第二章 生命表及其分析方法2016

昆虫生态学徐学62815981xnxu@

第二章生命表及其分析方法第三章昆虫单种群增长模型

第四章种间关系

一．生命表的概念

二．生命表的类型及参数1时间特征生命表及参数估计生殖力表的构建内禀增长率的含义及估计方法2

年龄特征生命表种群存活曲线种群趋势指数关键因子分析

四.

昆虫种群的r对策与Ｋ对策第二章

生命表及其分析方法

三．生命表的作用一．

生命表的概念昆虫生命表(lifetable)：采用田间系统调查或一定条件下的室内实验，系统观察并以一定的表格形式，记录某一种群在各个年龄或发育阶段的死亡数量、死亡原因和成虫阶段的繁殖数量。二．生命表的类型及参数

时间特征生命表(time-specificlifetable)

年龄特征生命表(age-specificlifetable)

1时间特征生命表(time-specificlifetable)及参数估计

是在年龄组配稳定的前提下，以特定时间为间隔（如天、周、月等）系统调查记载在时刻x

开始时的数量（或存活率）和x期间死亡数量。也包括各时间间隔内每一雌体的平均产雌数量。内禀增长率rm（intrinsicrateofincrease

)周限增长率λ（finiterateofincrease)净增殖率R0

（netreproductionrate)世代重叠的种群室内实验种群不能分析种群死亡的主要原因或关键因子生命期望表生殖力表生殖力表的构建桔全爪螨(Panonychuscitri)在25℃下的生殖力表6.66820.47631.10.43314132.914712.7022Σ9.44580.72661.40.519130000.011269.79200.81601.50.544120000.0372511.08911.00811.70.593110.88800.03701.00.0372410.19201.01921.60.637100.85100.03700.50.074239.00451.00051.50.66792.25060.10231.10.093228.22001.02751.50.68583.25710.15511.10.141214.71240.67320.90.74878.49200.42462.20.193208.11921.35321.70.79665.65440.26971.20.248195.97001.19401.50.79655.00400.27801.00.278183.26000.815010.81546.06560.35681.10.3261700.85936.65280.41581.10.3781600.91527.32600.48841.20.4071501.0001lxmxxlxmx每雌产雌率mx存活率lx年龄xlxmxxlxmx每雌产雌数mx存活率lx年龄x(天)庭园发丽金龟（Phylloperthahorticola）的生殖力表

（Laughlin1965)X（周）lxmxlxmxlxmxx01.0-幼期490.49-500.45-510.421.00.4221.42520.316.92.13110.76530.057.50.3820.14540.010.90.010.54∑2.94152.86内禀增长率的含义及估计方法粗略估计：rm=lnR0/T精确计算：内禀增长率rm

：对于在恒定环境下增长的种群，如果外界的食物、空间等条件对种群不加限制，那么种群将永远保持最适容度在该无限的空间中增长，同时可以期望，对于具有稳定年龄组配的种群，其最大出生率（b）和最小死亡率（d）将趋向常值，这个常值即种群的内禀增长率。Birch&Watson林昌善rm

的粗略估计t0:N=N0t:N=Ntt=T

R0：净增殖率每一世代种群的增殖倍数。

T：世代平均周期从亲代出生到子代出生的平均周期when

R0

=1,rm

=0,stablepopulationR0

<1,rm

<0,shrinkingR0

>1,rm

>0,growingrememberthatrm=b-dwhereb=births/unittimeandd=deaths/unittime.Sounitsofrmareindividualsproducedperunittime.rm

的粗略估计庭园发丽金龟桔全爪螨∑lxmx(R0)2.9412.7022∑lx

mxx152.86132.9147T51.993210.4639rm0.020740.2431rm

的精确计算

如计算值小于1，应增加-rm

值（rm绝对值减小）再算，如计算值大于1，应减小-rm

值（rm的绝对值增加）再算，直至等号右边为1为止，此时的rm值为精确计算值。xNxlxmxlxmxxlxmxe-rxe-rxlxmx050010001014000.80870.77922000.40370.2843500.10150.01240000.00.00.0560∑=1.075

∑e-rmxlxmx=1.075,sofirstestimateofrm

=0.72istoosmall.(Wearesummingtermsraisedtoanegativeexponentofr,so↑rmwill↓sum).Adjustestimateofrmandtryagain.Asaguess,tryrm=0.75xNxlxmxlxmxxlxmxe-rxe-rxlxmx050010001014000.80720.75522000.40230.2683500.10050.01140000.00.00.0500∑=1.034∑

e-rmxlxmx

=1.034,soestimateofrm

(0.75)isstilltoosmall.Increaseitandtryagain.Byrepeatingthisprocess,willhomeinonrm=0.776,whichgives∑

e-rmxlxmx

=1xNxlxmxlxmxxlxmxe-rxe-rxlxmx050010001014000.80600.73622000.40120.2543500.10970.01040000.00.00.0450∑=1.000rm=0.776

周限增长率λ

：表示种群在经过一个单位时间后为原数量的倍数。种群加倍时间t

：温度对生命表参数的影响温度对两种天敌生命表参数的影响DietEffectsonIntrinsicRateofIncreaseandRearingofProprioseiopsisasetus(Acari:Phytoseiidae)Authors:Emmert,C.J.;

Mizell,R.F.;

Andersen,P.C.;

Frank,J.H.;

Stimac,J.L.Source:

AnnalsoftheEntomologicalSocietyofAmerica,Volume101,

Number6,November2008,pp.1033-1040(8)Abstract:Theintrinsicrateofincrease(rm)wasestimatedforProprioseiopsisasetusMuma(Acari:Phytoseiidae)onpollenandtwopreyspeciesat35°C.Cattail,TyphalatifoliaL.,pollenandFrankliniellaoccidentalis(Pergande)wereequallyfavorableasfoodandsuperiortoTetranychusurticaeKoch.MorefemaleswereproducedinashortertimewhenP.asetuswasfedF.occidentalisorpollen;however,survivorshipwaslongerwhenthepredatorwasfedT.urticae.OnanF.occidentalisdiet,calculatedrmvaluesrangedfrom0.2789to0.2925.ThenetreproductiveratewasloweronadietofT.urticae(6.48)thanonpollen(9.76)oronF.occidentalis(21.39).ThefiniterateofincreasewasloweronT.urticae(1.1868)thanonpollen(1.3274)orF.occidentalis(1.3398).Themeangenerationtimewassimilaronthethreediets.Thatthedatareportedherewerecollectedat35°Cmakesitremarkable.P.asetusmayhavegreatpotentialasabiologicalcontrolagentofarthropods,especiallythripsunderclimaticconditionsofhightemperaturessuchasinprotectedculture.宽叶香蒲花粉西方花蓟马二斑叶螨rm0.28320.2789-0.29250.17130.2425R09.7621.396.48λ1.32741.33981.1868

一种植绥螨(Proprioseiopsisasetus)在35℃条件下取食不同食物后的生命表参数Emmertetal.2008.AnnalsoftheEntomologicalSocietyofAmerica,101(6),1033-1040不同的食物对生命表参数的影响不同食物对生命表参数的影响Parameters瘿螨Aceriadioscoridis叶螨Tetranychusurticae(nymphs)家蝇Muscadomestica(eggs)根结线虫Meloidoyneincognita(eggs)Netreproductionrate(R0)44.7322.5426.835.82Meangenerationtime(T)17.9223.4422.6226.00Intrinsicrateofincrease(rm)0.2120.1380.1450.065Finiterateofincrease(erm)61.07Timeto50%mortality(days)32454248No.observation(female)27252112Sexratio(female/total)38/5238/5431/4852/90Sexratio(female:male)2.71:12.36:11.82:11.37:1四种不同食料(A.dioscoridis,T.urticae,M.domesticaandM.incognita)饲养一种毛绥螨(L.athiasae)对其生命表参数的影响(27℃)(Abou-Awadetal.,2001)蔬菜R0rmλTGI黄瓜21.200.2081.23114.9616.96甘蓝21.280.1841.20217.4315.65豆11.020.1641.17815.629.49西红柿6.820.1001.10522.487.07辣椒1.340.0171.01716.191.13GI,生长指数,成虫羽化率与非成熟期(仅为幼虫期和蛹期)平均时间的比率(Sétamou,etal.,1999)西方花蓟马取食五种不同蔬菜植物的生命表参数及生长指数(Zhangetal.,2007)不同作物对生命表参数的影响GeraldoJoséNascimentodeVasconcelos·GilbertoJosédeMoraes·ÍtaloDelaliberaJúnior·MarkusKnapp,2008.LifehistoryofthepredatorymitePhytoseiulusfragariaeonTetranychusevansiandTetranychusurticae(Acari:Phytoseiidae,Tetranychidae)atfivetemperatures.ExpApplAcarol(2008)44:27–36AbstractTetranychusevansiBakerandPritchardandTetranychusurticaeKoch(Acari:Tetranychidae)areimportantpestsofSolanaceaeinmanycountries.SeveralstudieshavedemonstratedthatT.urticaeisanacceptablepreytomanypredatorymites,althoughthesuitabilityofthispreydependsonthehostplant.T.evansi,hasbeenshowntobeanunfavorablepreytomostpredatorymitesthathavebeentestedagainstit.ThepredatorPhytoseiulusfragariaeDenmarkandSchicha(Acari:Phytoseiidae)hasbeenfoundinassociationwiththetwospeciesinBrazil.TheobjectiveofthisworkwastocomparebiologicalparametersofP.fragariaeonT.evansiandonT.urticaeasprey.Thestudywasconductedunderlaboratoryconditionsat10,15,20,25and30°C.Atalltemperatures,survivorshipwasloweronT.evansithanonT.urticae.Nopredatorreachedadulthoodat10°Contheformerspecies;evenonthelatterspecies,onlyabout36%ofthepredatorsreachedadulthoodat10°C.Forbothprey,ingeneral,durationofeachlifestagewasshorter,totalfecunditywaslowerandintrinsicrateofpopulationincrease(rm)washigherwithincreasingtemperatures.TheslowerrateofdevelopmentofP.fragariaeonT.evansiresultedinaslightlyhigherthermalrequirement(103.9degree-days)onthatpreythanonT.urticae(97.1degree-days).Thevaluesofnetreproductionrate(R0),intrinsicrateofincrease(rm)andfiniterateofincrease(λ)weresignificantlyhigheronT.urticae,indicatingfasterpopulationincreaseofthepredatoronthispreyspecies.Thehighestvalueofrmofthepredatorwas0.154and0.337femaleperfemaleperdayonT.evansiandonT.urticae,respectively.TheresultssuggestedthatP.fragariaecannotbeconsideredagoodpredatorofT.evansi.ParameterpreyTemperature(°C)A15202530RegressionequationBR2(%)nC伊氏叶螨25283226-二斑叶螨36363542-T伊氏叶螨35.8±12.1b21.4±7.1b17.4±4.9a13.7±4.5aò=104.535-6.221T+0.107T293.8二斑叶螨40.6±11.1a28.1±7.7a15.6±4.3a12.5±3.2aò=112.410-6.166T+0.094T297.3R0伊氏叶螨5.5±1.8b6.1±1.8b8.5±2.5b7.3±2.3bò=-5.210+0.966T-0.018T275.7二斑叶螨24.5±6.7a39.7±10.9a29.7±8.2a34.1±8.6aò=-27.760+5.236T-0.108T277.6rm伊氏叶螨0.048±0.016b0.084±0.028b0.123±0.041b0.145±0.052bò=-0.049+0.007T98.4二斑叶螨0.023±0.014a0.161±0.044a0.273±0.076a0.337±0.085aò=-0.177+0.017T99.5λ伊氏叶螨1.049±0.358b1.088±0.347b1.131±0.3291.156±0.388bò=0.942+0.007T99.5二斑叶螨1.082±0.296a1.140±0.312a1.242±0.344a1.326±0.336aò=0.822+0.017T99.5

二斑叶螨的捕食螨天敌很多，而伊氏叶螨被认为不为大多捕食螨所取食。deVasconcelosetal.(2008)比较研究了小植绥螨（Phytoseiulusfragariae）以上二者为食在不同温度条件下的生命表参数。结果表明，这种小植绥螨同样不是伊氏叶螨的良好天敌。c:用于数据分析的个体数温度、猎物对生命表参数的影响AbstractThecommerciallyavailablestrainsofPhytoseiuluspersimilisAthias-Henriot,thebiologicalcontrolagentofTetranychusurticaeKoch,performpoorlyintheWesternMediterranean,probablybecausetheyarenotwelladaptedtolocalclimaticconditions.Forthatreason,effortsarebeingfocusedonthedevelopmentofabiologicalcontrolprogrammeusingnativephytoseiidmites.FourspeciesofredspidermitescanbefoundinvegetablecropsineasternSpain:T.urticae,

TetranychusturkestaniUgarovandNikolski,TetranychusludeniZacherandtherecentlyintroducedTetranychusevansiBakerandPritchard.Toevaluatetheirpotentialroleasbiologicalcontrolagents,thepresentstudyevaluatesthelife-historyoflocalpopulationsofNeoseiuluscalifornicus(McGregor)andP.persimiliswhenfedonT.urticae,T.turkestani,T.evansi,andT.ludeniinthelaboratory.ResultsindicatethatN.californicusandP.persimilisareabletofeedandcompletetheirdevelopmentonthefourtestedredspidermitespecies.ThepredatorsmayexhibitaparticularlyhighcapacityforpopulationincreasewhenfedonT.urticae,T.turkestani,andT.ludeni,thusmaybeabletoprovideeffectivecontrolofthesespeciesinthefield.WhenfedT.evansi,however,predatorperformancewaspoor;significantincreaseindevelopmentandpreovipositiontimes,andareductioninovipositionperiodandfecunditywererecorded.

TheresultantlowcapacityforpopulationgrowthsuggestspoorabilityofthetwotestedpredatorstosuppressT.evansipopulationsoncommercialcrops.ItisunlikelythereforethatP.persimilisandN.californicus,nowbeingwidelyusedtocontrolT.urticaeingreenhousecropsinCentralEurope,willbeabletohaltanyspreadofT.evansitogreenhousecropsintemperateareas.营养物质对生命表参数的影响生殖力生命表用途评价生物(有害生物或天敌)的增殖情况评价一种天敌对多种有害生物的作用评价多种天敌对一种有害生物的作用评价环境因素对生物的作用2.

年龄特征生命表

(age-specificlifetable)

以种群的年龄阶段（如虫态或虫龄）作为划分的标准，系统观察并记载不同发育阶段或年龄区组中的死亡数量、死亡原因以及成虫阶段的繁殖数量。世代隔离清楚的种群自然种群分析影响种群数量变动的关键因子估算种群趋势指数组建预测模型小菜蛾Plutellaxylostella

在晚季甘蓝上第三代生命表

Canada，Ottawa(Harcourt，1969)年龄组xx开始时活虫数

lx主要致死原因dxF在x期内的死亡虫数dxx期内的死亡率100qxx期内的存活率SxK值卵(N1)1154未受精141.20.990.0044幼虫I(1-4龄中期）1140降雨53647.00.530.2757

幼虫II(4龄中期至结茧）604小茧蜂Microplitisplutellus

14023.2降雨7712.7小计21735.90.640.1938预蛹387寄生蜂Diadegmainsularis19851.20.490.3098蛹189姬蜂Diadromusplutella5328.20.720.1427成虫136性（40.1%♀♀）2719.90.800.0969雌虫×2109光周期52.448.10.520.2840“正常雌虫”×256.6成虫死亡48.185.00.150.8239全世代总计8.5-1145.599.3雌蛾的实际平均产卵量:112粒,标准产卵量:216粒生命表参数计算期望卵量=1/2×（“正常雌蛾×2”）×标准产卵量=6113实际卵量=½×全世代总计×标准产卵量=½×8.5×216=918趋势指数=918/1154=0.80雌虫×2=成虫数×雌虫比×2=136

×40.1%×2=109“正常雌蛾×2”=（雌虫×2）×雌蛾的实际平均产卵量/标准产卵量

=109×112/216=56.6种群存活曲线

在某一特定时刻，种群中的同龄个体随时间移动而减少的现象，可以用一条曲线表示，这条曲线就是存活曲线TypeI:survivewelluntilsenescence(largemammals,humans)TypeIIisastraightline:constantprobabilityofdeath(birds)TypeIII:heavyjuvenilemortality(inverts,fish,plants)Survivorshipcurves:threegeneraltypesareoftendescribed(Deevey1947)Typicalsurvivalcurveforhumanpopulation.Noticethatthedropofsurvivorshipinearlyyearsduetoinfantmortalityhasdramaticallyreducedinrecenttimes.Theageingofthepopulationisalsoveryapparent,asthesurvivalcurvedropoccurslaterinlifeandbecomessharperafter1950.Survivorshipcurve:Survivorshipcurveskeeptrackofthefateofanygivenbirthcohort.Theyshowthepercentstilllivingatagivenage.Nowadaysinthedevelopedworldfewchildrendiebeforereproduction.InGreatBritainin1999only1%ofallchildrenbornalivediedbytheageoffive(comparedto10%inIndia,and35%inNiger).However,300yearsagoitwasquiteadifferentmatter,asthegraphaboveillustrates.IntheCityofYork(England)inthe17th.Century,only15%madeittothethresholdofreproduction(15yrs.).Only10%remainedalivebytheageoftwenty.Withsofewfemaleslivingtoreproduction,onlyahighfertilityratecouldmaintainthepopulation.Notethatchangeswitheconomicdevelopment,asshownbyNigerandIndia.NotealsotheimpactofbiasagainstfemalesinIndiaontheirsurvival--otherwise,India'scurvein1999isverysimilartoGreatBritain'sforthelate19th.C.(notshown).南美玉米苗斑螟(Elasmopalpuslignosellus)的３个世代的平均生存率（Smithetal.1989)新生若虫接虫后周数生存率％麦蚜（Sitobionavenae）的存活曲线(Thirakuptetal.1992)绵蜡蚧（Pulvinariaregalis）年生活史中的生存率(Speightetal.1999)

两种小花蝽在三种恒温条件下对西方花蓟马捕食的存活曲线及生殖率.种群趋势指数种群趋势指数（trendindexofpopulation)I值：下代数量和当代数量的比值

I=Ni+1/Ni（i=1，2，3，……表示世代）I＜1时，次代种群数量下降，即种群数量发展趋势下降I=1时，次代种群数量与当代相当，即种群数量保持原来水平I＞1时，次代种群数量上升，即种群数量发展趋势增长Morris（1963）&Watt(1961,1963)数学模式

SE…………….卵的存活率SL1

SL2……….幼虫各龄的存活率SPP……………预蛹的存活率SP……………..蛹的存活率SA…………….成虫的存活率P♀…………….雌虫占成虫总数的百分率PF…………….达到标准产卵量成虫占的百分率F……………标准产卵量

I=SE∙SL1∙SL2∙SL3∙∙∙∙∙∙SPP∙SP∙SA∙F∙PF∙P♀

在某虫期增加死亡率对I值的影响从I值成分中抽出一个组分对I值的影响I=S1∙S2∙S3∙S4∙∙∙∙∙∙Sn∙F∙PF∙P♀在某虫期增加死亡率对I值的影响I=S1∙S2∙S3∙S4∙∙∙∙∙Si∙∙∙∙∙∙∙Sn∙F∙PF∙P♀aiI(ai)=S1∙S2∙S3∙S4

∙∙(Si-ai)∙∙∙∙Sn∙F∙PF∙P♀I(ai)/I=(Si-ai)/Si=1-ai/Si令M(ai)=I(ai)/I，则：M(ai)=1-ai/SiI(a1,a2…an)=I(1-a1/S1)∙(1-a2/S2)∙∙∙(1-an/Sn)M(a1,a2…an)=(1-a1/S1)∙(1-a2/S2)∙∙∙(1-an/Sn)............(1)............(2)(2)/(1)释放赤眼蜂防治稻纵卷叶螟（庞雄飞，1980）已知:

第三代卵量（N3）与第二代卵量(N2)的比值I=N3/N2=5.2

第二代卵的自然寄生率平均为10%求：第二代卵期释放赤眼蜂后，使第三代卵量维持或低于第二代水平，问释放赤眼蜂后，卵的寄生率应该达到多少才行？I（a）≤1I（a）=I(1–a/Sa)≤15.2(1-a/(1-10%))≤1a≥0.7269卵的寄生率达到82.69%I(ai)/I=1-ai/Si散放赤眼蜂后稻纵卷叶螟生命表（广东，阳江海陵，庞雄飞等

1976）虫期x死亡原因dxF放蜂区对照区x期死亡率

100qxx期存活率

100Sxx期死亡率

100qxx期存活率

100Sx

卵捕食及其它46544654寄生（放蜂区放蜂）86143466幼虫1龄捕食及其它247612882-3龄捕食及其它24761288寄生861484.3315.674-5龄捕食及其它24761090寄生36642476预蛹寄生298199蛹捕食1486793寄生36.463.623.676.4雌虫百分率（P♀)=57.14%,

平均每雌产卵量（F∙PF)=74假定放蜂区的I值为It,对照区为Ic,Mt

为放蜂区It对对照区Ic增长的倍数Ic=0.54×0.66×0.88×0.88×0.1567×0.90×0.76×0.99×0.93×0.764×74×0.5714

=0.88It=0.54×0.14×0.76×0.76×0.14×0.76×0.64×0.98×0.86×0.636×74×0.5714

=0.067Mt=It/Ic=0.067/0.88=0.076校正死亡率组分分析法I

值分析法校正死亡率=78.79%I(ai)/I=1-ai/SiI(ai)/I=1-(0.86-0.34)/0.66=0.21

从I值成分中抽出一个组分对I值的影响I=S1∙S2∙∙∙∙∙∙

∙∙∙∙S（i-1）∙Si∙S（i+1）

∙∙∙∙∙∙∙∙Sn∙F∙PF∙P♀I(Si)=S1∙S2∙∙∙∙∙∙∙∙∙S（i-1）∙S（i+1）

∙∙∙∙∙Sn∙F∙PF∙P♀Si越大，M（Si）越小，表示Si对I值的作用相对较小Si越小，M（Si）越大，表示Si对I值的作用相对较大M(S1)=1/S1=1/0.54=1.85M(S2)=1/S2=1.52M(S3)=1/S3=1.14M(S4)=1/S4=1.14M(S5)=1/S5=6.38M(S6)=1/S6=1.11M(S7)=1/S7=1.32M(S8)=1/S8=1.01M(S9)=1/S9=1.08M(S10)=1/S10=1.31M(P♀)=1/SP♀

=1.75M(PF)=1/SPF=2.70M（S5）最大，即由于S5的作用，使I值增加了6.38倍。释放赤眼蜂后对稻纵卷叶螟的影响比较各组分对I值的作用M(S1)·M（S2）

········M(Sn)·M(PF)·M（SP♀）=♀

K=k1+k2+k3++++++++kn+KPF+kP♀♀令关键因子分析1.回归分析法（Morris-Watt1959)

关键因子（Keyfactors):

指对下代（或经历一段时间）种群数量变动起主导作用的因子2.K-值图解法（Varley&Gradwell1960)

二代稻纵卷叶螟生命表关键因子分析-回归分析法(古德祥等,1983)K1K2K3K4K5K6K7K8K9K0.28610.065700.67200.01240.194600.23910.26661.73650.21580.02110.01320.24360.04090.10830.00750.21270.05780.92090.23650.02050.03210.59710.02110.53790.01420.57650.02232.05820.44440.06520.03510.90710.0360.992500.466002.91390.31520.04390.03640.36700.05630.46180.04690.63970.05402.02120.23410.04180.03390.94160.01200.55020.02100.77020.01642.6212死亡原因卵被捕食卵被寄生

卵不孵化1-2被捕食1-2龄被寄生3-5龄被捕食3-5龄被寄生蛹被捕食蛹被寄生r0.6250.5140.6610.862-0.5800.9060.0440.676-0.401b0.0760.0150.0140.346-0.0170.4060.0010.216-0.057K值图表法若某个Ki值的变化趋势与总的K值变化趋势最为相似，则该Ki所代表的因子即为关键因子舞毒蛾Lymantriadispar(L)的生命表（新西兰，根据Campbell1981修改）StageMortalityfactorInitialno.ofinsectsNo.ofdeathsMorta