A test of temporal and spatial density dependence in the parasitism rates of introduced parasitoids on host, the arrowhead scale (Unaspis yanonensis) in stable host–parasitoids system

Abstract: In 1980, two species of parasitoid wasps (Aphytis yanonensis DeBach et Rosen and Coccobius fulvus Compere et Annecke) were introduced to Japan from China as biological control agents to combat the arrowhead scale (Unaspis yanonensis Kuwana). These introductions represent one of the most successful projects in the history of biological control in Japan. To examine whether density dependent parasitism was inevitable for success of biological control, we tried to detect temporal and spatial density dependence in parasitism rates using time‐series data of scale density, as well as parasitism, over a 16‐year period. The work was conducted in a Satsuma mandarin orange (Citrus unshiu Marc.) orchard in which we previously demonstrated that the system appeared to have stabilized after a decline in scale density following the introduction of the parasitoids. Earlier work also indicated that C. fulvus contributes most to the reduction in, and the stability of, scale density. In this study, we examined: (1) the relationship, on a whole‐orchard basis, between scale density and the rates of parasitism by A. yanonensis, C. fulvus, and a combination of the two species; (2) whether parasitism was positively correlated to scale density on a single‐tree basis among generations and (3) whether spatial density dependence was detectable within generations on an individual‐tree basis. Parasitism by A. yanonensis was temporally density‐dependent on scale population density at the whole‐orchard level, while parasitism by C. fulvus was not. Parasitism by A. yanonensis or by C. fulvus was rarely positively correlated to scale density at the single‐tree level, and spatial density‐dependence was hardly detected at all at this level. Most analyses of combined parasitism rates were similar to rates of parasitism by C. fulvus alone. Contrary to conventional wisdom of biological control theory, this study demonstrates that density dependence is not necessarily detected, even in a system in which a natural enemy has long held pest density stable at low levels.     

Is one parasitoid enough? A test comparing one with a pair of parasitoid species in the biological control of arrowhead scales

In classic biological control using natural enemies, the question of whether a single species or multiple species should be introduced has been a matter of debate. The introduction of two parasitoids, Aphytis yanonensis and Coccobius fulvus (Hymenoptera: Aphelinidae), to control the arrowhead scale, Unaspis yanonensis (Hemoptera: Diaspididae), which is a serious pest in Japanese citrus orchards, has been one of the most successful biological control projects in Japan. The success of this program may be explained by two alternative hypotheses: (1) the parasitoid species work complementarily, or (2) only one of them plays a major role. To test which hypothesis is applicable to this host-parasitoid system, we conducted caging experiments and observed temporal changes in the proportion of the parasitisms and the densities of arrowhead scales enclosed with one of the following combinations of parasitoids: (1) A. yanonensis and C. fulvus together, (2) A. yanonensis alone, (3) C. fulvus alone, or (4) neither parasitoid. Parasitisms in the cohorts with A. yanonensis and C. fulvus together and C. fulvus alone rapidly increased to approximately 70%; parasitism with A. yanonensis alone also increased slightly, although it remained consistently lower that those with A. yanonensis and C. fulvus together and C. fulvus. At the end of the experiment, parasitisms with A. yanonensis and C. fulvus together and C. fulvus alone were significantly higher than that with A. yanonensis alone. Parasitism by C. fulvus constituted most of (74%) the parasitism in the cohort with A. yanonensis and C. fulvus together. Further, only C. fulvus suppressed the population growth rates of scales significantly. These results suggest that C. fulvus alone successfully suppresses scale populations as efficiently as both species together do.     

卫矛矢尖蚧的研究

南京地区卫矛矢尖蚧unaspis euonymi Comstock一年发生3代．第一至三代若虫孵化盛期分别为5月中旬、6月下旬和8月中旬．各代雌成虫始见期分别为5月下旬,6月下旬和8月下旬．11月下旬开始以受精雌成虫在寄主上越冬．主要天敌有红点唇瓢虫Chilocorus kuwanae Silvestri、日本方头甲Cybocephalus nipponicus Enduody-Younga,黄蚜小蜂Aphytis sp．等．药剂防治用机油乳剂。冬季1：30-50倍液(防治越冬代雌成虫),夏季1：50-100倍液;40％氧化乐果1：1000倍液、25％亚胺硫磷乳剂1：800倍液等喷洒1至3代盛孵若虫,杀虫效果均可达95％以上．     

20种杀虫剂对柑橘矢尖蚧的室内毒力测定

柑橘矢尖蚧Unaspis yanonensis(Kuwana)是危害柑橘的重要害虫之一。为了筛选高毒有机磷农药的替代物,在室内条件下采用浸渍法分别测定20种杀虫剂原药对柑橘矢尖蚧1龄若虫的毒力。结果表明:供试杀虫剂对柑橘矢尖蚧的毒力大小依次为:甲氨基阿维菌素苯甲酸盐>阿维菌素>硝虫硫磷>噻嗪酮>杀扑磷>吡虫啉>乙酰甲胺磷>毒死蜱>喹硫磷>亚胺硫磷>稻丰散>敌百虫>残杀威>氟氯氰菊酯>甲氰菊酯>甲氧虫酰肼>苯氧威>高效氯氰菊酯>石硫合剂>矿物油。     

矢尖蚧发生动态及其寄生性天敌

柑桔矢尖蚧在福建闽北地区一年发生3代。以雌成虫和部分若虫及少数雄蛹越冬．一龄幼蚧盛期,第一代为4月中,下旬。第二代7月上．中旬,第三代9月中旬前后。二龄幼蚧盛期．第一代为5月上、中旬,第二代7月下旬,和第三代9月下旬．后期世代重叠．福建已知寄生蜂6种．其中一种为重寄生蜂,以矢尖蚧黄蚜小蜂为优势种．寄生生蜂自然寄生率最高达49.1%。     

Fertility schedules ofunaspis yanonensis (Hemiptera: Diaspididae) in relation to daily temperature

Fertility schedules of Unaspis yanonensis on citrus leaves were investigated from 1962 to 1971 for the first generation and from 1964 to 1971 for the second generation, at the Okitsu Branch of the Fruit Tree Research Station in Shizuoka Prefecture. The fertility curves were commonly characterized by a first major peak and a few subsequent minor peaks, although their shapes differed from year to year. Models estimating these fertility curves of each generation were constructed in relation to the fluctuations in average daily temperature, on the assumption that the curve was composed of two peaks, i.e., bimodal. Estimated curves were well fitted to observed ones.     

二双斑唇瓢虫对矢尖蚧的捕食作用

研究了二双斑唇瓢虫（Chilocorus bijugus Mulsant)对矢尖蚧（Unaspis yanonensis Kuwana)的捕食作用,结果表明,瓢虫雌成虫对矢尖蚧各虫态的功能反应呈Holling II型,瓢虫对矢尖蚧的捕食效应随捕食者个体间干扰作用的增加而下降,捕食作用率（E）随着瓢虫数增加呈幂函数下降曲线,温度对瓢虫的捕食效应具有显著的影响,寻找效率（a)和处置时间（Th)与温度之间呈二次函数关系,猎物密度对瓢虫生殖力的影响呈Logistic曲线。     

Cascading effects of a specialist parasitoid on plant biomass in a <Emphasis Type="Italic">Citrus</Emphasis> agroecosystem

We studied a specialist parasitoid (Coccobius fulvus Compere et Annecke; Hymenoptera: Aphelinidae), its host (the arrowhead scale, Unaspis yanonensis Kuwana; Hemiptera: Diaspididae) and the host plant (Citrus unshiu Marc; Rutaceae) to examine the indirect effects, via host–parasitoid interactions, of the parasitoid on plant biomass. We compared plant biomass and herbivore abundance in a system of two trophic levels (plants and herbivores) with a system of three trophic levels (plants, herbivores and parasitoids) using enclosure experiments in an agricultural setting. Each of eight young Citrus trees was infested with 40 scales and placed in an enclosure. We introduced three female parasitoids into half of the enclosures and monitored temporal changes in scale density and cumulative parasitism for the subsequent 11months. Plant biomass was then compared between treatment groups (parasitoids added) and controls (parasitoids excluded). During the experiment, cumulative parasitism increased rapidly in the parasitoid-addition enclosures to a maximum of 89%, and the number of live scales in the control enclosures was approximately 10-fold that in the treatment enclosures. At the end of the experiment, plant biomass was threefold higher in the parasitoid-addition enclosures than in the control enclosures. These results have two implications for terrestrial communities. First, specialist parasitoids, which are the principal natural enemies of most herbivorous insects, can trigger trophic cascades in the same way that generalist predators can. Second, cascading effects can be detected by observing changes in plant biomass. The latter finding is contrary to recent conclusions about top-down cascades (i.e. that trophic cascades are less likely to be observed when plant biomass, rather than plant damage, is considered as the plant-response variable).     

红点唇瓢虫对矢尖蚧的捕食作用(英文)

红点唇瓢虫（Chilocoruskuwanae）的产卵雌虫对矢尖蚧（Unaspisyanonensis）的捕食量显著大于雄性成虫。在25℃下,雌成虫日平均捕食矢尖蚧雌成蚧42．7头,而雄成虫的日捕食量仅为22．3头。但48h饥饿处理可消除此差异。要使雌成虫产卵,每天必须至少捕食15头矢尖蚧的雌成蚧。红点唇瓢虫成虫对矢尖蚧各虫态的功能反应均为HollingⅡ型。根据对雌成蚧的功能反应,在16－35℃范围内最大捕食量随温度的升高而升高,在37℃下则急剧下降。温度并不改变功能反应的类型。由红点唇＊虫成虫对雌成钛的捕食作用,估算得最低临界攻击温度、最佳攻击温度和最高临界攻击温度分别为10．58℃、31．53℃和3820℃。捕食空间增大或捕食者数量的增加都会使攻击率下降;但是捕食者密度变大时,捕食空间的影响将大大变小。红点唇瓢虫成虫对矢尖蚧的雄蛹表现出最大的猎物选择性。它们对矢尖蚧各虫态的捕食选择性依次为：雄蛹、2龄雄蚧、2龄雌蚧和1龄若虫。     

柑桔矢尖蚧生态学研究

1 996～ 1 999年 ,在浙江东南沿海自然变温下对矢尖蚧的发生历期与日平均气温的相关性、各虫态的发育起点温度和有效积温以及生命表等进行了系统的研究。第 1、2、3代雌性寿命分别为 1 0 9.5、1 1 6.2 (一年发生 2代的个体为2 5 6.5 d)和 2 60 .0 d左右 ;完成 1代所需历期分别为 70 .1± 3 .9、67.5± 9.8和 2 1 1 .4± 1 6.5 d。各虫态发生历期与日平均气温明显呈负相关。雌一龄、二龄、产卵前期和雄一龄、二龄、蛹 (包括预蛹 )的发育起点温度分别为 1 1 .5、9.6、8.7、1 4.7、1 3 .6和 1 3 .7℃ ,有效积温分别为 2 61 .8、2 83 .8、5 3 0 .4、1 65 .9、2 0 6.2和 94.0日度。第 1、2、3代每头雌成虫的产卵量分别为 93 .0± 42 .7、1 3 4.2± 65 .6和 1 96.3± 81 .9粒。第 2、3代存在世代重叠现象。第 1、2、3代的种群趋势指数避雨区分别为 2 2 .5 6、3 1 .1 7和 2 .48,露地区分别为 2 1 .0 9、2 9.76和 1 .96,表明在新区的种群自然增长能力很强。研究还揭示了在新区对矢尖蚧种群数量起控制作用的主要因子是自然死亡和失踪 ,因而必需从老发区引进、饲养释放和保护有效的寄生蜂等天敌。