白蜡虫优势寄生蜂行为学特征研究
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摘要
在白蜡生产系统中,危害白蜡虫及其寄主植物的害虫隶属于4目13科28种。白蜡虫寄生蜂有14种,隶属于5科,白蜡虫天敌是影响白蜡虫种虫繁育和进行白蜡生产的主要因素之一。白蜡虫寄生蜂不仅寄生于白蜡虫,而且还寄生于其它农林害虫。调查发现在中国传统白蜡虫种虫产区和白蜡产区,没有重大农林害虫发生。这一现象表明,在培育白蜡虫生产白蜡的同时,能繁殖出控制农林害虫的自然天敌。通过培育白蜡虫繁育寄生蜂,利用白蜡虫丰富的天敌资源和天敌寄生的广谱性,研究白蜡虫寄生蜂与寄主之间的信息传递机制、捕捉寄主能力,探求白蜡虫寄生蜂寻找寄主的传递机制和基本规律,以期为白蜡虫寄生蜂的综合控制、利用提供必要的科学指导,一方面通过控制寄生蜂提高虫、蜡的产量和质量,另一方面通过繁育白蜡虫,培育利用寄生蜂防治农林害虫,为构建农林复合生物防治新模式奠定理论基础。研究白蜡虫寄生蜂与寄主之间的信息传递机制,不仅在科学理论上有指导意义,而且在生产实践中更具有重要应用价值。
本论文“白蜡虫优势寄生蜂行为学特征研究”运用化学生态学原理,采用活体植株动态顶空套袋密闭循环式高效吸附捕集法,热脱附-气相色谱/质谱(TCT-GC/MS)联用技术分析了挥发性物质的化学成分组成及变化状况,为进一步合理开发利用白蜡资源提供新的科学依据和应用基础。利用白蜡虫寄生蜂对颜色的趋性,构建了白蜡虫及其优势寄生蜂的时间--空间生态位,论述了白蜡虫3种优势寄生蜂对白蜡虫的跟随作用和寄生效能,以期为白蜡虫寄生蜂的综合评价提供科学理论依据。
寄生蜂产卵器等外生殖器的形态结构特征决定了寄生危害白蜡虫虫期和种群动态发生消长规律。白蜡虫寄生蜂共寄生现象明显。白蜡虫阔柄跳小蜂♀﹕♂性比10.47﹕1,其产卵包括寻找寄主、刺孔、产卵和梳理4个过程。通常每雌蜂在一个寄主内只产卵1粒,极个别产卵2粒。不同浓度的蜂蜜水等补充营养均可延长白蜡虫阔柄跳小蜂雌、雄蜂寿命。白蜡虫阔柄跳小蜂对白蜡虫雌成虫和附有雌成虫的寄主枝条具正趋性,对蜡花产生忌避性。白蜡虫花翅跳小蜂对蜡被、蜡花和有虫枝条表现出强烈的趋性。
通过对白蜡虫及其3种优势寄生蜂的时空生态位研究表明,白蜡虫花翅跳小蜂与白蜡虫阔柄跳小蜂在时间生态位、空间生态位和时间-空间生态位上均有较大的重叠和比例相似性,在资源利用上存在明显的种间竞争。白蜡虫花翅跳小蜂在时间、空间和时间-空间一维、二维生态位宽度、生态位重叠及生态位比例相似性均比白蜡虫阔柄跳小蜂和中华花翅跳小蜂大,说明白蜡虫花翅跳小蜂与白蜡虫在时间上的同步性和在空间上的同域性较强。白蜡虫花翅跳小蜂在时间、空间和时间-空间二维生态位上与白蜡虫有较明显的跟随现象,表明这种寄生蜂对白蜡虫的寄生作用较大,在利用时间、空间资源上占有优势。
通过对白蜡虫寄生蜂对颜色的趋性与日活动规律及迁飞能力与活动范围的研究表明,白蜡虫寄生蜂对黄色有明显的趋向性,对颜色的选择性依次为黄色>蓝色>蓝绿色>绿色>白色>灰褐色>红色>黑色。白蜡虫寄生蜂主要在白蜡园寄主林中冠层与下冠层之间范围活动,与白蜡虫在寄主植物上的分布密切相关。在一天中的不同时段里,白蜡虫寄生蜂在6﹕00～9﹕00 am有一个活动高峰期,在15﹕00～18﹕00 pm出现一个次高峰期。白蜡虫寄生蜂在晴天活动较活跃,在阴天和雨天活动较少。白蜡虫优势寄生蜂的垂直飞行能力弱于其它寄生蜂,白蜡虫寄生蜂的垂直飞行能力有限,一般不超过树梢2m,与寄生蜂本身体小而轻及风速、风向、光照等有关。白蜡虫寄生蜂有一定的水平飞行能力,迁飞距离在60m左右。
通过对白蜡生产体系挥发物的采集、检测及白蜡虫寄生蜂与寄主之间的信息传递物质分析表明,白蜡虫寄主植物女贞枝条挥发物成分共检测出14种化合物,其总含量达88.29%,主要成分为烷烃类、烯烃类、羧酸类、酮类等化合物。附吊糖期白蜡虫雌成虫的女贞枝条挥发物成分共检测出15种化合物,其挥发物总含量达88.81%,与女贞枝条挥发物接近,主要成分为烷烃类、烯烃类、萜烯类、酮类等化合物。成熟白蜡虫雌成虫挥发物成分共检测出8种化合物,化合物种类明显低于女贞枝条与附吊糖期白蜡虫雌成虫的女贞枝条挥发物,但其挥发物总含量高达99.29%,明显高于女贞枝条与附吊糖期白蜡虫雌成虫的女贞枝条挥发物含量,表明成熟白蜡虫雌成虫挥发物成分较单一、稳定,主要成分为酯类、烷烃类等化合物。在田间通过不同挥发物对白蜡虫寄生蜂的引诱效果检验表明,烷烃类、醇类、羧酸类、酯类、萜烯类挥发物能不同程度地诱集白蜡虫3种优势寄生蜂及其它寄生蜂混合种群。因此,白蜡虫寄生蜂对寄主与寄主植物的挥发物具有趋性,且对0.1%浓度的混合挥发物及单物质挥发物趋性明显,表明白蜡虫寄生蜂与寄主之间存在信息传递,主要通过挥发物气味寻找寄主。
In the white wax production systems, pest insects of wax worms and their host plants belong to 28 species of 13 families of 4 orders. Predators of white-wax insect were one of the main factors to affect the breeding of insect species and wax production. The basic research on identification, occurrence rule and damage survey of parasitic wasps, belonging to 14 species of 5 families, have been done in this study and pest control gains some efficiency by using sets cage stocking of Ericerus pela (Chavannes). Not only on wax worms, the wasps but also are parasitic on other pests. It is surveyed that there is no occurrence of major pests in traditional areas of white wax production in China. By this phenomenon, it is indicated that the production of white wax scale also could breed natural enemies to control pests in agriculture and forest. By cultivating wax insect to breed parasitic wasps and using the abundant resources parasitic of natural enemies in broad scope host, the information transmission mechanism between the wasps and the host, the host ability to capture were studied to explore the basic rules of transfer mechanism in wasps finding hosts, which is expected to provide the necessary scientific guidance on control and utilization of parasitoids of white-wax insect. On the one hand, the wax production and quality were increased by controlling insect parasitoids. On the other hand, the theoretical basis of new models for biological control in agro-forestry was built through the cultivation and utilization of parasitic wasps to control pests. The study of information transfer mechanisms between parasitoids and it host white-wax insect not only is significance in theoretical science, but also benefits the practice of production.
In this paper, Studies on the behavior characteristic of dominant parasitic wasps of Ericerus pela (Chavannes), by the theory of chemical ecology, Using the Capture method of dynamic head-space collection from living plant in airtight and circulatory bagging, Techniques of TCT-GC/MS and Computer library search, the chemical compositions and changes of volatile in white wax production system were revealed objectively and scientifically, which will afford the new scientific evidence and application base for further rational development and utilization of white wax resources. Using the color trend of parasitoids, the temporal-spatial niches of E. pela (Chavannes) and its dominant parasitoids were constructed and the follow function and parasitic effectiveness of three dominant parasitoids were also discussed in order to accumulate the scientific evidence for comprehensive evaluation on the parasitic wasps of E. pela (Chavannes).
The stage of parasitic damage occurring in white-wax insects and dynamics rules of population are determined by morphological structure of parasitoid ovipositor of parasitic wasps. The symparasitism in E. pela (Chavannes) was found apparently in this study. The sex ratio (female: male) of Metaphycus ericeri is 10.47:1. Its ovipositing includes the process of host searching, holing, laying eggs and carding. Usually, the females lay only one egg or very few two eggs in a host. Different concentrations of honey water may both extend the longevity of parasitoids with nutritional supplements. Metaphycus ericeri express positive trends to female adults of E. pela (Chavannes) and host branches with female adults while shows the repellent nature to wax flower. Microterys sinicus shows strong positive trends to wax cover, wax flower and host branches with female adults.
One-dimensional niche breadth, niche overlap and percentage similarity of E. pela and its dominant parasitoid wasps (Microterys ericeri Ishii, M. sinicus Jiang, and Metaphycus ericeri Xu et Jiang), were analysed. Temporal niche breadths of E. pela and its three dominant parasitoids were E. pela > Metaphycus ericeri > M. ericeri > M. sinicus and spatial niche breadths were E. pela > M. ericeri > Metaphycus ericeri > M. sinicus. The niche overlap of the three dominant parasitoids were M. ericeri > M. sinicus > Metaphycus ericeri; the percentage similarities were M. ericeri > Metaphycus ericeri > M. sinicus. This suggests that M. ericeri and E. pela possesses the most similarity in the resources series, strong synchronism and sympatry, and the maximum resource utilization. M. ericeri and Metaphycus ericeri had a large overlap in temporal-spatial niches and proportion similarity. Interspecific competition for resource utilization between these species was obvious. E. pela (Chavannes)is an abundant and stable food of many parasitoids because of its large population and for longer periods.
In the studies of color trend, daily activities rules, migration capacity and range of parasitic wasps, the results showed that parasitic wasps of E. pela (Chavannes) obvious owned the trend of yellow color and the sequence of color selectivity were yellow> blue> blue-green> green> white> brown> red> black. The parasitic wasps mainly occurred in the middle and under layer of host plant garden, which closely related to the white wax worms distributing in host plants. In different times of the day, an activity peak of parasitic wasps appeared in 6:00 ~ 9:00 am with a second peak in the 15:00 ~ 18:00 pm. And parasitic wasps of E. pela (Chavannes) were more active in the sunny day than in the cloudy and rainy days. Vertical flight ability of dominant parasitoids of E. pela (Chavannes) was weaker than other parasitic wasps, generally no more than 2 meters beyond the treetop, which closely related to the small, light body of parasitic wasps and wind speed, direction and sunlight. The parasitic wasps have a certain ability of horizontal flight with migration distance limited within 60 meters.
Through volatile collecting, testing in wax production systems and analysis material of information transfer the between E. pela (Chavannes) and parasitic wasps, 14 compounds were detected in volatiles from host t branches, accounting for 88.29 percent of the total content. The main components include alkanes, alkenes, carboxylic acids and ketones. In the period of sugar hanging attached, 15 compounds were detected in volatiles from female adults and host branches. The volatiles were mainly composed of alkanes, alkenes, terpenes and ketones, taking 88.81% of total content and nearly as volatiles from privet branches. While only 8 compounds were detected from mature female adult of E. pela (Chavannes). The types of compound in the volatiles were significantly lower than those in volatile from host branches or from female adults and host branches in the period of sugar hanging attached, But the content of volatile accounted for 99.29% of total, significantly higher than the content of host branches or from female adults and host branches in the period of sugar hanging attached, which showed the volatiles from matured female adults are more simplex and stable, mainly composed of esters and hydrocarbons. The test results of luring parasitic wasps by using different volatile compounds in the field show that alkanes, alcohols, carboxylic acids, esters and terpene volatiles can allure three dominant parasitoids and other mixed populations of parasitic wasps in varying levels. The growth period of E. pela (Chavannes) and its relationship to the occurrence of parasitic wasps should be given priority think to luring. There is some axis to volatile component between host plant and parasitic wasps of E. pela (Chavannes), depending on taste to look for their hosts.
本论文“白蜡虫优势寄生蜂行为学特征研究”运用化学生态学原理,采用活体植株动态顶空套袋密闭循环式高效吸附捕集法,热脱附-气相色谱/质谱(TCT-GC/MS)联用技术分析了挥发性物质的化学成分组成及变化状况,为进一步合理开发利用白蜡资源提供新的科学依据和应用基础。利用白蜡虫寄生蜂对颜色的趋性,构建了白蜡虫及其优势寄生蜂的时间--空间生态位,论述了白蜡虫3种优势寄生蜂对白蜡虫的跟随作用和寄生效能,以期为白蜡虫寄生蜂的综合评价提供科学理论依据。
寄生蜂产卵器等外生殖器的形态结构特征决定了寄生危害白蜡虫虫期和种群动态发生消长规律。白蜡虫寄生蜂共寄生现象明显。白蜡虫阔柄跳小蜂♀﹕♂性比10.47﹕1,其产卵包括寻找寄主、刺孔、产卵和梳理4个过程。通常每雌蜂在一个寄主内只产卵1粒,极个别产卵2粒。不同浓度的蜂蜜水等补充营养均可延长白蜡虫阔柄跳小蜂雌、雄蜂寿命。白蜡虫阔柄跳小蜂对白蜡虫雌成虫和附有雌成虫的寄主枝条具正趋性,对蜡花产生忌避性。白蜡虫花翅跳小蜂对蜡被、蜡花和有虫枝条表现出强烈的趋性。
通过对白蜡虫及其3种优势寄生蜂的时空生态位研究表明,白蜡虫花翅跳小蜂与白蜡虫阔柄跳小蜂在时间生态位、空间生态位和时间-空间生态位上均有较大的重叠和比例相似性,在资源利用上存在明显的种间竞争。白蜡虫花翅跳小蜂在时间、空间和时间-空间一维、二维生态位宽度、生态位重叠及生态位比例相似性均比白蜡虫阔柄跳小蜂和中华花翅跳小蜂大,说明白蜡虫花翅跳小蜂与白蜡虫在时间上的同步性和在空间上的同域性较强。白蜡虫花翅跳小蜂在时间、空间和时间-空间二维生态位上与白蜡虫有较明显的跟随现象,表明这种寄生蜂对白蜡虫的寄生作用较大,在利用时间、空间资源上占有优势。
通过对白蜡虫寄生蜂对颜色的趋性与日活动规律及迁飞能力与活动范围的研究表明,白蜡虫寄生蜂对黄色有明显的趋向性,对颜色的选择性依次为黄色>蓝色>蓝绿色>绿色>白色>灰褐色>红色>黑色。白蜡虫寄生蜂主要在白蜡园寄主林中冠层与下冠层之间范围活动,与白蜡虫在寄主植物上的分布密切相关。在一天中的不同时段里,白蜡虫寄生蜂在6﹕00～9﹕00 am有一个活动高峰期,在15﹕00～18﹕00 pm出现一个次高峰期。白蜡虫寄生蜂在晴天活动较活跃,在阴天和雨天活动较少。白蜡虫优势寄生蜂的垂直飞行能力弱于其它寄生蜂,白蜡虫寄生蜂的垂直飞行能力有限,一般不超过树梢2m,与寄生蜂本身体小而轻及风速、风向、光照等有关。白蜡虫寄生蜂有一定的水平飞行能力,迁飞距离在60m左右。
通过对白蜡生产体系挥发物的采集、检测及白蜡虫寄生蜂与寄主之间的信息传递物质分析表明,白蜡虫寄主植物女贞枝条挥发物成分共检测出14种化合物,其总含量达88.29%,主要成分为烷烃类、烯烃类、羧酸类、酮类等化合物。附吊糖期白蜡虫雌成虫的女贞枝条挥发物成分共检测出15种化合物,其挥发物总含量达88.81%,与女贞枝条挥发物接近,主要成分为烷烃类、烯烃类、萜烯类、酮类等化合物。成熟白蜡虫雌成虫挥发物成分共检测出8种化合物,化合物种类明显低于女贞枝条与附吊糖期白蜡虫雌成虫的女贞枝条挥发物,但其挥发物总含量高达99.29%,明显高于女贞枝条与附吊糖期白蜡虫雌成虫的女贞枝条挥发物含量,表明成熟白蜡虫雌成虫挥发物成分较单一、稳定,主要成分为酯类、烷烃类等化合物。在田间通过不同挥发物对白蜡虫寄生蜂的引诱效果检验表明,烷烃类、醇类、羧酸类、酯类、萜烯类挥发物能不同程度地诱集白蜡虫3种优势寄生蜂及其它寄生蜂混合种群。因此,白蜡虫寄生蜂对寄主与寄主植物的挥发物具有趋性,且对0.1%浓度的混合挥发物及单物质挥发物趋性明显,表明白蜡虫寄生蜂与寄主之间存在信息传递,主要通过挥发物气味寻找寄主。
In the white wax production systems, pest insects of wax worms and their host plants belong to 28 species of 13 families of 4 orders. Predators of white-wax insect were one of the main factors to affect the breeding of insect species and wax production. The basic research on identification, occurrence rule and damage survey of parasitic wasps, belonging to 14 species of 5 families, have been done in this study and pest control gains some efficiency by using sets cage stocking of Ericerus pela (Chavannes). Not only on wax worms, the wasps but also are parasitic on other pests. It is surveyed that there is no occurrence of major pests in traditional areas of white wax production in China. By this phenomenon, it is indicated that the production of white wax scale also could breed natural enemies to control pests in agriculture and forest. By cultivating wax insect to breed parasitic wasps and using the abundant resources parasitic of natural enemies in broad scope host, the information transmission mechanism between the wasps and the host, the host ability to capture were studied to explore the basic rules of transfer mechanism in wasps finding hosts, which is expected to provide the necessary scientific guidance on control and utilization of parasitoids of white-wax insect. On the one hand, the wax production and quality were increased by controlling insect parasitoids. On the other hand, the theoretical basis of new models for biological control in agro-forestry was built through the cultivation and utilization of parasitic wasps to control pests. The study of information transfer mechanisms between parasitoids and it host white-wax insect not only is significance in theoretical science, but also benefits the practice of production.
In this paper, Studies on the behavior characteristic of dominant parasitic wasps of Ericerus pela (Chavannes), by the theory of chemical ecology, Using the Capture method of dynamic head-space collection from living plant in airtight and circulatory bagging, Techniques of TCT-GC/MS and Computer library search, the chemical compositions and changes of volatile in white wax production system were revealed objectively and scientifically, which will afford the new scientific evidence and application base for further rational development and utilization of white wax resources. Using the color trend of parasitoids, the temporal-spatial niches of E. pela (Chavannes) and its dominant parasitoids were constructed and the follow function and parasitic effectiveness of three dominant parasitoids were also discussed in order to accumulate the scientific evidence for comprehensive evaluation on the parasitic wasps of E. pela (Chavannes).
The stage of parasitic damage occurring in white-wax insects and dynamics rules of population are determined by morphological structure of parasitoid ovipositor of parasitic wasps. The symparasitism in E. pela (Chavannes) was found apparently in this study. The sex ratio (female: male) of Metaphycus ericeri is 10.47:1. Its ovipositing includes the process of host searching, holing, laying eggs and carding. Usually, the females lay only one egg or very few two eggs in a host. Different concentrations of honey water may both extend the longevity of parasitoids with nutritional supplements. Metaphycus ericeri express positive trends to female adults of E. pela (Chavannes) and host branches with female adults while shows the repellent nature to wax flower. Microterys sinicus shows strong positive trends to wax cover, wax flower and host branches with female adults.
One-dimensional niche breadth, niche overlap and percentage similarity of E. pela and its dominant parasitoid wasps (Microterys ericeri Ishii, M. sinicus Jiang, and Metaphycus ericeri Xu et Jiang), were analysed. Temporal niche breadths of E. pela and its three dominant parasitoids were E. pela > Metaphycus ericeri > M. ericeri > M. sinicus and spatial niche breadths were E. pela > M. ericeri > Metaphycus ericeri > M. sinicus. The niche overlap of the three dominant parasitoids were M. ericeri > M. sinicus > Metaphycus ericeri; the percentage similarities were M. ericeri > Metaphycus ericeri > M. sinicus. This suggests that M. ericeri and E. pela possesses the most similarity in the resources series, strong synchronism and sympatry, and the maximum resource utilization. M. ericeri and Metaphycus ericeri had a large overlap in temporal-spatial niches and proportion similarity. Interspecific competition for resource utilization between these species was obvious. E. pela (Chavannes)is an abundant and stable food of many parasitoids because of its large population and for longer periods.
In the studies of color trend, daily activities rules, migration capacity and range of parasitic wasps, the results showed that parasitic wasps of E. pela (Chavannes) obvious owned the trend of yellow color and the sequence of color selectivity were yellow> blue> blue-green> green> white> brown> red> black. The parasitic wasps mainly occurred in the middle and under layer of host plant garden, which closely related to the white wax worms distributing in host plants. In different times of the day, an activity peak of parasitic wasps appeared in 6:00 ~ 9:00 am with a second peak in the 15:00 ~ 18:00 pm. And parasitic wasps of E. pela (Chavannes) were more active in the sunny day than in the cloudy and rainy days. Vertical flight ability of dominant parasitoids of E. pela (Chavannes) was weaker than other parasitic wasps, generally no more than 2 meters beyond the treetop, which closely related to the small, light body of parasitic wasps and wind speed, direction and sunlight. The parasitic wasps have a certain ability of horizontal flight with migration distance limited within 60 meters.
Through volatile collecting, testing in wax production systems and analysis material of information transfer the between E. pela (Chavannes) and parasitic wasps, 14 compounds were detected in volatiles from host t branches, accounting for 88.29 percent of the total content. The main components include alkanes, alkenes, carboxylic acids and ketones. In the period of sugar hanging attached, 15 compounds were detected in volatiles from female adults and host branches. The volatiles were mainly composed of alkanes, alkenes, terpenes and ketones, taking 88.81% of total content and nearly as volatiles from privet branches. While only 8 compounds were detected from mature female adult of E. pela (Chavannes). The types of compound in the volatiles were significantly lower than those in volatile from host branches or from female adults and host branches in the period of sugar hanging attached, But the content of volatile accounted for 99.29% of total, significantly higher than the content of host branches or from female adults and host branches in the period of sugar hanging attached, which showed the volatiles from matured female adults are more simplex and stable, mainly composed of esters and hydrocarbons. The test results of luring parasitic wasps by using different volatile compounds in the field show that alkanes, alcohols, carboxylic acids, esters and terpene volatiles can allure three dominant parasitoids and other mixed populations of parasitic wasps in varying levels. The growth period of E. pela (Chavannes) and its relationship to the occurrence of parasitic wasps should be given priority think to luring. There is some axis to volatile component between host plant and parasitic wasps of E. pela (Chavannes), depending on taste to look for their hosts.
引文
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