Effect of floral and extrafloral resources on predation of Aphis spiraecola by Harmonia axyridis on Apple

Flowering companion plants and plants producing extrafloral nectar are being proposed to enhance biological control in apple (Malus domestica Borkh.) orchards. This experiment evaluated the impact of floral and extrafloral resources on predation of spirea aphid (Aphis spiraecola Patch) on apple by adult Harmonia axyridis (Pallas) under greenhouse conditions. Predation of spirea aphids was not affected by the presence of flowering buckwheat (Fagopyrum esculentum) as compared with either buckwheat with flowers removed or an uninfested apple shoot. However, there was a significant reduction in predation of spirea aphids on an apple shoot in the presence of a peach shoot with extrafloral nectar glands compared with either a peach shoot of the same cultivar without nectar glands or an uninfested apple shoot. These findings demonstrate that alternative food resources potentially could interfere with rates of biological control and, therefore, need to be carefully evaluated before incorporating in an orchard design. More studies are needed to adequately gauge the net effects of adding floral or extrafloral resources in orchards for conservation biological control.     

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.     

Predation and functional response of the multi-coloured Asian ladybeetle Harmonia axyridis on the adult Asian citrus psyllid Diaphorina citri

Diaphorina citri (Hemiptera: Liviidae) is a vector for the citrus Huanglongbing greening disease, which can reduce crop yields. Harmonia axyridis (Coleoptera: Coccinellidae) is known to prey on D. citri nymphs; we investigated the predation capacity and functional response of H. axyridis on adult D. citri. H. axyridis larvae exhibited Holling’s type II functional response to different densities of adult D. citri. For second, third and fourth instar H. axyridis larvae, the successful attack rates were 0.75?±?0.22, 1.06?±?0.11, and 1.04?±?0.26, respectively; the handling times were 18.08?±?5.37, 0.45?±?0.07, and 0.24?±?0.07?h, respectively; and the estimated maximum predation rates were 4, 54, and 102, respectively. Fourth instar larvae had the lowest handling time. Adult H. axyridis consumed more D. citri adults maintained at 15–20°C than at 30–35°C, with an optimal temperature of 17.28°C. More adults were consumed during the day than at night, with peak consumption between 12:00 and 14:00?h, not differing significantly differ between predator genders. Adult H. axyridis reared on the aphid Acyrthosiphon pisum were tested for D. citri predation; these H. axyridis had successful attack rates, handling times, and estimated maximal predation rates of 0.66?±?0.18, 0.82?±?0.26?h, and 29, respectively. Conversely, those fed and tested on D. citri had corresponding rates of 0.99?±?0.17, 0.38?±?0.12?h, and 63, respectively. Significant H. axyridis predation on adult D. citri indicated its enhanced role in suppressing both nymph and adult D. citri.     

Suppression of aphids by augmentative release of larvae of flightless Harmonia axyridis

The release of the larvae of flightless ladybird beetles could extend the duration of effective control because of the longer presence of adult stages, and the per individual production cost for juvenile stages is lower than that for adult stages. A preliminary release experiment was conducted to assess the effectiveness in controlling two aphid species, Aphis gossypii and Aulacorthum solani, using second instars of a flightless strain of Harmonia axyridis. The number of A. gossypii was suppressed in greenhouses that contained the flightless strain compared with greenhouses that contained the wild‐type strain. In one of two replicates, more flightless adults were observed on the plants for longer than wild‐type adults. However, it was not clear whether the extension of the residence period contributed to the effectiveness in controlling aphids directly, because the population of ladybirds in the greenhouses consisted of both larvae and adults during parts of the experimental period. In flightless H. axyridis, the release of larvae was more effective in suppressing A. solani than the release of adults. These results suggest that it may be more effective to release larvae of flightless H. axyridis than wild‐type larvae or flightless adults.     

Inferring the origin of populations introduced from a genetically structured native range by approximate Bayesian computation: case study of the invasive ladybird Harmonia axyridis

Correct identification of the source population of an invasive species is a prerequisite for testing hypotheses concerning the factors responsible for biological invasions. The native area of invasive species may be large, poorly known and/or genetically structured. Because the actual source population may not have been sampled, studies based on molecular markers may generate incorrect conclusions about the origin of introduced populations. In this study, we characterized the genetic structure of the invasive ladybird Harmonia axyridis in its native area using various population genetic statistics and methods. We found that native area of H. axyridis most probably consisted of two geographically distinct genetic clusters located in eastern and western Asia. We then performed approximate Bayesian computation (ABC) analyses on controlled simulated microsatellite data sets to evaluate (i) the risk of selecting incorrect introduction scenarios, including admixture between sources, when the populations of the native area are genetically structured and sampling is incomplete and (ii) the ability of ABC analysis to minimize such risks by explicitly including unsampled populations in the scenarios compared. Finally, we performed additional ABC analyses on real microsatellite data sets to retrace the origin of biocontrol and invasive populations of H. axyridis, taking into account the possibility that the structured native area may have been incompletely sampled. We found that the invasive population in eastern North America, which has served as the bridgehead for worldwide invasion by H. axyridis, was probably formed by an admixture between the eastern and western native clusters. This admixture may have facilitated adaptation of the bridgehead population.     

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.     

Intraguild interactions and aphid predators: biological efficiency of Harmonia axyridis and Episyrphus balteatus

Introductions of the harlequin ladybird Harmonia axyridis into Belgium for aphid biological control have been followed by declines in native aphid natural enemies. We first examined, in laboratory and field conditions, the impact of larval mobility of either H. axyridis or Episyrphus balteatus, the most abundant native hoverfly in central Europe, on aphid suppression. The hoverfly larvae consumed more aphids (Megoura viciae) in a short term, whereas the ladybird ones were more efficient in a long term. We second investigated the intraguild interactions between H. axyridis and E. balteatus larvae and adults. In larva pairings between the two species in laboratory microcosms containing aphid (M. viciae) infested broad bean (Vicia faba) plants, H. axyridis had an intraguild predation (IGP) advantage over the hoverfly. When conspecific larvae were paired together on aphid‐infested plants, no cannibalism between them was detected. The presence of either H. axyridis or E. balteatus larvae on aphid‐infested plants negatively influenced the ovipositional behaviour of H. axyridis and E. balteatus females; lower numbers of laid eggs were recorded compared to control treatment. Moreover, eggs laid by E. balteatus females were also dropped as victims of predation mainly by H. axyridis larvae. Our results suggest that while the exotic ladybird was more efficient in aphid biological control, larvae and eggs of the native hoverfly species face increased IGP by H. axyridis, which would contribute, as a consequence, to the decline in E. balteatus population following invasion.     

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.     

Ecology and biological control application of multicoloured Asian ladybird, Harmonia axyridis: A review

The ecology of and biological control by multicoloured Asian ladybird beetle, Harmonia axyridis (Pallas) are reviewed. Our emphasis is on assembling and interpreting information on general characteristics, invasion and establishment, sexual activity, foraging and predation, development, survival and reproduction, predator-predator interactions, natural enemies, biocontrol, non-target effects and status of H. axyridis as a pest of fruits. Colonization of H. axyridis for aphid biocontrol in the USA have been successful in terms of its establishment, but its abundance is turning out to be a nuisance to humans. Its harmful non-target impact on beneficial organisms, humans and native species is becoming a debatable issue. The question on its present position, whether it is a biocontrol agent or pest, is a critical issue and discussed. Inferences from the empirical data are made and new avenues for future research are suggested.     

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。     

Spatial distribution and fixed-precision sampling plans for the ladybird Harmonia axyridis in sweet corn

To improve the utilization of the multicolored Asian lady beetle, Harmonia axyridis (Pallas), as a natural enemy in integrated pest management (IPM) programs for sweet corn, Zea mays L., fixed-precision enumerative sequential sampling plans for this coccinellid were developed and validated. Data were collected from sweet corn plots during 2000 to 2004, with means ranging from 0.02 to 11.83 H. axyridis (larvae and adults combined) per plant. Taylor’s power law parameters from the regression of log variance versus log mean suggested an aggregated distribution for larvae and the combined count of larvae and adults in sweet corn. For IPM purposes, a plan was developed for H. axyridis larvae and adults to provide a desired precision level of 0.25 (SEM/mean), resulting in an average sample number of 65 whole plants. However, for the purposes of ecological or applied research (with desired precision = 0.10), an average sample number of 205 plants was necessary.     

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。     

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.     

标记-重捕技术研究果园异色瓢虫种群动态

利用标记-重捕技术,对果园异色瓢虫的种群动态进行了估测,结果表明,6月14日至6月28日生防美国杂交杏李园异色瓢虫的种群数量为1058．97头,存活率为0．5534,新增数量为525．48头,将理论标记异色瓢虫在种群中的总数和实际标记释放总数进行配对样本t检验,95％置信区间上下限相差不超过59．43头,试验结果和理论值精度较高。     

Antennal sensory organs and glands of the harlequin ladybird,Harmonia axyridis

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is a sub‐cosmopolitan species. Native to Asia, it has been released during the 20th century for classical and augmentative biological control of several herbivorous insects, mostly aphids and coccids. Despite its recognized positive impact on biological control, H. axyridis is now considered among the most dangerous invasive species in Europe and in most places where it has established. This is mostly due to its ability to reduce the populations of native predatory species of the same trophic guild. When exploring a new area, H. axyridis adults use semiochemical cues to acquire information about the habitat. Presumably, these cues are perceived by the sensilla located on the antennae. Surprisingly, in spite of the huge literature existing on H. axyridis, the antennal sensory organs have been poorly characterized. Here, we used scanning and transmission electron microscopy (SEM, TEM) techniques to study H. axyridis antennae, with focus on the various types of sensilla and their distribution in male and female individuals. The presence of various classes of antennal sensilla belonging to the main types described in insects (chemoreceptors, mechanoreceptors, and thermo‐hygroreceptors) was highlighted, as well as the widespread presence of antennal glands. The investigations showed some peculiar characteristics not known in Coccinellidae, such as the concentration of sensory structures at the level of the distal part of the apical antennomere and the discovery of antennal glands associated with it. No sexual dimorphism was revealed, neither for the general structure of the antenna (similar number of antennomeres and presence of modifications), nor for the total length and width of the antenna, the relative size of the antennomeres, the types of antennal sensilla, of their distribution and abundance. The potential relevance of these sensory structures and antennal glands, reported for the first time in Coccinellidae, is discussed in the context of intra‐ and interspecific communication.     

Effect of temperature on the development and consumption of Phaenochilus kashaya (Coleoptera: Coccinellidae), a predator of the cycad aulacaspis scale,Aulacaspis yasumatsui

The cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), is a serious pest of ornamental cycads in the southeastern United States. In Florida, CAS was first reported in Miami in 1996 and is now present in 43 Florida counties. Chemical control of CAS can be effective, but it is expensive, and insecticides must be frequently and regularly applied. Recent surveys of natural enemies in Thailand identified a new potential biological control agent of CAS. The objective of this study was to examine the temperature requirements and consumption rates of Phaenochilus kashaya Giorgi and Vandenberg (Coleoptera: Coccinellidae). Survival, developmental time and number of scales consumed by P. kashaya were measured at four constant temperatures (20, 25, 30 and 35°C). In addition, adult longevity and consumption were recorded at three constant temperatures (20, 25 and 30°C). Phaenochilus kashaya completed development to adulthood at temperatures ranging from 20 to 30°C. The highest survival rate (48%) was obtained at 25°C, and the accumulated degree-days for this species was 714. Phaenochilus kashaya is a voracious predator of CAS, consuming 380 scales during the larval stage and an estimated 4700 scales during the adult stage. Adult longevity varied from 59 days at 30°C to 220 days at 20°C for females, and 31 days at 30°C to 148 days at 20°C for males. The intrinsic rate of increase (r _m) was 0.34 at 20°C, which is high compared with other coccinellid species. According to this study, P. kashaya has the potential to be an effective biological control agent of the CAS in Florida.     

Decline in native ladybirds in response to the arrival of Harmonia axyridis: early evidence from England

1. Harmonia axyridis (Coleoptera: Coccinellidae) is an invasive non‐native ladybird in Europe, where it was introduced as a biological control agent of aphids and coccids. 2. This study assesses changes to ladybird species assemblages, in arboreal habitats, over a 3‐year period encompassing the invasion phase of H. axyridis in eastern England. The effects of H. axyridis and other factors (weather and prey availability) on native ladybirds are assessed. 3. Harmonia axyridis increased from 0.1% to 40% of total ladybirds sampled, whilst native aphidophagous species declined from 84% to 41% of total ladybirds. The actual number of native aphidophagous ladybirds per survey decreased from a mean of 19.7 in year 1, to 10.2 in year 3. 4. Three ladybird species in particular experienced declines: Adalia bipunctata, Coccinella septempunctata, and Propylea quattuordecimpunctata. Harmonia axyridis was the most abundant species by the end of the study. 5. The decline in native aphidophagous ladybirds could be attributed to competition for prey and intraguild predation of eggs, larvae, and pupae by H. axyridis. Physiological and behavioural traits of H. axyridis are likely to confer an advantage over native ladybird species.     

Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China

Abstract 1. We observed native populations of Harmonia axyridis (Pallas) around Beijing, China, over 2 years and performed choice and no‐choice mating tests between melanic and succinic (non‐melanic) beetles in the laboratory. 2. Succinic phenotypes outnumbered melanics by 5:1 in autumn, but melanics became equally abundant in spring, supporting previous inferences that melanism is advantageous in winter, but costly in summer. 3. Female H. axyridis expressed mate preference overtly, by rejecting less‐preferred phenotypes, and cryptically, by retaining their eggs for longer periods after matings with less‐preferred males, ostensibly to replace their sperm. 4. Succinic pairs formed more quickly in the spring generation, and melanic pairs in the autumn, and the time to copula was affected by both male and female phenotype. The strength of mate preference was contingent on female phenotype, suggesting melanic alleles had pleiotropic effects. 5. Whereas pair formation was under female control, the duration of copula was under male control and lasted longer in the autumn generation than in the spring. Copulations in the choice test tended to be shorter between similar phenotypes, suggesting that males invested more in dissimilar females when alternative mates were available. 6. Although spring and autumn generations were raised under identical conditions, significant contrasts were observed in their reproductive behaviour. 7. Two alternative hypotheses are advanced to explain why gender‐specific reproductive behaviours might vary between generations: maternally‐mediated epigenetic factors that influence the expression of genes in progeny as a function of maternal environment, and linkage disequilibria among alleles that cycle in frequency seasonally as a function of assortative mating.     

Bad side of a good beetle: the North American experience with Harmonia axyridis

The multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is a generalist predator of aphids and other soft-bodied insects and has been utilized in biological control programs around the world. Over the last two decades, this species has spread throughout much of the continental USA and southern Canada. Despite the benefits it offers as a biological control agent, H. axyridis is perhaps most well known for its adverse impacts. In this paper we provide a review of the North American experience with H. axyridis, focusing on these adverse impacts, which can be classified into three general categories: impacts on non-target arthropods, impacts on fruit production and impacts as a household invader. The impacts of H. axyridis on non-target arthropods and, to lesser extent, the impacts as a household invader possibly could have been anticipated, due to its generalist feeding preferences and overwintering behavior in Asia. However, it is unlikely that the impacts on fruit production could have been anticipated. Therefore, even in retrospect, it is difficult to predict the potential impacts that an introduced natural enemy might have in its adventive range.