Weak photoperiodic response facilitates the biological invasion of the harlequin ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae)

Photoperiodic regulation of reproductive diapause in two invasive and two native populations of Harmonia axyridis and in one native population of Harmonia yedoensis was investigated in laboratory at 20°C, five photoperiods (day length of 10, 12, 14, 16 and 18 h) and two diets: (i) eggs of the Angoumois grain moth Sitotroga cerealella and (ii) the green peach aphid, Myzus persicae. Laboratory strains originated from native populations of H. axyridis from Irkutsk province of Siberia and H. yedoensis from South Korea showed a strong photoperiodic response: under short photoperiods (10–14 h and 10–12 h for H. axyridis and H. yedoensis, correspondingly), all females which fed on eggs and most of those fed on aphids did not start to lay eggs during 40 days after emergence, while under long photoperiods, all females fed on aphids and most of those fed on eggs oviposited. The photoperiodic response of H. axyridis from South Korea was less strong: on the both diets, the range of the photoperiodic response (the difference in the proportion of ovipositing females between the treatments with long and short days) was ca 40%. In the European (Czech Republic) and in the Caucasian (Sochi region, Russia) invasive populations of H. axyridis, the photoperiodic response was very weak: the proportion of females that started oviposition (when fed on aphids) or at least reproductive maturation (when fed on eggs) during 40 days after emergence was close to 100%, independently of the photoperiodic conditions. Obviously, instead of a rapid micro‐evolutionary adaptation of the critical day length to a new climate, the invasive populations of the harlequin ladybird decrease their dependence on photoperiod and thus the weak photoperiodic response of SE Asian population of H. axyridis can be considered as a pre‐adaptation further developed during the invasion.     

不同色斑型异色瓢虫COⅠ和COⅡ基因序列系统进化分析

利用线粒体COⅠ和COⅡ基因序列分析法,研究13种色斑型异色瓢虫Harmonia axyridis Pallas的系统分化,计算核苷酸的使用频率,并构建分子系统树,以便更好地探索异色瓢虫色斑变异的规律。结果表明,在用于分析的COⅠ基因和COⅡ基因的序列中,共有31个变异位点,29个简约信息位点,A+T平均约占71.2%,大部分碱基改变为颠换。13种不同色斑型异色瓢虫的遗传距离为0.0021～0.0328,平均相似系数为0.0193。分别采用邻接法(NJ)、最大简约法(MP)、最小进化法(ME)构建系统发育树。分子系统树显示:各类群间遗传分化很小,鞘翅颜色相同的瓢虫均已较高的置信值聚在一起;双月斑变型和显明变种亲缘关系最近,重名变种为原始类群,分化较早,与其它12种不同色斑异色瓢虫亲缘关系较远。     

Differential sensitivity of an invasive and an indigenous ladybeetle to two reduced‐risk insecticides

The invasive multicoloured Asian lady beetle, Harmonia axyridis (Pallas), and the indigenous twelve spotted lady beetle, Coleomegilla maculata DeGeer (Col., Coccinellidae), are two important generalist predators commonly found in apple orchards in Quebec, Canada. Both species are exposed to two reduced‐risk insecticides, recently adopted to control codling moth, Cydia pomonella (L.) (Lep., Tortricidae) in south‐eastern Canada. Chlorantraniliprole (Altacor^® 35 WG), an anthranilic diamide insecticide, causes paralyses of the muscle cells by interfering with the insect ryanodine receptors, whereas novaluron (Rimon^® EC 10), a benzoylphenyl urea, inhibits the chitin synthesis. The aim of this study was to compare the sensitivity of both the invasive ladybeetle H. axyridis and the indigenous C. maculata to reduced‐risk insecticides through the assessment of lethal effect on eggs and larvae following topical contact, ingestion of treated prey and exposure to fresh residues, at field rates (50.75 g a.i./ha chlorantraniliprole and 100 g a.i./ha novaluron) in laboratory conditions. Eggs of both species were not affected. Following 6 days of residual contact, chlorantraniliprole and novaluron caused more than 98% mortality to larvae of both ladybeetle species. In topical contact and ingestion trials, chlorantraniliprole caused less than 18% mortality to larvae of the two species after 6 days following exposure. Novaluron had a drastically different impact on the two predators. It did not affect the indigenous C. maculata, whereas it killed 91% and 96% of H. axyridis individuals after 6 days, respectively, following topical contact and ingestion. These results illustrate a differential sensitivity to novaluron between two relatively close species (subfamily Coccinellinae), a potential impact on the invasion process by H. axyridis, and consequently on the ladybeetle assemblage in the field.     

Role of photoperiod in the development and reproduction of Harmonia axyridis (Coleoptera,Coccinellidae)

To improve the laboratory mass rearing and the winter greenhouse use of the multicoloured Asian lady beetle Harmonia axyridis (Pallas) (Coleoptera, Coccinellidae), the effects of photoperiod on the development and fecundity of H. axyridis were investigated at 20°C and photoperiods ranging from 0L:24D to 24L:0D. We foun'd photoperiod to significantly affect the pre-imaginal development and reproduction of H. axyridis. The development of the pre-imaginal stage was significantly faster at shorter photoperiods (0–14?h) than at longer ones (16–24?h). The proportion of females laying eggs within the first 30 days after eclosion, the mean number of eggs per ovipositing female, and the long axial length of the first oocyte were all lower at shorter photoperiods (6–12?h) than at longer ones (14–24?h). These results suggest that adult H. axyridis may encounter problems in preying and reproducing in the winter conditions of unheated and unilluminated greenhouses in areas with temperatures lower than 20°C and day lengths shorter than 12?h.     

异色瓢虫与隐斑瓢虫的区别及其色斑型和横脊的频率

异色瓢虫Harmonia axyridis(Pallas)和隐斑瓢虫H.yedoensis(Takizawa)是亲缘关系很近的姐妹种,在中国(北京到南岭,西至甘肃南部和西藏东部)、日本(北海道以南)和南韩等地同域分布,常常在松树等植物上共存。长期以来,隐斑瓢虫被认为是异色瓢虫的一个异名。有时从外形上很难区分这2种瓢虫,因此对于依据野外数据,分析异色瓢虫色斑型和鞘翅横脊的发生频率及小进化会产生一些错误。本文从没有隐斑瓢虫分布的东北地区的材料及其他数据,报道异色瓢虫的色斑型及鞘翅横脊发生频率。异色瓢虫的花斑型在我国东部地区(从东北至广东北部,西至甘肃和云南)发生率很低,而在新疆西北部发生率较高。鞘翅横脊的发生率从东北佳木斯的98.85%降低到云南大理的78.26%。本文列出了区分这2种瓢虫的形态特征及自然分布。如果鞘翅具横脊,则属于异色瓢虫,但如果标本来自2种瓢虫的共存区,鞘翅没有横脊,鞘翅的斑纹呈花斑型、四窗型或二窗型,则很难从外部形态上对它们进行鉴定。但这2种瓢虫的幼虫很容易区分。本文还提供了2种瓢虫的雄性外生殖器形态图、幼虫和成虫。     

Nitrogen fertiliser affects the functional response and prey consumption of Harmonia axyridis (Coleoptera: Coccinellidae) feeding on cereal aphids

Predator–prey interactions are influenced by nitrogen availability. Wheat (Triticum aestivum cv. Solstice) plants were provided with four levels of nitrogen and examined the responses of coccinellid predator, Harmonia axyridis to cereal aphids, Rhopalosiphum padi and Sitobion avenae. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. In no‐choice feeding trials, H. axyridis consumed more aphids on low fertilised plants, suggesting a compensatory consumption to overcome reduced biomass (lower aphid size). Total biomass devoured by H. axyridis on all nitrogen fertiliser treatments was not statistically different. Logistic regression analysis of the proportion of prey consumed demonstrated that all developmental stages (larval and adult) of H. axyridis exhibited the type II functional response on all nitrogen fertiliser treatments. The rate of successful search (a′) of third and fourth instars and adults were the same across all fertiliser treatments suggesting that nitrogen fertilisation did not affect a′. Maximum handling time for the first instars of H. axyridis on R. padi (3.81 h^?1) and S. avenae (4.59 h^?1) was on the highest nitrogen treatment while minimum handling time was for the adults of H. axyridis on R. padi (0.20 h^?1) and S. avenae (0.20 h^?1) on the lowest nitrogen treatment. Handling time varied at varying fertiliser treatments within all instars and affected the predator's efficiency. The functional response curve, rate of successful search and handling time provide the information needed to understand the predator–prey interaction between H. axyridis and these cereals aphids. This could lead to the development of a better strategy for the biological control of R. padi and S. avenae at any particular level of nitrogen fertiliser regime in the field crops.     

异色瓢虫幼虫的食物搜索行为

本试验采用Nakamuta(1982)装置研究两种光照条件下异色瓢虫幼虫的食物搜索行为,结果表明：(1)摄食刺激均能激发搜索行为由广域型转换为地域集中型;(2)摄食时间越长,地域集中型搜索时间(GUT)值越大;(3)摄食的最后一个食饵大小决定GUT的长短;(4)光照对搜索行为有影响;(5)饥饿度对GUT的长短有一定影响。     

异色瓢虫不同龄期幼虫踪迹对受孕雌虫产卵的影响

本文通过室内试验研究了异色瓢虫Harmonia axyridis (Pallas)不同龄期幼虫踪迹对受孕雌虫在猎物寄主上停留时间和产卵的影响。结果表明:受孕雌虫倾向于在猎物丰富的植株上停留。幼虫踪迹抑制了受孕异色瓢虫雌虫的停留时间,但2龄幼虫踪迹和4龄幼虫踪迹对受孕雌虫停留时间的影响差异不显著。异色瓢虫雌虫将卵产于有蚜虫的植株上,幼虫踪迹对受孕雌虫的产卵抑制作用显著,但2龄和4龄幼虫踪迹的抑制效果差异不明显。     

异色飘虫幼虫对桃大尾蚜的捕食作用

试验研究了异色飘虫二－四龄幼虫对桃大尾蚜低龄和高龄若蚜的捕食作用,对低龄若蚜的捕食作用,三龄幼虫的捕食量在猎物密度为40头以上时显著高于二龄幼虫,四龄幼虫的捕食量在猎物密度为70头以上时显著高于三龄幼虫。对高龄若蚜的捕食作用,三龄幼虫的捕食量在猎物密度为20头以上时显著或极显著高于二龄幼虫,四龄幼虫的捕食量在猎物密度为30头以上时显著高于三龄幼虫。异色飘虫二－四龄幼虫对桃大尾蚜低龄和高龄若蚜的捕食功能反应均为HollingⅡ型。用a/Th来评价天敌对害虫的控制能力,异色飘虫蚴虫龄期愈大,对桃大尾蚜控制作用愈强。     

Symbiont‐mediated chemical defense in the invasive ladybird Harmonia axyridis

The volatile alkylpyrazines methyl‐ and methoxypyrazines (MPs) present in the reflex bleeds of coccinellid beetles such as the harlequin ladybird beetle Harmonia axyridis are important semiochemicals that function in antipredatory defense behavior. Pyrazines have also been coadapted from a primarily defensive role into pheromones that function in intraspecific communication, attraction, and aggregation behavior. However, the biosynthesis of MPs in ladybird beetles is poorly understood. Here, we tested the hypothesis that MPs could be produced by microbial symbionts in H. axyridis, which generates four different MPs. The evaluation of tissue‐specific MP production showed that MP concentrations were highest in the gut tissue and hemolymph of the beetles rather than the fat body tissue as the presumed site of MP biosynthesis. Furthermore, manipulation of gut microbiota by antibiotic‐containing diets resulted in a lower MP content in adult beetles. The analysis of the bacterial community of the digestive tract revealed the presence of bacteria of the genera Serratia and Lactococcus which are reportedly able to produce MPs. In line with the known diet‐dependent production of MP in H. axyridis, we determined that the presence or relative abundance of some of the potential MP producers (Enterococcus and Staphylococcus) is also diet‐dependent. We hypothesize a potential role of the microbiota in MP production in H. axyridis as a possible example for outsourcing the synthesis of ecologically important semiochemicals to its gut bacteria.     

外源补充多巴胺对异色瓢虫突变体gr生长发育及鞘翅颜色的影响

异色瓢虫Harmonia axyridis是我国常见的一种捕食性天敌昆虫,其基于黑色素的体色在环境适应中发挥着重要作用.多巴胺(dopamine,DA)是昆虫合成黑色素的一类重要前体物质,同时在调节生理活动方面具有重要作用,但尚不清楚多巴胺在异色瓢虫生长发育中的作用.本实验室前期发现一异色瓢虫浅色突变体gr,其体色的黑化程度明显浅于野生型.本研究以该突变体为研究对象,以添加不同浓度DA的人工饲料(10 mg/g(D-H)、1 mg/g(D-M)、与0.1 mg/g(D-L))饲养幼虫,测定生长发育相关参数及新羽化成虫的鞘翅颜色参数.结果显示:桃蚜Myzus persicae饲喂至2龄的gr取食对照饲料后无法成功发育至3龄,而取食添加DA的饲料后可发育至4龄,但不同饲料间的存活率无显著差异,同时D-L处理的3龄幼虫发育时间显著短于D-H与D-M.桃蚜饲喂至4龄后取食人工饲料,D-M处理的化蛹率、羽化率、雌雄虫体重及鞘翅黑色斑点的Brightness值均高于其它两种饲料及对照处理,并且D-L处理新羽化成虫鞘翅黑色斑点的Brightness值最小(颜色最黑).上述结果表明DA在异色瓢虫幼虫生长发育中具有重要作用,但受浓度影响;幼虫期补充DA可影响成虫鞘翅颜色.     

异色瓢虫成虫对麦二叉蚜的捕食作用模型

通过异色瓢虫雌,雄成虫对麦二叉蚜捕食作用研究,得出：1．异色瓢虫雌、雄成虫对麦二叉蚜捕食作用的功能反应均是HollingⅡ型反应;2．异色瓢虫捕食麦二叉蚜时,同等数量的异色瓢虫雌雄混合的干扰作用大于同一性别成虫间的干扰作用;3．异色瓢虫雌、雄成虫均喜好捕食低龄若蚜．     

Fine structural analysis of the fibrillar adhesion apparatus in ladybird beetle

The attachment system on the ladybird beetle Harmonia axyridis is composed of a pair of pretarsal claws and adhesive pads at the tarsal segments. The claws, which are connected to the pretarsal segment, are mainly used to hold the rough substrates by their apical diverged hooks. In contrast, the adhesive pads have an adhesive function when landing on smooth surfaces. They are interspersed at the ventral adhesive pad of each tarsomere, and are composed of two kinds of hairy setae. The discoid tip seta (DtS) is located at the central region of each adhesive pad. The DtS has a spoon‐shaped endplate with a long and narrow shaft. In contrast, the pointed tip seta (PtS) is interspersed along the marginal regions of each adhesive pad, and has a hook‐shaped spine near the tip. In the present study, we found numerous fine cuticular pores beneath the setae, which seem to be related to the secretion of some adhesive fluids. It may be deduced that ladybird beetles can attach to smooth surfaces more effectively by employing adhesive fluids filling in surface crevices to overcome problems cause by their larger size endplates.     

不同温度条件下异色瓢虫四龄幼虫亲缘关系对自残行为的影响

在14、18、22、24℃及30℃下对异色瓢虫Harmonia axyridis(Pallas)的不同同血缘的四龄幼虫自残行为及其对生长发育的影响进行研究。结果表明,异色瓢虫四龄幼虫自残发生概率受环境温度条件影响显著。在22-24℃处理范围内同血缘个体间自残发生概率均显著小于非同血缘个体。自残滞后时间同样受环境温度条件影响显著,且在同一温度条件下同血缘个体间均显著小于非同血缘处理。个体间相互攻击率不受环境温度变化影响,而在高于22℃下受血缘关系影响显著。此外,总受攻击次数在不同温度处理间差异显著。同血缘个体间自残后死亡率随温度增加而减少,均在30%以上。而非同血缘个体间自残后死亡率不受温度变化影响,均在70%以上。自残后残存个体的化蛹率及成虫羽化率均不受环境温度变化及个体间血缘关系的影响,化蛹率除30℃处理外均在90%以上,而在各温度下两血缘处理的羽化率均大于70%。     

Detecting seasonal variation in composition of adult Coccinellidae communities

1. Following its recent global spread, Harmonia axyridis (Coleoptera: Coccinellidae), an aphid predator of East Palearctic origin, dominated local coccinellid communities. While long‐term trends in this change are well documented, little is known about seasonal changes. 2. Seasonal changes in communities of adult Coccinellidae were investigated in central Europe (Prague), from April to November in 2010–2014. The beetles were collected at 2‐weekly intervals by net‐sweeping particular stands of trees (Tilia spp., Acer spp. and Betula sp.). The numbers of beetles caught were recalculated per unit effort, and average species abundance and community richness, dominance and diversity were calculated for each 2‐week period. 3. Frequencies of 21 established species varied throughout the vegetative season and general trends in this variation were similar in each of the 5 years of this study. In May–June the abundant species was Adalia decempunctata, which was later replaced by Harmonia axyridis as the most abundant species in late summer. Coccinella septempunctata colonized trees occasionally in summer. This sequence of species substitution occurred every year and was reset during winter. Species richness decreased and dominance increased during the course of each vegetative season. The diversity of the coccinellid community (Shannon index H′) significantly decreased during the course of the vegetative season. This was caused by a gradual increase in the dominance of H. axyridis, the abundance of which explained 88% of variation in H′. 4. The existence of a significant seasonal difference indicates that it is important to take this into consideration when comparing coccinellid communities.     

异色瓢虫的应用研究概况

从异色瓢虫Harmoniaaxyridis(Pallas)的生物学和生态学特性 ,包括色斑变化、生活史、繁殖、食性、捕食、越冬、贮存、人工饲养、田间动态以及在生物防治上的应用等方面概述了异色瓢虫的研究进展。并对异色瓢虫在农业生产特别是果园、菜地、温室花卉上的应用前景作了讨论 ,指出异色瓢虫是一种很有利用前景和产业化生产价值的天敌生物     

异色瓢虫幼虫对桃一点斑叶蝉捕食作用的研究

异色瓢虫在自然界中普遍存在,且可捕食多种小型害虫,在生物防治中占有重要位置。本文就异色瓢虫不同龄期幼虫对不同龄期（虫期）桃一点斑叶蝉捕食作用的研究结果,这时以虫治虫工作提供了理论依据。主要结果为：（1）异色飘虫幼虫对桃一点斑叶蝉的功能反应属HollingⅡ型。（2）随着捕食者和猎物年龄的增加,瞬间玫击率增大,处理时间增长。同时干扰常数亦增大。（3）在最适温度范围内,温度升高可促进取食活动。（4）空间的大小不能改变功能反应类型,仅能使参数数值发生变化。（5）异色瓢虫幼虫喜好选择高龄桃一点斑叶蝉若虫捕食。     

越冬代七星瓢虫和异色瓢虫的飞翔能力

在室内用飞行磨吊飞技术测定了七星瓢虫和异色瓢虫越冬代自然种群的飞翔能力。个体间飞翔特性差异极大。36.5%的七星瓢虫和23.7%的异色瓢虫每次飞翔都不到30分钟, 而这两种瓢虫中都有累计飞翔20小时以上的个体。最长持续飞翔时间达8小时以上和12小时以上的个体在七星瓢虫中分别为9.4%和1.2%, 在异色瓢虫中分别为21.1%和13.2%。大于30分钟的飞翔累计时间为6小时以上和12小时以上的个体在七星瓢虫中分别为20.0%和7.1%, 异色瓢虫则达47.4%和28.9%。雌性个体在卵发育期间表现出很强的飞翔能力。吊飞124小时, 飞翔能力的最高纪录是, 七星瓢虫累计飞翔22小时27分, 飞翔距离94.464km;异色瓢虫为24小时, 92.550km;飞翔速度都达到2m/秒。     

A comparative study of larval cannibalism in three species of ladybird

Abstract. 1. Cannibalism was studied under laboratory conditions in three species of Coccinellidae (Coleoptera): Cycloneda sanguinea , Olla v-nigrum , and Harmonia axyridis . Larval cannibalism varied among species, C. sanguinea  > H. axyridis  ≥  O. v-nigrum .
2. Larvae of all species cannibalised more in response to reduced food availability (14 h starvation daily) than in response to reduced food quality (dry Ephestia eggs).
3. Larvae of H. axyridis cannibalised siblings at lower rates than non-siblings, but larvae of C. sanguinea and O. v-nigrum did not. Rates of cannibalism increased significantly with increasing size disparity among larvae of all three species.
4. Cannibalism in C. sanguinea and H. axyridis , but not in O. v-nigrum , increased with larval density, suggesting that not all attacks on conspecifics are driven by hunger.
5. Costs, not benefits, were the predominant effects of exclusively and partially cannibalistic diets. A diet of conspecific eggs yielded survival equivalent to the Ephestia egg diet for H. axyridis and O. v-nigrum , but developmental time was extended and adults were smaller. Larvae of C. sanguinea had superior survival and faster development on conspecific eggs than on the Ephestia egg diet. Only H. axyridis larvae survived as well on diets comprised exclusively or partially of conspecific larvae as on Ephestia eggs, although they developed more slowly.     

Impact of plant architecture and extraguild prey density on intraguild predation in an agroecosystem

Several models and experimental studies conducted in confined environments have shown that intraguild predation (IGP) can modulate population abundances and structure communities. A number of ecological and abiotic factors determine the nature and frequency of IGP. This study examined the effect of plant architecture and extraguild prey density, and their interactions, on the occurrence of IGP between two species of ladybird, Harmonia axyridis (Pallas) and Propylea quatuordecimpunctata L. (both Coleoptera: Coccinellidae). Theoretical concepts predict that IGP levels would increase with a decrease of both extraguild prey density and plant structural complexity. We conducted a factorial experiment in an open soybean field into which coccinellid larvae were introduced in experimental plots for a period of 5 days. We tested two levels of soybean aphid [Aphis glycines Matsumara (Hemiptera: Aphididae)] density, low and high (100 and 1 000 aphids per plot, respectively), and two levels of plant complexity, low (by removing half of the branches from the soybean plants) and high (by leaving plants intact). We used species‐specific molecular markers to detect the presence of P. quatuordecimpunctata in the digestive tract of H. axyridis. Molecular gut‐content analysis of H. axyridis revealed that rates of IGP were higher (20%) at low aphid density than at high aphid density (<6%). Decreased plant complexity did not impact the frequency of IGP. In accordance with existing literature, this study demonstrates that IGP is amplified at low extraguild prey density. We conclude that considering environmental factors, such as extraguild prey density, is crucial to improve our ability to predict the impact of intraguild predation on community structure and, from an applied perspective, biological control.