族群生态学Populationecology群体群落课件

Chapter6

PopulationGrowth106.*生態學的分科：

以生物組織水準來分個體生態學Autecology種群（族群）生態學Populationecology群體（群落）生態學Synecology: communityecology生態系統生態學Ecosystemecology206.*OutlineTabulatingchangesinpopulationagestructurethroughtimeTime-specificlifetablesAge-specificlifetables306.*OutlineFecundityschedulesandfemalefecundity,andestimatingfuturepopulationgrowthPopulationgrowthmodelsDeterministicmodelsGeometricmodels406.*OutlinePopulationgrowthmodels(cont.).LogisticmodelsStochasticmodels506.*DemographyandPopulationGrowthDemography族群統計學、人口統計學ThequantitativedescriptionofapopulationPopulation族群Agroupofconspecificsinhabitsaspecificplaceataspecifictime.

.Thedemographicandgeneticpopulationsarenotnecessarilythesame..Demographicdistinctionandgeneticdifferencearenotnecessarilycorresponding.606.*DemographyandPopulationGrowth.Populationcharacteristics.Quantitativeparameters.Populationdifference:density,agedistribution.Populationgrowthandquantitativemethods.Populationdeterminationfactors.Offspringproductionandenergyinvest.Populationgeneticsvariation.Intrapopulationbehavior706.*Populationdensity族群密度.abundant豐富的.common普遍.rare稀有.endangered瀕危(絕)的.extinct絕滅的eg.Zaccopachycephalus

粗首獵(溪哥)endemictoTaiwan,butcommoninwestcoasts806.*Samplingmethods:.Traps陷阱.Fecalpellets糞便.Vacuolizationfrequencies鳴叫頻度.peltrecords生皮記錄.Catchperuniteffort(CPUE)單位努力漁獲量.Percentagegroundcover遮蔽度.Frequencyofabundancealongtransectsorinquadrantsofknownarea.穿越線或四分區豐度。.Feedingdamage食害程度。.Roadsidespottinginastandarddistance單位長度內視察的記錄。906.*Estimatesofabsolutedensity-1.Proportionofpopulationmarked=(no.ofanimalsmarkedatt1)/(totalno,ofanimalsinpopulation).Proportionofsamplemarked=(no.ofmarkedanimalscapturedatt2)/(totalno.ofanimalscapturedatt2)1006.*Estimatesofabsolutedensity-2Ifrandomsampling,

Proportionofpopulationmarked=proportionofsamplemarkedt1標示個體/族群數量﹦t2再捕獲標示個體數/t2捕獲數（已知數）（未知數） （已知數） （已知數）

Assumptions:1.

標記不會對於個體有增加死亡率的危險2.

標記不會影響再捕捉的機率（記憶與學習）3.

實驗期間族群內個體沒有移入或移出的問題4.

實驗期間沒有死亡或新生的變化1106.*櫻花鉤吻鮭1206.*SalmonWatch—SkinDiving1306.*SalmonWatch—SkinDivingandCounting1406.*七家灣溪流域櫻花鉤吻鮭歷年族群變化圖重大天災以紅色圖說標於圖中歷年最高數量3428尾2000秋繁殖季節遭逢象神颱風1994年以來最低族群數量346尾1506.*頭前溪毛蟹1606.*Dispersion分散情形.randomdistribution:闊葉林中的樹.aggregated:草地上的韓國薊.hyperdispersed,regularin dispersion:繁殖區的海鳥1706.*雪山冷杉林1806.*Statisticalmethods:-1.PoissondistributionPx=axe-a/x! P=Poissonprobability x=No.ofoccurrences a=Themeannumberofoccurrences e=Thebaseofthenaturallog1906.*蘭嶼熱帶森林2006.*Statisticalmethods:-2方差/平均數比率指標Ifs2/x

(mean/variance)>1

hyperdispersionIfs2/x

(mean/variance)<1

aggregationSignificanttest

(n-1)s2/x

x:meanS2:variance2106.*LifeTables壽命表是一種記載和描述出生後的一定數量的個體，隨著生命過程而損失，減少的規律的一種方法，是描述種群的死亡與存活過程的一種有效手段。2206.*LifeTables壽命表1.agespecific(cohortanalysis)群對分析,年齡群特性。Short-livedanimal：

2.time-specific(static)靜態分析,時間特性。long-livedanimals.

2306.*age-specificlifetable

特定年齡壽命表=dynamiclifetable=horizontallifetable以一特定世代群體的命運為根據，跟蹤該世的整個生活史，每隔一定時間統計存活數與死亡數。

.優點.可反映出與密度相關的死亡率與繁殖率變化。.分析種群數量變動與調節的機制。

缺點.不易掌握全世代的生物資料。2406.*Timespecificlifetable

特定時間壽命表=StationaryorVerticallifetable由不同年齡組組成的群體及估算的各年齡組的死亡率為根據。

優點.可在短時間內獲得所需資料缺點.不同世代會有不同歷史背景.推理不易來自隨機的取樣.種群結構不易穩定

2506.*Compositelifetable

綜合壽命表取不同來源的資料平均值作壽命表

(B)birth種群大小 death(D)

dn=B-DNetreproductiverate:RoRo=lxmxln=survivingproportionmx=age-specificfertility2606.*四種族群年齡結構的組成圖2706.*已開發和開發中國家在1995～2025年之間的年齡結構2806.*三種不同的存活曲線模型2906.*TypeITypeIITypeIIIManymammalsManybirds,smallmammals,lizards,turtlesManyinvertebratesAgeNumberofsurvivors(n)(logscale)x10001001010.13006.*DeterministicModels

宿命論模式DiscreteGenerations不連續世代

GeometricCarryingCapacitylogistick-n尚未利用於種群增長的潛力

3106.*StochasticModels推計學模式DiscreteGeneration

OverlappingGenerations3206.*細菌的對數生長3306.*海豹族群的成長模式3406.*族群成長率與族群數量增加之間的關係3506.*邏輯模型所預測的族群成長情形3606.*指數成長和對數成長的比較3706.*LifeTablesDateonnumbersofindividualsateachageConstructlifetablesDemonstratetheagestructureofapopulationLifetableconstruction–demography3806.*LifeTablesLifetableconstruction–demography(cont.).HowapopulationwillgrowConstructionofdataFollowacohortfrombirthstodeaths(agespecificlifetable)3906.*LifeTablesTime-specificlifetableSnapshot–agestructureatasinglepointintime(time-specificlifetable)Usefulinexamininglong-livedanimalsEx.DallMountainSheep(Figure6.1andTable6.1)4006.*00.511.522.533.5Age(years)n(logscale)x1011423456789101112134106.*4206.*LifeTablesUsefulparametersinthelifetables(Deevey1947)x=ageclassorintervalnx=numberofsurvivorsatbeginningofageintervalx.dx=numberoforganismsdyingbetweenageintervals=nx–nx=1lx=proportionoforganismssurvivingtothebeginningofageintervalx=ns/n04306.*LifeTablesUsefulparametersinthelifetables(Deevey1947)(cont.).qx=rateofmortalitybetweenageintervals=dx/nsex=themeanexpectationoflifefororganismsaliveatthebeginningofagexLx=averagenumberaliveduringanageclass=(ns+ns+1)/2Tx=intermediatestepindetermininglifeexpectancy=SLx4406.*LifeTablesex=themeanexpectationoflifefororganismsalive(cont.).Ex=Tx/nxAssumptionsthatlimittheaccuracyoftime-specificlifetablesEqualnumberofoffspringareborneachyearFavorableclimateforbreeding?Aneedforanindependentmethodforestimatingbirthratesofeachageclass4506.*LifeTablesAssumptionsthatlimittheaccuracyoftime-specificlifetables(cont.).Asaresult,age-specificlifetablesaretypicallyreportedAge-specificlifetablesNeededforshort-livedorganismsTime-specificlifetablesbiasedtowardthestagecommonatthemoment4606.*LifeTablesFollowsonecohortorgenerationPopulationcensusesmustbefrequentandconductedoveralimitedtimeEx.Table6.2andFigure6.34706.*4806.*00.511.522.533.5n(logscale)

xAge(years)12763454906.*LifeTablesGeneraltypesofsurvivorshipcurves(Figure6.4)TypeIMostindividualsarelostwhentheyareolderVertebratesororganismsthatexhibitparentalcareandprotecttheiryoungSmalldipatyoungageduetopredators5006.*LifeTablesTypeIIAlmostlinearrateoflossManybirdsandsomeinvertebratesTypeIIILargefractionarelostinthejuvenilestagesInvertebrates,manyplants,andmarineinvertebratesthatdonotexhibitparentalcare5106.*LifeTablesTypeIII(cont.).LargelossesduetopredatorsComparisonintheaccuracyoflifetablesFigure6.55206.*ReproductiveRateFecundityAge-specificbirthratesNumberoffemaleoffspringproducedbyeachbreedingfemaleFecundityschedulesFecundityinformationinlifetable5306.*ReproductiveRateDescribereproductiveoutputandsurvivorshipofbreedingindividualsFecundityschedules(cont.).Ex.Table6.35406.*5506.*ReproductiveRateTablecomponentslx=survivorship(numberoffemalessurvivingineachageclassmx=age-specificfecundityRo=population’snetreproductiverate=Slxmx

5606.*ReproductiveRateRx=population’snetreproductiverate=SlxmxRo=1;populationisstationaryRo>1;populationisincreasingRo<1;populationisdecreasingTable6.35706.*5806.*ReproductiveRateVariationinformulaforplantsAge-specificfecundity(m)iscalculateddifferentlyFx=totalnumberofeggs,seeds,oryoungdepositednx=totalnumberofreproducingindividualsmx=Fx/nxFigure6.5x5906.*6006.*ReproductiveRateVariationinformulaforplants(cont.).Table6.46106.*6206.*ReproductiveRateVariationinformulaforplants(cont.).Figure6.66306.*DeterministicModels:GeometricGrowthPredictingpopulationgrowthNeedtoknow;RoInitialpopulationsizePopulationsizeattimetPopulationsizeoffemalesatnextgeneration=Nt+1=RoNt6406.*DeterministicModels:GeometricGrowthPopulationsize(cont.).Ro=netreproductiverateNt=populationsizeoffemalesatthisgeneration6506.*DeterministicModels:GeometricGrowthDependencyofRoRo<1;populationbecomesextinctRo=1;populationremainsconstantPopulationisatequilibriumNochangeindensity6606.*DeterministicModels:GeometricGrowthDependencyofRRo>1;populationincreasesEvenafractionaboveone,populationwillincreaserapidlyo6706.*DeterministicModels:GeometricGrowthRo>1;populationincreases(cont.).Characteristic“J”shapedcurveGeometricgrowthFigure6.76806.*100200300400500Populationinsize(N)0Generations30R=1.200R=1.150R=1.100R=1.0501020N+1=RNt0t6906.*DeterministicModels:GeometricGrowthRo>1;populationincreases(cont.).Something(e.g.,resources)willeventuallylimitgrowthPopulationcrashFigure6.8a7006.*19101920193019401950Numberofreindeer2000150010005000Year7106.*DeterministicModels:GeometricGrowthRo>1;populationincreases(cont.).Figure6.8b7206.*DeterministicModels:GeometricGrowthRo>1;populationincreases(cont.).Figure6.8c7306.*DeterministicModels:GeometricGrowthHumanpopulationgrowthPriortoagricultureanddomesticationofanimals(~10,000B.C.)Averageannualrateofgrowth:~0.0001%7406.*DeterministicModels:GeometricGrowthAftertheestablishmentofagriculture300millionpeopleby1A.D.800millionby1750Averageannualrateofgrowth:~0.1%7506.*DeterministicModels:GeometricGrowthPeriodofrapidpopulationgrowthBegan1750From1750to1900Averageannualrateofgrowth:~0.5%From1900to1950Averageannualrateofgrowth:~0.8%From1950to2000Averageannualrateofgrowth:~1.7%7606.*DeterministicModels:GeometricGrowthPeriodofrapidpopulationgrowthReasonsforrapidgrowthAdvancesinmedicineAdvancesinnutritionTrendsingrowth(Figure6.9)7706.*1830193019601975198719982009202020332046210001234567891011131214Billionsofpeople2-5millionYearsago7,000BC6,000BC5,000BC4,000BC3,000BC2,000BC1,000BC1AD1,000AD2,000AD3,000ADYear4,000AD7806.*DeterministicModels:GeometricGrowthHumanpopulationstatisticsPopulationisincreasingatarateof3peopleeverysecondCurrentpopulation:over6billionUNpredictspopulationwillstabilizeat11.5billionby2150DevelopedcountriesAverageannualrateofgrowthfrom1960-1965:1.19%7906.*DeterministicModels:GeometricGrowthHumanpopulationstatistics(cont.).Averageannualrateofgrowthfrom1990-1995:0.48%DevelopingcountriesAverageannualrateofgrowthfrom1960-1965:2.35%8006.*DeterministicModels:GeometricGrowthHumanpopulationstatistics(cont.).DevelopingcountriesAverageannualrateofgrowthfrom1990-1995:2.38%8106.*DeterministicModels:GeometricGrowthFertilityratesAveragenumberoflivebirthstypicallybornebyawomanduringherlifetime(Table6.6)8206.*8306.*DeterministicModels:GeometricGrowthFertilityrates(cont.).Theoreticreplacementrate:2.0Actualreplacementrate:2.1Declineinfertilityrate1960-1965:~5.01990:3.38406.*DeterministicModels:GeometricGrowthOverlappinggenerationsManyspeciesinwarmclimatesreproducecontinuallyandgenerationsoverlap.8506.*DeterministicModels:GeometricGrowthOverlappinggenerations(cont.).RateofincreaseisdescribedbyadifferentialequationdN/dt=rN=(b–d)NN=populationsizet=timer=percapitarateofpopulationgrowth8606.*DeterministicModels:GeometricGrowthOverlappinggenerations(cont.).RateofincreaseisdescribedbyadifferentialequationdN/dt=rN=(b–d)Nb=instantaneousbirthrated=instantaneousdeathratedN=therateofchangeinnumbers8706.*DeterministicModels:GeometricGrowthRateofincreaseisdescribedbyadifferentialequation(cont.).dN/dt=therateofpopulationincreaseProducesa“J”shapedcurvePlotwiththenaturallogarithm,producesastraightline(Figure6.8)8806.*01234520406080100r=0.02r=0.01r=0(equilibrium)Time(t)In(N)8906.*DeterministicModels:GeometricGrowthrisanalogoustoRoInastablepopulationR=(lnRo)/TcLogisticGrowthOccursinpopulationswhereresourcesareorcanbelimitingLogisticgrowthequationsTcgenerationtime9006.*DeterministicModels:GeometricGrowthLogisticgrowthequationsdN/dt=rN[(K-N)/K];ordN/dt==rN[1-(N/K)]dN/dt=Rateofpopulationchanger=percapitarateofpopulationgrowthN=populationsizeK=carryingcapacity9106.*DeterministicModels:GeometricGrowth(K-N)/K=unusedresourcesremainingS-ShapedCurve:Figure6.11AsNgetslarger,theamountofresourcesremaininggetssmallerWhenN=K,zerogrowthwilloccur9206.*PopulationsizeKTime9306.*DeterministicModels:GeometricGrowthLogisticgrowthassumptionsRelationbetweendensityandrateofincreaseislinearEffectofdensityonrateofincreaseisinstantaneousEnvironment(andthusK)isconstantAllindividualsreproduceequallyNoimmigrationandemigration9406.*DeterministicModels:GeometricGrowthTestingassumptionsEarlylaboratoryculturesPearl1927Figure6.129506.*150300450600750AmountofyeastK=66502468101214161820Time(hrs)9606.*DeterministicModels:GeometricGrowthComplexstudiesandtemporaleffects(cont.).Figure6.139706.*200400600800TimeNNumberper12gramsofwheatLogisticcurvepredictedbytheoryTime(weeks)50100180CallandraoryzaeRhizoperthadominica9806.*DeterministicModels:GeometricGrowthDifficultyinmeetingassumptionsinnatureEachindividualaddedtothepopulationprobablydoesnotcauseanincrementaldecreasetorTimelags,especiallywithspecieswithcomplexlifecyclesKmayvaryseasonallyand/orwithclimate9906.*DeterministicModels:GeometricGrowthDifficultyinmeetingassumptionsinnature(cont.).OftenafewindividualscommandmanymatingsFewbarrierstopreventdispersalEffectoftimelagsRobertMay(1976)10006.*DeterministicModels:GeometricGrowthEffectoftimelags(cont.).IncorporatedtimelagsintologisticequationdN/dt=rN[1-(Nt-t/K)]dN/dt=Rateofpopulationchanger=percapitarateofpopulationgrowthN=populationsizeK=carryingcapacity10106.*DeterministicModels:GeometricGrowthdN/dt=rN[1-(Nt-t/K)](cont.).Nt-t=timelagbetweenthechangeinpopulationsizeanditseffectonpopulationgrowth,thenthepopulationgrowthattimetiscontrolledbyitssizeatsometimeinthepast,t-tNt-t=populationsizeinthepastEx.r=1.1,K=1000andN=900Notimelag,newpopulationsize10206.*DeterministicModels:GeometricGrowthNotimelag,newpopulationsize(cont.).dN/dt=1.1x900(1–900/1000)=99Newpopulationsize=900+99=99910306.*DeterministicModels:GeometricGrowthNotimelag,newpopulationsize(cont.).Withtimelag,whereapopulationis900,althoughtheeffectsofcrowdingarebeingfeltasthoughthepopulationwas800StillbelowKdN/dt=1.1x900(1–800/1000)=198Newpopulationsize=900+198=1098PossibleforapopulationtoexceedK10406.*DeterministicModels:GeometricGrowthEffectofresponsetimei.Inverselyproportionaltor=(1/r)Ratiooftimelag(t)toresponsetime(1/r)orrtcontrolspopulationgrowthrtissmall(<0.368)Populationincreasessmoothlytocarryingcapacity(Figure6.14)10506.*KTime(t)SmoothresponseKTime(t)Numberofindividuals(N)KTime(t)DampedoscillationsperiodamplitudeStablelimitcycler

`small(<0.368)r

t

medium(>0.368,<1.57)rtlarge(>1.75)Numberofindividuals(N)Numberofindividuals(N)10606.*DeterministicModels:GeometricGrowth

rtislarge(>1.57)PopulationentersintoastableoscillationcalledalimitcycleRisingandfallingaroundKNeverreachingequilibriumrtisintermediate(>0.368and<1.57)PopulationsundergooscillationsthatdampenwithtimeuntilKisreached

10706.*DeterministicModels:GeometricGrowthSpecieswithdiscretegenerationsLogisticequationNt+1=Nt+rNt[1–(Nt/K)]Indiscretegenerations,thetimelagis1.0rissmall(2.0)PopulationgenerallyreachesKsmoothlyrisbetween2.0and2.449Populationentersastabletwo-pointlimitcyclewithsharppeaksandvalleys10806.*DeterministicModels:GeometricGrowthSpecieswithdiscretegenerationsLogisticequationrisbetween2.449and2.570

MorecomplexlimitcyclesFigure6.1510906.*NNNtttrsmall(2.000–2.499)rmedium(2.499–2.570)rlarge(>2.570)11006.*DeterministicModels:GeometricGrowthSpecieswithdiscretegenerationsLogisticequation(cont.).rislargerthan2.57LimitcyclesbreakdownPopulationgrowsinacomplex,non-repeatingpatterns,knowas‘chaos’11106.*StochasticModelsModelsarebasedonprobabilitytheoryFigure6.1611206.*00.100.200.30Proportionofobservations68101214Populationsize11306.*StochasticModelsStochasticmodelsofgeometricgrowthdN/dt=rN=(b–d)NIfb=0.5,d=0,andN0=10,integralformofequation11406.*StochasticModelsStochasticmodelsofintegralformofequationNt=N0ertSofortheaboveexample,Nt=10x1.649=16.4911506.*StochasticModelsStochasticmodelsofgeometricgrowth(cont.).PathofpopulationgrowthFigure6.1711606.*PopulationdensityTimeExtinctionPossiblestochasticpath11706.*StochasticModelsProbabilityofextinction=(d/b)N0ThelargertheinitialpopulationsizeThegreaterthevalueofb–dThemoreresistantapopulationistoextinction11806.*StochasticModelsIntroducebiologicalvariationintocalculationsofpopulationgrowthMorerepresentativeofnatureMorecomplicatedmathematics11906.*AppliedEcology1992JohnsHopkinsstudyDevelopedcountries70%ofcouplesusecontraceptivesDevelopingcountries~45%ofcouplesusecontraceptivesAfrica,14%12006.*AppliedEcologyDevelopingcountries(cont.).Asia,50%LatinAmerica,57%12106.*AppliedEcology1992JohnsHopkinsstudy(cont.). China1950sand1960sFertilitywassixchildrenperwoman1970sGovernmentplanningandincentivestoreducepopulationgrowth12206.*AppliedEcologyChina(cont.).199075%usebirthcontrolFertilityratedroppedto2.2Othergovernments1976,only97governmentssupportedfamilyplanning12306.*AppliedEcologyOthergovernments(cont.).1988,125governmentssupportedfamilyplanningAsof1989,in31countries,coupleshavenoaccesstofamilyplanning12406.*AppliedEcologyWomenWomenindevelopingcountrieswantfewerchildrenInvirtuallyeverycountryoutsideofSaharanAfrica,thedesiredsnumberofchildrenisbelow312506.*AppliedEcologyCountriesconcernedaboutlowgrowthratesSomeWesternEuropeancountriesandotherdevelopedcountriesTotalfertilityhasdroppedbelowthereplacementlevelof2.112606.*AppliedEcologyCountriesconcernedaboutlowgrowthrates(cont.).ReducedpopulationsconcernsAffectpoliticalstrengthEconomicstructure12706.*

Summary

Lifetablesprovideinformationonhowpopulationsarest