A natural flightless mutation in the ladybird, Harmonia axyridis

A homozygous flightless strain, obtained by selection from a laboratory population of Harmonia axyridis Pallas (Coleoptera, Coccinellidae), has been reared under controlled conditions since 1982 after sampling in China. The adults of this flightless strain have normal elytra and wings but drop almost vertically when they attempt to fly. The origin of this mutation, whether laboratory or natural, is discussed. As the mutation does not affect the fitness of the beetles, this flightless strain can be used in biological control. Releasing flightless adults rather than larvae should provide the potential for more continuous control of aphids by both larvae and adults.     

亚致死剂量杀虫剂对异色瓢虫繁殖力的影响

为协调害虫的化学防治与生物防治,测定了吡虫啉、鱼藤酮、氰戊菊酯、阿维菌素、抗蚜威和印楝素6种杀虫剂亚致死剂量对异色瓢虫成虫繁殖力的影响．结果表明,各处理异色瓢虫每雌每日产卵量为40～55粒,吡虫啉接触法和饲喂法预处理及鱼藤酮和印楝素饲喂预处理后,均显著低于对照．所有处理间每雌累积产卵量差异不显著．6种药剂2种方式处理异色瓢虫后,其所产卵的孵化率均低于对照,从卵发育至蛹期的累积存活率均显著降低．吡虫啉两种方式处理和氰戊菊酯接触处理,从卵至蛹期的总发育历期显著延长,鱼藤酮两种方式处理和氰戊菊酯、阿维菌素、抗蚜威、印楝素饲喂法处理的总发育历期明显比对照缩短,而鱼藤酮和阿维菌素接触处理的与对照差异不显著．药剂直接浸渍处理异色瓢虫卵后,各处理卵的孵化率均较对照降低．氰戊菊酯处理的l龄幼虫存活率也明显较对照低,2龄至蛹期的存活率也有所下降．鱼藤酮和氰戊菊酯处理的总发育历期显著延长．这些研究结果显示,低剂量的杀虫剂对异色瓢虫繁殖力的影响很复杂,药剂的类型、作用方式、毒性高低以及给药途径等均影响到试验结果．     

Ecology of two-spotted ladybird, Adalia bipunctata: a review

Abstract:  Adalia bipunctata (Linnaeus) is a generalist aphidophagous ladybird having a wide prey range. It exhibits complex polymorphism. We reviewed the information on its general characteristics, polymorphism, sexual activity, foraging behaviour, food range and prey suitability, growth and development, heterospecific interactions including natural enemies, and its biocontrol potential. Although a poor biocontrol agent (only successful against aphid, Dysaphis plantaginea in Switzerland), it was used in the past as a good model to test various hypotheses and models facilitating ecological understanding in insects. In the present review, the empirical data are interpreted and inferences drawn and a checklist of its prey is provided.     

An artificial non-flying mutation to improve the efficiency of the ladybird Harmonia axyridis in biological control of aphids

The use of coccinellids in the biological control of aphids is restricted to the release of larvae because adults tend to fly away. Non-flying adults may stay longer in one place and so they and their progeny could give longer term protection to plants. This work is an attempt to produce a non-flying population by the use of a chemical mutagen and selection of adults with wing malformations through their subsequent generations. These adults are characterized by open elytra and extended wings. Some general features of this mutation were disclosed. The mutation is either unexpressed or results in malformed wings. It also seems recessive and lethal when homozygous. The adults with the mutation suffered a high level of mortality and a drastic reduction in reproductive capacity that prevents their mass rearing for biological control. This study revealed a negative relationship between wing malformations and reproductive capacity. Nevertheless, when adults with the mutation were released in greenhouses containing cucumbers infested with the aphid Aphis gossypii, they remained on the plants in higher numbers and laid eggs over a longer period of time than the control adults but their progeny were less numerous.     

Testing semiochemicals from aphid,plant and conspecific: attraction of Harmonia axyridis

Abstract Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is an invasive specie affecting the dynamics and composition of several guilds. Nowadays, no biological control method is available to reduce the populations of this harmful coccinellid. Attractants and semiochemicals seem to be the best alternative but only few studies have tested the impact of semiochemicals on this Asian lady beetle. In this work, through wind‐tunnel experiments, semiochemicals from aphids (Z,E‐nepetalactone, [E]‐β‐farnesene, α‐pinene and β‐pinene), from coccinellids ([‐]‐β‐caryophyllene) and from the nettle Urtica dioica L. were evaluated as potential attractants. The nettle volatile compounds ([Z]‐3‐hexenol and [E]‐2‐hexenal) were extracted using a Clevenger Apparatus^® and identified by headspace gas chromatography–mass spectroscopy. In the wind‐tunnel experiments, the main components of the aphid alarm pheromone as well as a component of the aphid sexual pheromone strongly attracted both sexes of the Asian lady beetle while (‐)‐β‐caryophyllene only attracted few individuals and had no impact on the males. The nettle extract as well as the (Z)‐3‐hexenol oriented both males and females to the odor source. The (E)‐2‐hexenal was shown to have no effect on females even if this green leaf volatile attracted males. Because Z,E‐nepetalactone was identified as the most efficient attractant in the wind‐tunnel experiments, this volatile was also tested in a potato field where H. axyridis has been showed to respond to this semiochemical. This study highlighted that Z,E‐nepetalactone orientated the Asian lady beetle H. axyridis under natural conditions, indicating that this volatile compound could certainly help for an efficient biological control approach against this invasive specie.     

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。     

Toxicity of two ecdysone agonists, halofenozide and methoxyfenozide, against the multicoloured Asian lady beetle Harmonia axyridis (Col., Coccinellidae)

Abstract: The insecticidal activity of two ecdysone agonists, methoxyfenozide (RH-2485) and halofenozide (RH-0345), was tested against last-instar larvae of the natural predator Harmonia axyridis (Col., Coccinellidae). In addition, the relative weight gain of the larvae after application was followed. Both products proved to be equally toxic at concentrations of 25, 50 and 100 mg/l. The ecdysteroidal activity of the compounds caused premature induction of larval moulting, cessation of feeding and incomplete pupation in affected larvae. Although, compared with previous results with methoxyfenozide and halofenozide in target pests such as the Colorado potato beetle, Leptinotarsa decemlineata , these compounds caused mortality only after application at relatively high concentrations.     

异色瓢虫生物生态学研究进展

原产于亚洲的异色瓢虫[Harmonia axyridis Pallas(Coleoptera:Coccinellidae)]是最重要的捕食性瓢虫种类之一,被广泛应用于农林业害虫的生物防治.本文综合分析了国内外近几十年对异色瓢虫的研究成果,对其重要的生物学及生态学特性进行了归纳,主要涉及生活史、繁殖行为及策略、捕食行为和自残行为等,分析了相关研究对其应用前景的影响.基于异色瓢虫的人工饲养、农药交互作用以及入侵生态影响等的总结,提出了预防异色瓢虫造成生态不平衡的具体措施.     

Experimental evidence for the phenotypic impact of admixture between wild and biocontrol Asian ladybird (Harmonia axyridis) involved in the European invasion

Hybridization can fuel evolutionary processes during biological invasions. The harlequin ladybird Harmonia axyridis has long been used as a biocontrol agent before the species became invasive worldwide. Previous analysis based on microsatellite data has shown that European invasive populations bear traces of admixture between an eastern North American source, which is at the origin of the worldwide invasion, and biocontrol strains used in Europe. In this study, we tested the hypothesis that this early admixture event may have fostered the European invasion by impacting on the phenotypes of wild European populations. Mean life history traits of experimental F(1) hybrids are compared with pure parental sources and wild European crosses. Our results reveal a biased impact whereby North American beetles benefitted from being admixed with European biocontrol strains. Resemblance between experimental hybrids and wild European invasive crosses further suggests a long-lasting effect of admixture that may still be at work and fostering invasiveness.     

温度对异色瓢虫生长发育和繁殖的影响

为探索饲养异色瓢虫Harmonia axyridis（Pallas）的最适温度,了解其对高温的耐受性,组建了异色瓢虫在15,20,25,30和35℃下的实验种群生命表。结果表明：各虫态的发育速率随温度的升高而加快,在15℃下世代的发育历期最长（65.5d）,而30℃时仅为22.4d,20℃和25℃时分别为39.0d和23.0d。卵、幼虫、蛹和世代的发育起点温度分别为9.8,8.2,11.6和8.5℃,有效积温分别为41.1,225.6,81.6,432.6日.度。成虫寿命随温度升高而逐渐缩短,15℃时最长,为77.7d,在30℃下最短,为36.3d;单雌产卵量在25℃时最高,平均为2193.0粒,15℃时最低,为626.0粒。在25℃下,异色瓢虫的种群趋势指数和内禀增长力均较高,分别为916.7和0.15,表明25℃是异色瓢虫生长、繁殖的最适温度。     

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.     

Invasion history, habitat preferences and phenology of the invasive ladybird Harmonia axyridis in Belgium

Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe, H. axyridis is considered to be an invasive alien species because of its potential to disrupt native ladybird communities. Since 1999, the Belgian Ladybird Working Group mapped all Belgian Coccinellidae and recorded data on substratum plants and habitat. The first feral H. axyridis population in Belgium was recorded in 2001, but the expansion rate is decreasing because it now colonised the whole country. Recorded occupancy in Belgium showed an average rate of increase of 189% between 2002 and 2006. In Belgium, H. axyridis occurred in a wide range of habitats, including those of high conservation value. However, habitat and land cover analysis showed that H. axyridis is more frequently found in urbanised landscapes than in semi-natural landscapes. A niche overlap analysis based on plant use data showed that the potential to affect native species is higher for generalist, deciduous and coniferous tree ladybird species than for heathland and wetland specialist species. Phenology data showed that H. axyridis is able to reproduce later in the year than native species. Based on recorded distribution, ecology and phenology, we discuss the potential of H. axyridis to disrupt native ladybird assemblages in Belgium.     

Honeydew volatile emission acts as a kairomonal message for the Asian lady beetle Harmonia axyridis (Coleoptera: Coccinellidae)

Abstract The Asian lady beetle Harmonia axyridis Pallas is considered as an invasive species in most territories where it has been introduced. Because aphid honeydew acts as an attractant for many aphid predators and parasitoids, the objectives of this work were to collect and identify the volatile compounds released from the aphid excretory product to evaluate how these semiochemicals could affect the H. axyridis foraging behavior. Twelve volatile chemicals were identified from the Megoura viciae Buckton honeydew including four alcohols, three ketones, three aldehydes, a pyrazine and a monoterpene. The volatiles 3‐methyl‐1‐butanol and 3‐methyl‐butanal were highlighted as the two most abundant semiochemicals released from the M. viciae honeydew. Vicia faba L. plants treated with crude honeydew attracted more than 80% of the tested individuals with 40% of attracted beetles located on the plant. Four volatile compounds (3‐hydroxy‐2‐butanone, 3‐methyl‐butanal, 3‐methyl‐1‐butanol and limonene) were also highlighted to attract more than 75% of the coccinellids toward the odor source and to locate about 35% of them on the plants. Limonene was the most efficient attractant since 89% of the H. axyridis responded to this odor. The use of the identified semiochemicals as well as the composition of an artificial honeydew could certainly be helpful to control the dispersal of the Asian lady beetle H. axyridis.     

Melanic through nature or nurture: genetic polymorphism and phenotypic plasticity in Harmonia axyridis

Abstract Individuals can adapt to heterogeneity in their environment through either local adaptation or phenotypic plasticity. Colour forms of the ladybird Harmonia axyridis are a classic example of local adaptation, in which the frequency of melanic forms varies greatly between populations. In some populations, there are also large seasonal changes in allele frequency, with melanism being costly in summer and beneficial in winter. We report that the non‐melanic morph of H. axyridis dramatically increases its degree of melanization at cold temperatures. Furthermore, there is genetic variation in reaction norms, with different families responding to temperature in different ways. Variation at different spatial and temporal scales appears to have selected for either genetic or phenotypically plastic adaptations, which may be important in thermoregulation. As melanism is known to have a large effect on fitness in H. axyridis, this plasticity of melanization may have hastened its spread as an invasive species.     

Mating refusal and its significance in females of the ladybird beetle, Harmonia axyridis

ABSTRACT. . Mating receptivity in females of the ladybird, Harmonia axyridis Pallas (Coleoptera: Coccinellidae), was studied with reference to internal and environmental conditions. Females sometimes reject copulation: they run away, or shake off the male and bend the abdomen upwards. Newly emerged females show a greater likelihood of accepting copulation with an advance in the stage of ovarian development, this being related to age. Females usually mate repeatedly, but frequently show refusal behaviour under unfavourable food conditions. Dissections reveal that females which accept copulation have a greater number of mature eggs in the ovary than do rejectors. Results also suggest that the food conditions are a major environmental factor, directly controlling mating receptivity.     

Phenotypic variation in invasive and biocontrol populations of the harlequin ladybird, Harmonia axyridis

Despite numerous releases for biological control purposes during more than 20 years in Europe, Harmonia axyridis failed to become established until the beginning of the 21st century. Its status as invasive alien species is now widely recognised. Theory suggests that invasive populations should evolve toward greater phenotypic plasticity because they encounter differing environments during the invasion process. On the contrary, populations used for biological control have been maintained under artificial rearing conditions for many generations; they are hence expected to become specialised on a narrow range of environments and show lower phenotypic plasticity. Here we compared phenotypic traits and the extent of adaptive phenotypic plasticity in two invasive populations and two populations commercialized for biological control by (i) measuring six phenotypic traits related to fitness (eggs hatching rate, larval survival rate, development time, sex ratio, fecundity over 6 weeks and survival time of starving adults) at three temperatures (18, 24 and 30°C), (ii) recording the survival rate and quiescence aggregation behaviour when exposed to low temperatures (5, 10 and 15°C), and (iii) studying the cannibalistic behaviour of populations in the absence of food. Invasive and biocontrol populations displayed significantly different responses to temperature variation for a composite fitness index computed from the traits measured at 18, 24 and 30°C, but not for any of those traits considered independently. The plasticity measured on the same fitness index was higher in the two invasive populations, but this difference was not statistically significant. On the other hand, invasive populations displayed significantly higher survival and higher phenotypic plasticity when entering into quiescence at low temperatures. In addition, one invasive population displayed a singular cannibalistic behaviour. Our results hence only partly support the expectation of increased adaptive phenotypic plasticity of European invasive populations of H. axyridis, and stress the importance of the choice of the environmental parameters to be manipulated for assessing phenotypic plasticity variation among populations.     

人工扩繁代异色瓢虫卵和成虫最适冷藏条件的探讨

探讨了人工扩繁异色瓢虫时卵和成虫的最适冷藏条件。卵的最适保存温度是 1 0℃ ,冷藏 1 5d内孵化率平均在 60 %以上。成虫的最适保存温度是 1 0℃ ,如将刚羽化的成虫直接在该温度下冷藏 ,冷藏 40d时存活率能保持在 5 0 %左右 ;如将刚羽化成虫先在 1 5℃、0L∶2 4D条件下处理 1 6d,然后置于 1 0℃下冷藏 ,经冷藏 70d后存活率接近 1 0 0 % ,冷藏 90d后存活率仍高于 70 %。表明经一定条件下预处理后再冷藏成虫能保持较高的存活率和较长的存活期 ,可满足人工扩繁时对成虫的冷藏要求。     

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。     

Sibling cannibalism in the ladybird beetleHarmonia axyridis: Fitness consequences for mother and offspring

The adaptive significance of sibling cannibalism was analyzed in the ladybird beetle Harmonia axyridis at two prey densities. Possible costs and benefits were considered from three points of view: the mother, the cannibal and the victim. Cannibals ate both infertile and fertile eggs, thereby increasing in body length and survival rate with the intensity of sibling cannibalism. The cannibalistic trait was clearly beneficial to the cannibal when aphid density was low. However, it was not always beneficial when aphid density was high and the victims were full siblings. The altruistic behavior of being a victim was beneficial only when the victim was cannibalized by full siblings at low aphid density. The mother attained almost equal fitness at low aphid density, regardless of the intensity of sibling cannibalism. This suggests that sibling cannibalism is not maladaptive for the mother. At high aphid density, however, mother's fitness decreased with the intensity of sibling cannibalism, indicating that sibling cannibalism is maladaptive for the mother when larval food availability is high.     

Genetic estimations for body size characters,development period and development rate in a coccinelid beetle,Harmonia axyridis

Heritabilities and genetic correlations for body size characters and development period in a coccinellid beetle,Harmonia axyridis were estimated by a sib-analysis experiment. Positive genetic correlations were detected between size characters and development rate. If this is upheld in the field, genetic variation would be eliminated, as the loci affecting the characters are supposed to be fixed. However, the results indicated moderate heritabilities for all characters. Possible explanations for the results are discussed.