大斑啄木鸟对光肩星天牛幼虫捕食的功能反应和数值反应.pdf

动物学报 54(6):1106-1111,2008Acta Zoologica Sinica 2008-02-12收稿,2008-09-20 接受*霍英东教育基金会高等院校青年教师基金项目(No1101025)、十一五科技支撑计划课题/林业重大生物灾害防控新技术产业化与示范0(No12006BADO8A12)、长江学者和创新团队发展计划(No1IRT0607)Foundation items:The Fok Ying Tong Education Foundation forYong Teachers in Colleges and Universities(No1101025),National Science-Technology Support of China during the 11thFive-Year Plan Period(No12006BADO8A12),Program for Changjiang Scholars and Innovative Research Team in University(No1IRT0607)*通讯作者(Corresponding author)1 E-mail:wenjb bjfu1edu1cn Z2008 动物学报 Acta Zoologica Sinica大斑啄木鸟对光肩星天牛幼虫捕食的功能反应和数值反应*矫振彪1 万 涛1 温俊宝1*胡加付1 骆有庆1 张林生2 付林巨21.北京林业大学省部共建森林培育与保护教育部重点实验室,北京 1000832.内蒙古自治区乌拉特前旗森林病虫害防治检疫站,乌拉特 前旗 014400Functionalresponse and numericalresponse of great spottedwoodpeckerPicoidesmajoronAsianlonghornedbeetleAnoplophora glabripennis larvae*JIAO Zhen-Biao1,WAN Tao1,WEN Jun-Bao1*,HU Jia-Fu1,LUO You-Qing1,ZHANG Lin-Sheng2,FU Lin-Ju21.Key Laboratory for Silviculture andConservation,Ministry of Education,Beijing Forestry University,Beijing 100083,China2.The Forest Disease and Pest Control station of Wulate Qianqi,Inner Mongolia Autonomous Region,Wulate Qianqi 014400,ChinaAbstract The Asian longhorned beetle Anoplophora glabripennis is a wood-boring beetle that,upon severe outbreaks,causesgreat damage in the artificial shelter-forest of the Three-NorthernAreas.The great spottedwoodpecker Picoides major is one of thenatural predators of A1 glabripennis.P1 major is also a common species in Wulate Qianqi of the Inner Mongolia AutonomousRegion and iswidely distributed in the artificial shelter-forest.The wood-boring beetle is an important food source of P1 major,especially during the winter.Despite the potentialof P1 major to control A1 glabripennis populations,surprisingly little is knownabout their predator-prey interactions.This information is essential to predicting the efficacy of P1 major as a control agent to A1glabripennis.Therefore,it is important to study how the predator responds to changes in prey density.From 2006 to 2008,fifteenstudy plots were selected,and the predation of P1 major on A1 glabripennis was examined with sample plot and traceinvestigation,the latter being in the field.The results showed that the function response of P1 major to A1 glabripennis wasHolling,with a non-linear regression equation.The increase in predation capacity correlated with the increase in A1glabripennis density,within a certain density range.As the density of A1 glabripennis increased,the predatory capability had apositive acceleration phase,then a negative acceleration phase,and finally a saturation state.Likewise,as the density of A1glabripennis increased,the predation rate initially increased,peaked,and was reduced in the end,with a non-linear regressionequation.The results indicated that the numerical response was a positive reaction to density.In other words,the populationdensity of P1 major increased as the density of A1 glabripennis increased Acta Zoologica Sinica 54(6):1106-1111,2008.Key words Anoplophora glabripennis,Picoides major,Predation,Functional response,Numerical response关键词 光肩星天牛 大斑啄木鸟 捕食 功能反应 数值反应 光肩星天牛 Anoplophora glabripennis(Motsch.)是我国重要林木蛀干害虫,造成了巨大的经济损失(张星耀、骆有庆,2003)。大斑啄木鸟 Picoidesmajor(L.)是林业蛀干类害虫的重要天敌,关于啄木鸟对蛀干类害虫的控制作用国内已有一些报道,冬季在杨树人工林内主要以光肩星天牛等蛀干类害虫的幼虫为食(郑作新,1958;张仲信,1981,1986,1992;高瑞桐等,1994)。鸟类对农林害虫控制作用的研究,国外也有部分报道(Greenberg etal.,2000;Tremblay et al.,2001;Mols and Visser,2002;Low and Connor,2003)。功能反应和数值反应是用来描述捕食者的作用及其数量如何依赖于猎物的密度变化而变化的,从而估计在被食者密度上升时捕食者的作用强度及其控制 效 果(郑 汉业、夏 乃 斌,1995;孙 儒泳,2001)。在国内,目前对于功能反应和数值反应的研究主要是对捕食性螨类和捕食性昆虫等体型相对较小的动物,且主要在室内进行实验(赵志模等,1988;任顺祥等,2001;任月萍、刘生祥,2006),对于体型相对较大的鸟类及其它脊椎动物的功能反应和数值反应研究在国内较少(楚国忠,1989;陶双伦等,2003)。而国外近年来关于鸟类、兽类、草食型动物、水生生物等捕食作用的研究较多(Gross et al.,1993;Norris and Johnstone,1998;Fritzet al.,2001;Jeschke et al.,2002;Biro et al.,2003;Stillman and Simmons,2006;Wong and Barbeau,2006)。在研究鸟类对森林害虫的作用时,许多学者强调鸟类对害虫功能反应和数值反应的重要性,并通过对这些反应的分析,估计在不同害虫发生水平及害虫不同发育阶段鸟类的捕食作用(Buckner,1967;Gaye et al.,1970;楚国忠,1989)。我国西北地区光肩星天牛危害严重,大斑啄木鸟作为其重要的捕食性天敌,近年来在有些地区种群呈增长趋势(李刚等,2000)。Jeschke et al.(2002)认为,研究捕食者和猎物间的捕食关系,可以预测两者种群变动趋势。本文通过研究两者之间功能反应和数值反应,为进一步研究大斑啄木鸟对光肩星天牛的控制作用及两者种群互动关系提供依据。1 材料与方法111 研究区域自然概况乌拉特前旗位于内蒙古自治区河套平原东部,地处北纬 40 b28 c-41b16c,东经 108b11c-109b54c,属中温带大陆性气候。试验地段选择在乌拉特前旗河套灌区,主要水源为黄河引流的灌溉水。试验地段主要以农田防护林网为主,辅以村庄四旁林及部分片林。主要树种有杨树(Populus spp.)、旱柳(Salix matsudana)、榆 树(Ulmusspp.)、沙 枣(Elaeagnusangustifolia)、红柳(Tamarixramosissima)等多种乔木,森林覆被率 5%(郑丽颖等,2006)。112 研究方法11211 功能反应2005 年 8 月开始,在内蒙古自治区乌拉特前旗开展了大斑啄木鸟的种群数量、动态、取食行为、生态位、捕食选择等等方面的研究(郑丽颖等,2006;万涛等,2008)。2006 年 12 月至2007年1 月及 2007年 12 月至 2008年 1月,根据蛀干害虫危害程度的划分标准(薛贤清,1992),按光肩星天牛轻、中、重三种危害程度(有虫株率 10%为轻度危害,10%0 说明功能反应的类型为Hholling 型。应用 SPSS1310 软件对光肩星天牛虫口总数和大斑啄木鸟的捕食量进行曲线回归(S 模型回归法),回归方程为:Ne=exp(41323-391758 P No)()R2=01951,模型的回报精度达 79186%。图 1 不同光肩星天牛虫口密度下大斑啄木鸟的捕食量Fig 11 ThepredatorycapacityofP1majoronA1glabripennis 由图 1 也可以看出两者间捕食关系呈 Holling型,即为脊椎动物型或 S 型。随着光肩星天牛虫口密度的增加,大斑啄木鸟的捕食量在虫口密度较小时增长较快,但当虫口密度较高时增速变小;即大斑啄木鸟对光肩星天牛幼虫的捕食功能反应开始有一个正加速期,随后是负加速期,而后达到饱和水平。以光肩星天牛虫口密度为横坐标,大斑啄木鸟的捕食率为纵坐标,应用软件 Sigmaplot 1010 做图并进行回归分析得图 2。回归分析的方程为:y=14128+185103exp-0157 lnx71432x()1108动 物 学 报54卷 图 2 捕食率与光肩星天牛虫口密度的关系Fig12 The predation ratio ofP1major on A1glabripennis R2=0186。其中:x 为光肩星天 牛的虫口密度(头P 株),y 为大斑啄木鸟对光肩星天牛的捕食率。模型回报精度可以达到 88141%。由图 2 可以看出,大斑啄木鸟的捕食率随光肩星天牛虫口密度的增加呈现先增加后下降的趋势。根据回归方程,可以计算不同虫口密的捕食率。当虫口密度在0-311 头 P 株时,捕食率会随着虫口密度的增加而增加,在虫口密度为 311 头 P 株时捕食率达到最大值 5219%。当虫口密度大于 311 头P 株时捕食率会随着虫口密度的增加而减小,当虫口密度大于1912 头P 株时捕食率小于 20%。当虫口密度为211-414 头P 株时,捕食率大于 50%,即有一半以上的天牛幼虫被捕食。212 数值反应通过对不同光肩星天牛危害程度的 16 个样地的调查,得到不同样地的光肩星天牛危害有虫株率和大斑啄木鸟的种群密度,计算出无光肩星天牛危害和轻、中、重三种光肩星天牛危害等级下大斑啄木鸟的平均种群密度,应用最小显著差数法对数据进行差异显著性检验,结果见图 3。轻度危害区和没有天牛危害区的大斑啄木鸟种群密度差异不显著(P 0105),说明光肩星天牛虫口密度较小时,其对大斑啄木鸟种群密度影响不大。中度危害区和轻度危害区的大斑啄木鸟的种群密度差异显著(P 0105)但和轻度危害区差异显著(P 0105)。说明大斑啄木鸟的种群密度随着光肩星天牛虫口密度的增加而增加,数值反应的类型为正密度反应。图 3 不同光肩星危害程度下大斑啄木鸟的种群密度最小显著差数法;小写字母表示组间最小显著差数法检验P 0105;误差线为标准偏差。Fig 13 The population density of P1 major in fourdifferent damage classificationLeast significant difference;lowercase P 0105;the error line isthe standard deviation.3 讨 论光肩星天牛虫口密度较小时,大斑啄木鸟的捕食量和捕食率都很低,因为大斑啄木鸟接触到光肩星天牛幼虫的几率较小,从而影响了捕食效率;并且大斑啄木鸟可以取食节肢动物、植物种子等多种食物(郑作新,1958;张仲信,1986;高瑞桐等,1994;Michalek and Miettinen,2003),此时大斑啄木鸟可以通过取食其它食物资源或者增加取食的范围来增加取食量(Simon,1995)。捕食者和猎物接触较少时,捕食者对猎物不能形成条件反射快速发现猎物(郑汉业、夏乃斌,1995),大斑啄木鸟具有较 高的智力水 平及很强 的记忆及 学习能力(Michalek andMiettinen,2003),在光肩星天牛数量比较少时,鸟虫之间的接触较少,不能建立条件反射以很快地发现和识别食物。当光肩星天牛虫口密度由较低水平增加时,由于大斑啄木鸟寻食到光肩星天牛的几率增大并且频繁的接触使得大斑啄木鸟捕食光肩星天牛的能力也大大加强,因此大斑啄木鸟对光肩星天牛的捕食量显著增加。此时大斑啄木鸟对光肩星天牛幼虫的捕食率也显著增加,并维持在一个较高的水平。大斑啄木鸟与光肩星天牛之间的数值反应为正密度反应。总体上,大斑啄木鸟的种群密度会随着光肩星天牛的密度增加呈现增长趋势。当光肩星天牛的虫口密度较高时,不仅可以为大斑啄木鸟的生存和繁殖提供充足的食物来源,而且由于光肩星天11096期矫振彪等:大斑啄木鸟对光肩星天牛幼虫捕食的功能反应和数值反应 牛危害而枯死、腐朽的枯立木,也为大斑啄木鸟的营巢定居提供便利条件(B tler et al.,2004)。由于Holling 型功能反应在猎物密度上升到一定的阈值之前一直受到密度制约,在平均猎物密度低 于这一 阈值 时,将可 保证种 群的 稳定 性(Holling,1959;陈均平、张洪德,1986;Hosseiniet al.,2005)。当光肩星天牛虫口密度较低时,大斑啄木鸟的控制作用能有效抑制其虫口密度,此时可通过各种措施(悬挂人工鸟巢招引、加强人为保护)稳定大斑啄木鸟的种群(杨奋勇等,2006),达到长期持续控制光肩星天牛的目的。光肩星天牛虫口密度较高时,大斑啄木鸟用于搜索猎物的时间减少,搜寻猎物的成功率增加;但其在特定时间内的食量是有限的,并且虫口密度较高时大斑啄木鸟趋向于取食进入木质部的大幼虫(万涛等,2008),使得大斑啄木鸟的取食量(啄食幼虫数量)没有显著增加。此时,捕食量没有显著增加,捕食率反而呈下降趋势,对光肩星天牛的控制作用减弱,将无法保证种群的稳定性(陈均平、张洪德,1986;Hosseini et al.,2005),需要结合其它措施(多树种配置、砍伐、截干)进行光肩星天牛的综合控制。参考文献(References)Biro PA,Post JR,Parkinson EA,2003.From individuals to populations:prey fish risk-taking mediates mortality in whole-system experiments.Ecology 84(9):2419-2431.Buckner CH,1967.Avian and mammalian predators of forest insects.Biocontrol 12:491-501.B tler R,Angelstam P,Ekelund P,Schlaepfer R,2004.Dead woodthreshold values for the three-toed woodpecker presence in boreal andsub-Alpine forest.Biological Conservation 119(3):305-318.Chen JP,Zhang HD,1986.The qualitative analysis of two species predator-prey model with Hollingc s type functional response.AppliedMathematics and Mechanics 7(1):73-80(In Chinese).Chu GZ,1989.Growth and 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