortcomingsmeanintermsofdetermininghowpredatorsandpreyinteract?,DiscussionQuestion#8,Agreatmanyfishstocksseemtohavebeenoverfished.Howdoyouthinkwecouldpreventoverfishing?Whatbiologicalinformationdoweneedtohave,andhowcanwegetitwhenwecantseethepopulationinquestion?,Predator-PreyModels,Preyisoclineshavecharacteristichumpshape(cont.)Belowtheisocline,preypopulationsincreasePredatorisoclinesThresholddensity,wherepredatorpopulationwillincreasePredatorpopulationcanincreasetocarryingcapacity,Predator-PreyModels,Predatorisoclines(cont.)MutualinterferenceorcompetitionbetweenpredatorsMorepreyrequiredforagivendensitypredatorPredatorisoclinesslopestowardtherightSuperimposepreyandpredatorisoclinesFigure10.5,Predator-PreyModels,Superimposepreyandpredatorisoclines(cont.)Onestablepointemerges:theintersectionofthelinesThreegeneralcasesInefficientpredatorsrequirehighdensitiesofprey(Figure10.5a),Dampedoscillations,Preyisocline,Predatorisocline,a),Predator-PreyModels,Threegeneralcases(cont.)Amoderatelyefficientpredatorleadstostableoscillationsofpredatorandpreypopulations(Figure10.5b),Stableoscillations,Populationdensity,Predatorequilibriumdensity,b),Predator-PreyModels,Threegeneralcases(cont.)Ahighlyefficientpredatorcanexploitapreynearlydowntoitslimitingrareness(Figure10.5c),Increasingoscillations,Predatordensity,c),Predator-PreyModels,AllbasedonhowefficientpredatorisShiftinisoclinesPreystarvation(shifttoleft)Foodenrichment(shifttoright)(Figure10.5d),K1,increasestoK1*withenrichment,Prey,Predator,Predatorisoclineremainsunchanged,“Theparadoxofenrichment”,Preyisoclinechanges,K1,K1*,d),Predator-PreyModels,Foodenrichment(shifttoright)(cont.)CarryingcapacitychangesPredatorisoclinechanges“paradoxenrichment”:Increasesinnutrientsorfooddestabilizesthesystem,Predator-PreyModels,FunctionalresponseHowanindividualpredatorrespondstopreydensitycanaffecthowpredatorsinteractwithprey(Figure10.6),I,II,III,Numberofpreyeatenperpredator,Preydensity,Predator-PreyModels,Functionalresponse(cont.)ThreetypesTypeI:IndividualsconsumemorepreyaspreydensityincreasesTypeII:Predatorscanbecomesatiatedandstopfeeding,orlimitedbyhandlingtime.,Predator-PreyModels,Threetypes(cont.)TypeIII:Feedingrateissimilartologisticcurve;lowatlowpreydensities,butincreasesquicklyathighdensitiesChangesinpreyconsumptionFunctionalresponsechanges(Figure10.7),Preda
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