北方地区重大迁飞性害虫的监测与种群动态分析
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摘要
我国地处东亚季风区，农作物重要害虫大部分都具有远距离迁飞的习性。迁飞性害虫以其突发性、暴发性、毁灭性的特点始终威胁着我国的粮食安全。近年来，由于全球气候异常与耕作方式变更等原因，近年来，新发害虫二点委夜蛾连年暴发成灾，重大迁飞性害虫草地螟、小地老虎和黏虫等也呈现间歇性、局部大面积暴发的态势。为了提高迁飞性害虫的监测预警水平，及时开展有效防控，减少农业损失和药剂防治带来的一系列问题，本文利用自行设计的高空诱捕器和自动分时段取样的探照灯诱虫器等监测设备，在北京延庆、吉林公主岭和内蒙古锡林浩特等多个监测点获得的多年野外虫情，对我国北方多种夜行性昆虫的探照灯诱虫数量与高空取样结果之间的关系，多种昆虫的扑灯类型和迁飞性昆虫的上灯节律及其对轨迹分析的潜在影响进行了研究。针对新发害虫二点委夜蛾和重大迁飞性害虫小地老虎和黏虫，除分析探照灯下的种群动态以外，又结合垂直监测昆虫雷达获得空中虫情、卵巢解剖数据、地面和空中气象资料、全国农技推广中心收集的各地虫情数据等，综合运用GrADS、HYSPLIT、ArcMap等图形处理与统计分析软件，重点研究了种群动态与各种气象因素之间的关系，系统分析了种群波动与空中风场之间的关系，并探讨了二点委夜蛾的虫源性质及其迁飞的可能性。本文主要获得了以下几方面的结果：
1.新设计的空中诱捕器可在空中风力不足以张开拖网的情况下进行空中取样，并可有效防止诱捕昆虫的逃脱。试验表明，该诱捕器在200~400m高度可捕获隶属于半翅目Hemiptera、鳞翅目Lepidoptera、鞘翅目Coleoptera等7目的多种昆虫；高空取样结果证实了探照灯诱虫器在辅助雷达进行迁飞目标识别中的作用。
2.探照灯分时段诱捕数据表明，直翅目的东方蝼蛄、北京油葫芦和鞘翅目步甲类等优势种属于非整夜扑灯型，脉翅目的大草蛉和丽色草蛉，鳞翅目的草地螟、玉米螟、甜菜白带野螟、小菜蛾和大多数夜蛾科种类都属于整夜扑灯型。不同种类其扑灯节律明显不同，迁飞习性是影响其在探照灯诱捕器内出现节律的最重要因素之一。当轨迹模拟限制在10h以内时，分时段参数化所得的结果并不是传统方法所得结果在距离上的简单缩短，而是涉及更多的虫源地、可能的降落地点和运转路径。但随着轨迹模拟时间的延长，两种方法得到的虫源地信息基本相同。
3.综合分析表明，二点委夜蛾可以在北京延庆地区越冬。二点委夜蛾在河北栾城1年发生4代，北京延庆1年发生3代，两地的雄虫诱集总量均极显著高于雌性，实际性比显著或极显著小于1的天数，在两地所占的比例分别达到了58%和30%。种群波动影响性比变化，当种群处于上升期时，二点委夜蛾成虫的性比极显著高于1，当种群进入平缓期后，性比接近于1或极显著小于1。北京延庆探照灯诱集的一代成虫数量不符合正态分布，一代成虫出现两个明显高峰，高峰期诱虫数量与空中925hPa的偏南气流密切相关。北京周边地区的监测数据也表现出不同程度的同期增长，初步判断二点委夜蛾是一种兼性迁飞昆虫，北京延庆一代成虫除包含本地种群外，还包括从河北和天津迁飞或扩散而来的个体。
4.室内饲养和吊飞发现，小地老虎的发育历期和飞行能力可能导致性比发生变化。一代发生区，3-4月份的温度和降水对当年小地老虎的发生程度具有重要影响，4月份温度偏高，降水量较大的年份小地老虎发生重，反之较轻；6月下旬小地老虎迁飞高峰期的雷达回波高度主要在500m以下，300~400m是其主要的飞行高度。垂直雷达监测和夜间定时取样均表明小地老虎可以整夜迁飞。风场分析表明，有利的天气系统可促进小地老虎的迁飞与危害。
5.2008-2012年，各年累计诱集黏虫量分别为5,105头、3,981头、11,385头和50,584头，诱集数量在年度间存在极显著差异，2012年是黏虫诱集数量最高的一年。气象分析表明，适宜黏虫繁殖的气象条件是导致2012三代黏虫大暴发的直接原因。7月份适宜黏虫扩散的天气系统是三代黏虫暴发的外在动力。北京延庆监测点二代黏虫成虫诱集数量与空中气流的运转方向密切相关，轨迹分析显示华北北部地区三代黏虫大暴发的虫源大部分是从周边地区迁飞扩散而来，是迁入种群和本地种群在适宜气象条件下共同繁殖的结果，与东北黏虫种群关系不大。东北地区三代黏虫的虫源大部分来自本地，受偏西气流和降雨的影响，东北地区二代成虫不能有效迁出，黑龙江北部二代成虫在偏南气流的运载下向南迁飞，受锋面天气影响降落在黑龙江、吉林交界处，与东北地区北部和华北地区迁入的成虫汇集后，造成了东北局部地区的严重发生。三代黏虫回迁期，由于受东北和华北地区地形和季风的影响，很难形成合适的偏北气流，9月中下旬，华北、华中及华南地区农作物接近成熟期，基于上述因素，预测4代黏虫不会在全国大面积暴发，后期实际发生情况与预期结果基本一致。
China is located in the East Asian monsoon area, where many key crop pests are migrants.Irregular outbreaks of these migratory pest insects cause severe damages to crop, and therefore, threatenagricultural production and food security. There was a sudden outbreak of Proxenus lepigone a new pestinsect of maize in China in2011, and intermittent heavy occurrence of the meadow moth Loxostegesticticalis L., black cutworm Agrotis ypsilon Rottemberg, armyworm Mythimna seperata in recent years,due to the anomalies of global climate and changes in tillage practices, etc. This study aims to improvethe level of monitoring and forecasting on migratory pest insects and benefit integrated management,and also to prevent losses in agriculture and reduce many social problems caused by chemical control.In the former part, catches from aerial samples by the self-designed aerial trap and by searchlight trapwere compared, and whether the nocturnal insects could be trapped during the whole night were test onmany species based on the data of many years obtained through searchlight traps with automatictime-switches that have been deployed in Yanqing Beijing, Gongzhuling Jilin and Xilinhaote InnerMongolia, Also, the dynamics of catch sizes during the whole night and related potential effect on thetrajectory analysis were studied. Focusing on the black cutworm and armyworm, this study integratedthe data from searchlight traps with the data obtained by vertical-looking radar, included condition ofovarian development, ground and aerial meteorological data from weather station or internet, and pestinformation from the National Agro-Technical Extension and Service Centre. Then we integratedapplication of the wind field analysis, trajectory simulation and geographic information system softwaresuch as GrADS, HYSPLIT and ArcMap to analyze these data and investigate the influence of variousmeteorological factors on the population dynamics and the relationship between population fluctuationsand wind field. Also, this study explored the possibility of be migrantion of P. lepigone, and its sourceplace based on the data mentioned above. The results were as follows:
1. this newly designed aerial trap for high-flying insects could function when wind speed is notstrong enough to open the tow net, and is able to prevent the catches from escaping. Aerial samplingtests showed that this aerial trap could catch many individuals from7orders such as Hemiptera,Lepidoptera, Coleoptera etc. at the height of200~400m above the ground level. For the migratory pestinsects including L. sticticalis, M. separatar, Helicoverpa armigera Hubner, that were trapped in aerialsamples, the catch sizes in searchlight trap were consistently in peaked periods, which confirmed thatsearch light trap could be a practicable tool assisting radar in identification of the high-flying targets.
2. Dominant species of Orthoptera and Coleoptera only appeared in some time-sections at night,but Sympetrum Croceolum and Chrysopa formosa from Neuroptera, L. sticticalis, Pyrausta nubilalis,Hymenia recurvalis, Plutella xyllostella and many noctuids from Lepidoptera could be trapped duringthe whole night. Dynamic rhythm of catch sizes differed in different species. Distant migratory behaviorwas one of the most important factor the dynamic of specific species. When limiting the function timefor only1night, the trajectory simulation showed that the results obtained as parameterized withdifferent starting time involved more insect source place and possible landing sites, not simply thetruncated distance as generated by the traditional methods. Howerver, source places obtained by bothmethods tend to be similar with the extension of the function time.
3. In2012, the minimum temperature in Yanqing of Beijing was higher than the supercooling point(-25.35℃) of the mature larvae with cocoon. With other evidences taken into consideration, theresultsindicate that the mature larvae with cocoon of P. lepigone could therefore overwinter in YanqingBeijing. Monitoring data confirm that P. lepigone has occurred4generations per year in LuanchengHebei province and3generations per year in Yanqing Beijing. the total amount of trapped males wassignificantly higher than that of female in both sites. Nights when sex ratio of P. lepigone wassignificantly smaller than or close to1accounted for58%and30%in Yanqing and Luanchengrespectively. Variation in population sizes influences the sex ratio that changes in same trend for bothsites, that is, the sex ratios were significantly higher than1when population size rise, but close to1orsignificantly smaller than1in the the steady periods. The first generation of trapped P. lepigone adultsdid not form the normal distribution in Yanqing, and two obvious peaks appeared in the this generation.Variations in population sizes were related to the wind field at925hPa. With comprehensiveconsideration of the synchronous spurts in other light traps around Beijing, we conclude that the trappedfirst generation was composed not only the local individuals, but also the migrants or dispersers fromHebei and Tianjin，therefore, P. lepigone is a facutrative migratory species.
4. Indoors feeding observation showed that there were no significantly differences in the totaldevelopment duration between sexes. Tests on potential flight ability indicated that the males are ofmore powerful flight ability. So, total development duration and flight ablilty may lead to the changes insex ratioes. In the regions where first generation occurred, temperature and precipitation in March andApril had heavy influences on the occurrence of A. ypsilon, and it seemed that higher temperature andmore rains benefit the development of A. ypsilon. In the late June, the observation with entomologicalradar confirmed that most moths of A. ypsilon generally flew below500m, ant that the biggest densitywas at height of300~400m. The observation that moths of A. ypsilon can fly during the whole nightwere approved by both radar observation and timely samples of the catches from searchlight trap.Migrants of A. ypsilon could be windborne in that every peaks in trapped moths were alwaysaccompanied with favorable wind field and velocity, furthermore, the northern cyclones frontal weatherand heavy rainfall would cause massive landing.
5. In2008~2012, total trapped moths of armyworm were5105,3981,11385and50584,respectively, among which a significantly difference was identified. For armyworm, the biggest catchsizes was in2012and more than7300moths were trapped at one night which had nerver occurredbefore since this monitoring station has functioned. In conclusion, favorable weather condition were themain factors leading to the outbreak of larvae of3rd generation. Taken the relationship between catchsizes and wind fields, most of trapped moths were thought to be immigrants from surrounding regions.Therefore, appropriate weather condition for migrants in July promoted the outbreak of the3rd-generation larvae in2012. In North China, the outbreaks were mainly the result of the jointreproduction of immigrants and the local adults under the favorable environment condition and littleconcerned with adult population in Northeast China. For northeast China, the source moths of3rd-generation larvae in Northeast China were mainly from the local, because the local2nd-generationmoths in Northeast China failed to immigrate into the North China and had to stay in the natal areas ormake short southern migration due to westerly flow and rainfall in late July. The southward 2nd-generation moths in Heilongjiang mainly landed at the junction of Heilongjiang and Jilin provincesas a result of wind shear associated with heavy precipitation, lead to the serious outbreak at possiblelanding places, together with the immigrants from the North China. During the return migration seasonof3rd-generation adults, considering the influence of the terrain and monsoon in Northeast and NorthChina, we determined that only on some occasion the wind field can carry moths into North China fromNortheast China. In addition, almost all fall crops were in harvest at that time. Combining theseconsiderations together, we forecasted there was little chance for4th-generation larvae causing heavydamage to crop, which was proved to be true by the actual occurrence of4th larvae.
1.新设计的空中诱捕器可在空中风力不足以张开拖网的情况下进行空中取样，并可有效防止诱捕昆虫的逃脱。试验表明，该诱捕器在200~400m高度可捕获隶属于半翅目Hemiptera、鳞翅目Lepidoptera、鞘翅目Coleoptera等7目的多种昆虫；高空取样结果证实了探照灯诱虫器在辅助雷达进行迁飞目标识别中的作用。
2.探照灯分时段诱捕数据表明，直翅目的东方蝼蛄、北京油葫芦和鞘翅目步甲类等优势种属于非整夜扑灯型，脉翅目的大草蛉和丽色草蛉，鳞翅目的草地螟、玉米螟、甜菜白带野螟、小菜蛾和大多数夜蛾科种类都属于整夜扑灯型。不同种类其扑灯节律明显不同，迁飞习性是影响其在探照灯诱捕器内出现节律的最重要因素之一。当轨迹模拟限制在10h以内时，分时段参数化所得的结果并不是传统方法所得结果在距离上的简单缩短，而是涉及更多的虫源地、可能的降落地点和运转路径。但随着轨迹模拟时间的延长，两种方法得到的虫源地信息基本相同。
3.综合分析表明，二点委夜蛾可以在北京延庆地区越冬。二点委夜蛾在河北栾城1年发生4代，北京延庆1年发生3代，两地的雄虫诱集总量均极显著高于雌性，实际性比显著或极显著小于1的天数，在两地所占的比例分别达到了58%和30%。种群波动影响性比变化，当种群处于上升期时，二点委夜蛾成虫的性比极显著高于1，当种群进入平缓期后，性比接近于1或极显著小于1。北京延庆探照灯诱集的一代成虫数量不符合正态分布，一代成虫出现两个明显高峰，高峰期诱虫数量与空中925hPa的偏南气流密切相关。北京周边地区的监测数据也表现出不同程度的同期增长，初步判断二点委夜蛾是一种兼性迁飞昆虫，北京延庆一代成虫除包含本地种群外，还包括从河北和天津迁飞或扩散而来的个体。
4.室内饲养和吊飞发现，小地老虎的发育历期和飞行能力可能导致性比发生变化。一代发生区，3-4月份的温度和降水对当年小地老虎的发生程度具有重要影响，4月份温度偏高，降水量较大的年份小地老虎发生重，反之较轻；6月下旬小地老虎迁飞高峰期的雷达回波高度主要在500m以下，300~400m是其主要的飞行高度。垂直雷达监测和夜间定时取样均表明小地老虎可以整夜迁飞。风场分析表明，有利的天气系统可促进小地老虎的迁飞与危害。
5.2008-2012年，各年累计诱集黏虫量分别为5,105头、3,981头、11,385头和50,584头，诱集数量在年度间存在极显著差异，2012年是黏虫诱集数量最高的一年。气象分析表明，适宜黏虫繁殖的气象条件是导致2012三代黏虫大暴发的直接原因。7月份适宜黏虫扩散的天气系统是三代黏虫暴发的外在动力。北京延庆监测点二代黏虫成虫诱集数量与空中气流的运转方向密切相关，轨迹分析显示华北北部地区三代黏虫大暴发的虫源大部分是从周边地区迁飞扩散而来，是迁入种群和本地种群在适宜气象条件下共同繁殖的结果，与东北黏虫种群关系不大。东北地区三代黏虫的虫源大部分来自本地，受偏西气流和降雨的影响，东北地区二代成虫不能有效迁出，黑龙江北部二代成虫在偏南气流的运载下向南迁飞，受锋面天气影响降落在黑龙江、吉林交界处，与东北地区北部和华北地区迁入的成虫汇集后，造成了东北局部地区的严重发生。三代黏虫回迁期，由于受东北和华北地区地形和季风的影响，很难形成合适的偏北气流，9月中下旬，华北、华中及华南地区农作物接近成熟期，基于上述因素，预测4代黏虫不会在全国大面积暴发，后期实际发生情况与预期结果基本一致。
China is located in the East Asian monsoon area, where many key crop pests are migrants.Irregular outbreaks of these migratory pest insects cause severe damages to crop, and therefore, threatenagricultural production and food security. There was a sudden outbreak of Proxenus lepigone a new pestinsect of maize in China in2011, and intermittent heavy occurrence of the meadow moth Loxostegesticticalis L., black cutworm Agrotis ypsilon Rottemberg, armyworm Mythimna seperata in recent years,due to the anomalies of global climate and changes in tillage practices, etc. This study aims to improvethe level of monitoring and forecasting on migratory pest insects and benefit integrated management,and also to prevent losses in agriculture and reduce many social problems caused by chemical control.In the former part, catches from aerial samples by the self-designed aerial trap and by searchlight trapwere compared, and whether the nocturnal insects could be trapped during the whole night were test onmany species based on the data of many years obtained through searchlight traps with automatictime-switches that have been deployed in Yanqing Beijing, Gongzhuling Jilin and Xilinhaote InnerMongolia, Also, the dynamics of catch sizes during the whole night and related potential effect on thetrajectory analysis were studied. Focusing on the black cutworm and armyworm, this study integratedthe data from searchlight traps with the data obtained by vertical-looking radar, included condition ofovarian development, ground and aerial meteorological data from weather station or internet, and pestinformation from the National Agro-Technical Extension and Service Centre. Then we integratedapplication of the wind field analysis, trajectory simulation and geographic information system softwaresuch as GrADS, HYSPLIT and ArcMap to analyze these data and investigate the influence of variousmeteorological factors on the population dynamics and the relationship between population fluctuationsand wind field. Also, this study explored the possibility of be migrantion of P. lepigone, and its sourceplace based on the data mentioned above. The results were as follows:
1. this newly designed aerial trap for high-flying insects could function when wind speed is notstrong enough to open the tow net, and is able to prevent the catches from escaping. Aerial samplingtests showed that this aerial trap could catch many individuals from7orders such as Hemiptera,Lepidoptera, Coleoptera etc. at the height of200~400m above the ground level. For the migratory pestinsects including L. sticticalis, M. separatar, Helicoverpa armigera Hubner, that were trapped in aerialsamples, the catch sizes in searchlight trap were consistently in peaked periods, which confirmed thatsearch light trap could be a practicable tool assisting radar in identification of the high-flying targets.
2. Dominant species of Orthoptera and Coleoptera only appeared in some time-sections at night,but Sympetrum Croceolum and Chrysopa formosa from Neuroptera, L. sticticalis, Pyrausta nubilalis,Hymenia recurvalis, Plutella xyllostella and many noctuids from Lepidoptera could be trapped duringthe whole night. Dynamic rhythm of catch sizes differed in different species. Distant migratory behaviorwas one of the most important factor the dynamic of specific species. When limiting the function timefor only1night, the trajectory simulation showed that the results obtained as parameterized withdifferent starting time involved more insect source place and possible landing sites, not simply thetruncated distance as generated by the traditional methods. Howerver, source places obtained by bothmethods tend to be similar with the extension of the function time.
3. In2012, the minimum temperature in Yanqing of Beijing was higher than the supercooling point(-25.35℃) of the mature larvae with cocoon. With other evidences taken into consideration, theresultsindicate that the mature larvae with cocoon of P. lepigone could therefore overwinter in YanqingBeijing. Monitoring data confirm that P. lepigone has occurred4generations per year in LuanchengHebei province and3generations per year in Yanqing Beijing. the total amount of trapped males wassignificantly higher than that of female in both sites. Nights when sex ratio of P. lepigone wassignificantly smaller than or close to1accounted for58%and30%in Yanqing and Luanchengrespectively. Variation in population sizes influences the sex ratio that changes in same trend for bothsites, that is, the sex ratios were significantly higher than1when population size rise, but close to1orsignificantly smaller than1in the the steady periods. The first generation of trapped P. lepigone adultsdid not form the normal distribution in Yanqing, and two obvious peaks appeared in the this generation.Variations in population sizes were related to the wind field at925hPa. With comprehensiveconsideration of the synchronous spurts in other light traps around Beijing, we conclude that the trappedfirst generation was composed not only the local individuals, but also the migrants or dispersers fromHebei and Tianjin，therefore, P. lepigone is a facutrative migratory species.
4. Indoors feeding observation showed that there were no significantly differences in the totaldevelopment duration between sexes. Tests on potential flight ability indicated that the males are ofmore powerful flight ability. So, total development duration and flight ablilty may lead to the changes insex ratioes. In the regions where first generation occurred, temperature and precipitation in March andApril had heavy influences on the occurrence of A. ypsilon, and it seemed that higher temperature andmore rains benefit the development of A. ypsilon. In the late June, the observation with entomologicalradar confirmed that most moths of A. ypsilon generally flew below500m, ant that the biggest densitywas at height of300~400m. The observation that moths of A. ypsilon can fly during the whole nightwere approved by both radar observation and timely samples of the catches from searchlight trap.Migrants of A. ypsilon could be windborne in that every peaks in trapped moths were alwaysaccompanied with favorable wind field and velocity, furthermore, the northern cyclones frontal weatherand heavy rainfall would cause massive landing.
5. In2008~2012, total trapped moths of armyworm were5105,3981,11385and50584,respectively, among which a significantly difference was identified. For armyworm, the biggest catchsizes was in2012and more than7300moths were trapped at one night which had nerver occurredbefore since this monitoring station has functioned. In conclusion, favorable weather condition were themain factors leading to the outbreak of larvae of3rd generation. Taken the relationship between catchsizes and wind fields, most of trapped moths were thought to be immigrants from surrounding regions.Therefore, appropriate weather condition for migrants in July promoted the outbreak of the3rd-generation larvae in2012. In North China, the outbreaks were mainly the result of the jointreproduction of immigrants and the local adults under the favorable environment condition and littleconcerned with adult population in Northeast China. For northeast China, the source moths of3rd-generation larvae in Northeast China were mainly from the local, because the local2nd-generationmoths in Northeast China failed to immigrate into the North China and had to stay in the natal areas ormake short southern migration due to westerly flow and rainfall in late July. The southward 2nd-generation moths in Heilongjiang mainly landed at the junction of Heilongjiang and Jilin provincesas a result of wind shear associated with heavy precipitation, lead to the serious outbreak at possiblelanding places, together with the immigrants from the North China. During the return migration seasonof3rd-generation adults, considering the influence of the terrain and monsoon in Northeast and NorthChina, we determined that only on some occasion the wind field can carry moths into North China fromNortheast China. In addition, almost all fall crops were in harvest at that time. Combining theseconsiderations together, we forecasted there was little chance for4th-generation larvae causing heavydamage to crop, which was proved to be true by the actual occurrence of4th larvae.
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