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.     

Demographic traits of Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae) reared on Aphis fabae Scopoli (Hemiptera: Aphididae)

Lady beetles are the most important predatory species among arthropods; thus, studying their population parameters gives a clear picture about their life span. The demography of the lady beetle, Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae), was studied under laboratory condition, on Aphis fabae at 27?°C based on the age-stage, two-sex life table. The means and standard errors were estimated by the bootstrap techniques. The results showed that the intrinsic rate of increase (r) and the finite rate of increase (λ) were 0.18 and 1.20 d^?1, respectively. The net reproduction rate (R ₀) and the gross reproduction rate were 232.49 and 442.48 offspring/individual, respectively. The mean generation time (T) was 29.03?days. The first peak of reproductive value in females was on the 18th day, which coincides with the total pre-oviposition period counted from birth. The life expectancy curve (e_xj ) of a new hatched female adult was 60?days. The results demonstrated that growth, survival and development of the predator can be described accurately by use of the age-stage, two-sex life table.     

Target-oriented dissemination of Beauveria bassiana conidia by the predators,Harmonia axyridis (Coleoptera: Coccinellidae) and Chrysoperla carnea (Neuroptera: Chrysopidae) for biocontrol of Myzus persicae

Dissemination of microbial biocontrol agents via predators may have advantages for safe spore dispersal to targeted pests with the added benefit of predation. A laboratory study was conducted to test the target-oriented dissemination of conidia of Beauveria bassiana using larvae of both the multicoloured Asian lady beetle (Harmonia axyridis) and common green lacewing (Chrysoperla carnea) for control of aphids. Maximum dry conidial attachment occurred within approximately 7 min after exposure. After release of the treated predators on leaves of Chinese cabbage, within 12 hours lacewing larvae dispersed 89% of the attached conidia while Asian lady beetles dispersed 93%. Both predators dispersed conidia up to 2.4 m from the release site. Leaf disk bioassays were conducted to compare two application methods; the dissemination of conidia of B. bassiana by predators and the direct application of conidial suspensions. Mortality in sprayed aphids was 91±2.1% compared to 88±2.1 and 84±4.2%, respectively, when conidia were disseminated by lacewings and lady beetles. Predation was not affected in treated lacewing larvae whereas there was a 20% reduction in predation by lady beetle larvae. It appears that B. bassiana can be effectively delivered using certain insect predators.     

Overdispersion of Allothrombium pulvinum larvae (Acari: Trombidiidae) parasitic on Aphis gossypii (Homoptera: Aphididae) in cotton fields

Abstract.

• 1 Dispersion patterns of the protelean parasite, Allothrombium pulvinum Ewing, among individuals of an aphid host, Aphis gossypii Glover, were examined during spring 1991 in several cotton fields in Jiangsu Province, China.
• 2 The variance-to-mean ratios (i.e. dispersion index) of larval mites per host were greater than 1, indicating that the mite parasites were overdispersed among aphid hosts. The variance increased with the mean according to the power law, variance = 1.51 mean¹⁰⁶, which explained 99.7% of the variation in the data.
• 3 The negative binomial distribution adequately describes the patterns of larval mite dispersion among aphid hosts in eight out of ten populations. The degree of clumping (1/k) decreased curvilinearly with parasite density (mites per host).
• 4 Mites were more clumped among adult aphids than among immature ones.
• 5 Ecological and evolutionary consequences of mite overdispersion within host populations are discussed. The role of Allothrombium in pest control is also discussed.

     

Attraction of the predator,Harmonia axyridis (Coleoptera: Coccinellidae), to its prey,Myzus persicae (Hemiptera: Aphididae), feeding on Chinese cabbage

The predatory multicolored Asian lady beetle, Harmonia axyridis, was attracted to volatiles released from Chinese cabbage infested by the green peach aphid, Myzus persicae, in T-tube olfactometer choice tests. However, lady beetle adults and larvae did not respond to clean air, Chinese cabbage alone or green peach aphid alone. Of different prey densities, H. axyridis adults were most attracted to Chinese cabbage infested by 60 M. persicae adults after 24 h. However, H. axyridis larvae were not attracted to Chinese cabbage infested by M. persicae. Mechanically damaged Chinese cabbage attracted neither lady beetle adults nor larvae. Predatory adults were attracted to 60 M. persicae adults after 24 and 48 h, and to 90 M. persicae adults after 12 h, suggesting that the predatory response depends on the prey density. Lady adult beetles did not prefer the volatiles induced by Diamondback moth, Plutella xylostella, indicating that specific host insect specificity attracts respective natural enemies. It can be explained that the volatile compounds emitted from the host plant as a result of herbivore attack preferred by the specific insect species.     

Pathogenicity of hyphomycet fungi to aphids Aphis gossypii Glover and Myzus persicae (Sulzer) (Hemiptera: Aphididae)

The aphids Aphis gossypii and Myzus persicae are cosmopolitan, poliphagous and damage cultivated plants. The effects of the entomopathogenic fungi Beauveria bassiana (isolate IBCB 66), Metarhizium anisopliae (isolate IBCB 121), Paecilomyces fumosoroseus (isolate IBCB 141) and Lecanicillium (= Verticillium) lecanii (isolate JAB 02) on third instar nymphs of A. gossypii and M. persicae were evaluated in the laboratory at 25 degrees C, 70 +/- 10% RH and 12h photophase. The aphids were transferred to petri dishes with a foliar disk (cotton or pepper) with a layer of 1 cm tick of agar-water. The fungi were applied in a suspension containing 1.0 x 106 to 1.0 x 108 conidia/ml. In the control treatment 1 ml of sterilized water was added to the foliar disks. The mortality of aphids was evaluated daily. B. bassiana and M. anisopliae caused 100% mortality at the seventh day after inoculation, for both species. L. lecanii was the fungus that provided mortality later in the aphids and M. persicae was more susceptible to both fungi than A. gossypii.     

Walking activity of flightless Harmonia axyridis (Coleoptera: Coccinellidae) as a biological control agent

The use of flightless strains of the multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), established via artificial selection, can be highly effective as a biological control agent for aphids. However, flightless H. axyridis must depend on walking for dispersion. Therefore, data on the walking activity levels in flightless strains are important for the development of effective methods when releasing these agents in the field. Results of measurement of walking activity levels using an infrared actograph showed that walking activity levels during the daytime (but not nighttime) in both sexes of pure flightless strains tended to be lower than those of control strains. We also found that walking activity levels during the daytime for the F1 generation of hybrid strains, produced by reciprocal crossing between two pure flightless strains, were approximately equal to those of pure strains; the reduction in walking activity levels was not recovered by hybrid vigor. Our results indicate that the reduction in walking activity levels in the pure flightless strains was not caused merely by inbreeding depression stemming from the artificial selection process. Instead, potentially flight ability and walking activity levels in this species may be controlled by the pleiotropic effect of a gene.     

Efficacy and nontarget effects of reduced-risk insecticides on Aphis glycines (Hemiptera: Aphididae) and its biological control agent Harmonia axyridis (Coleoptera: Coccinellidae)

The efficacy of three reduced-risk insecticides (pyrethrins, insecticidal soap, and narrow-range mineral oil) was determined for nymphs and adults of the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), an exotic pest of North American soybean, Glycine max (L.) Merr. These insecticides also were evaluated for nontarget effects on one of the aphid's key biological control agents, multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), including first and third instars, pupae, and adults. A Potter Spray Tower was used to conduct direct spray laboratory bioassays. Results indicated that although pyrethrins and narrow-range mineral oil caused 100% mortality to A. glycines nymphs and adults at 72 h posttreatment, insecticidal soap caused equivalent mortality to only the nymphs during the same time period. However, A. glycines adult mortality due to the insecticidal soap (83.3%) was significantly greater than the control. Pyrethrins were highly toxic to first instars of H. axyridis (98% mortality), but they had no effect on third instars, pupae, or adults. Mineral oil and insecticidal soap were moderately lethal to first (48.9 and 40% mortality, respectively) and third (31.9 and 38.8% mortality, respectively) instars of H. axyridis, but they had no effect on pupae and adults. Our results suggest that pyrethrins, insecticidal soap, and narrow-range mineral oil may prove useful for soybean aphid management in organic soybean due to efficacy against the aphid with differential nontarget effects on select stages of H. axyridis. Additional studies will be necessary to elucidate the efficacy of these insecticides under field conditions.     

仲夏棉蚜种群崩溃机制研究进展

棉蚜是世界性分布的棉花害虫,仲夏棉蚜种群崩溃是生态控制棉蚜危害的重要途径,为棉蚜预测和防治提供了新思路与切入点,但其崩溃机制尚未明确。导致棉蚜种群崩溃的可能因素主要包括:(1)植物次生代谢物影响棉蚜的发育和繁殖,但不足以导致棉蚜种群崩溃;(2)暴风雨对棉蚜种群有一定的影响,但无法解释连续多年和多个地点发生的种群崩溃现象;(3)高温显著降低棉蚜的存活和繁殖,且与仲夏棉蚜种群崩溃期相吻合,高温很可能是棉蚜种群崩溃的关键影响因子;(4)高密度条件下激烈的种内竞争导致棉蚜繁殖率降低,有翅蚜比例增加;(5)天敌除具有直接捕食或寄生作用外,同时间接影响若蚜生长发育和刺激有翅蚜形成,对棉蚜种群后期崩溃发挥重要作用。本文对可能导致仲夏棉蚜种群崩溃的主要因素分别进行论述,探讨其对棉蚜存活、繁殖和有翅蚜形成的影响,旨在为探明仲夏棉蚜种群崩溃机制提供参考。     

Implications of phenotypic variation of Myzus persicae (Hemiptera: Aphididae) for biological control on greenhouse pepper plants

Variation in vulnerability to natural enemies, reproductive rate and insecticide resistance among phenotypes of Myzus persicae (Sulzer) has been shown to have the potential to disrupt biological control and IPM of this species, and movement of particularly troublesome phenotypes in international horticultural trade could be cause for concern. Three important components of fitness, vulnerability to parasitoids, reproduction and insecticide resistance were determined in three clones of M. persicae originating from prevalent phenotype populations on pepper crops in greenhouses in British Columbia, Canada. One of these phenotypes appeared to be consistently involved in outbreaks in commercial operations. These clones were also characterized for their DNA microsatellite genotype and compared with genotypes of M. persicae from Europe. The clone involved in outbreaks in commercial greenhouses showed reduced vulnerability to parasitoids, and a higher reproductive rate compared to the other two clones suggesting that these traits may have been involved in outbreaks. As in M. persicae European clones, a higher reproductive rate was correlated with a lack of esterase‐based resistance (primarily to organophosphates and, to some extent, to carbamates and pyrethroids). However, microsatellite analysis demonstrated that the three clones investigated in British Columbia had unique genotypes, and therefore there was no evidence for their movement in international trade.     

European monitoring of resistance to insecticides in Myzus persicae and Aphis gossypii (Hemiptera: Aphididae) with special reference to imidacloprid

The susceptibility to several insecticides of 16 and 8 strains of Myzus persicae Sulzer and Aphis gossypii Glover, respectively, received from different European countries in 2001 was investigated. Most of the strains were derived from places known for their aphid resistance problems to conventional insecticides before imidacloprid was introduced. In many regions and agronomic cropping systems imidacloprid has been an essential part of aphid control strategies for a decade, and therefore the susceptibility of aphid populations to imidacloprid using FAO-dip tests and diagnostic concentrations in a leaf-dip bioassay was checked. Additional insecticides tested were cyfluthrin (chemical class: pyrethroid), pirimicarb (carbamate), methamidophos and oxydemeton-methyl (organophosphates). Diagnostic concentrations (LC99-values of reference strains) for each insecticide were established by dose response analysis using a new leaf-disc dip bioassay format in 6-well tissue culture plates. Virtually no resistance to imidacloprid in any of the field-derived populations of M. persicae and A. gossypii was detected. In contrast, strong resistance was found to pirimicarb and oxydemeton-methyl, and to a lesser extent also to cyfluthrin. Two strains of A. gossypii exhibited reduced susceptibility to imidacloprid when tested directly after collection. However, after maintaining them for six weeks in the laboratory, the aphids were as susceptible as the reference strain. The diagnostic concentration of methamidophos did not reveal any resistance in M. persicae, but did so in four strains of A. gossypii.     

Comparison of the life tables and predation rates of Harmonia dimidiata (F.) (Coleoptera: Coccinellidae) fed on Aphis gossypii Glover (Hemiptera: Aphididae) at different temperatures

The life histories and predation rates of the ladybird beetle Harmonia dimidiata (F.) were compared among beetles kept at 15, 20, and 25 °C. The beetles were fed on Aphis gossypii Glover and were maintained at 70 ± 10% RH and a 14:10 (L:D) h photoperiod. According to the age-stage, two-sex life table, the net reproductive rates (R₀) were 147.4, 98.7, and 62.5 offspring for beetles kept at 15, 20, and 25 °C, respectively. Additionally, we employed both the jackknife and bootstrap techniques for estimating the means, variances, and standard errors of the population parameters. The sample means of R₀ and the other population parameters obtained using the bootstrap technique fit a normal distribution, but the jackknife technique generated biologically meaningless zero values for R₀. The net predation rates were 10963, 13050, and 7492 aphids for beetles kept at 15, 20, and 25 °C, respectively. For a comprehensive comparison of predation potential, we incorporated both the finite rate and the predation rate into the finite predation rate. When both the growth rate and the predation rate were considered, our results showed that H. dimidiata is a more efficient biological control agent for A. gossypii at 20 and 25 °C than at 15 °C.     

Mating rates and the prevalence of male‐killing Spiroplasma in Harmonia axyridis (Coleoptera: Coccinellidae)

Maternally inherited bacteria that kill male but not female hosts during embryogenesis have been widely reported in invertebrates. Harmonia axyridis is one of the species infected by male‐killing Spiroplasma. The presence of male‐killers in host populations can lead to the occurrence of extremely female‐biased sex ratios. Furthermore, infected females may have fewer chances to mate if males can discriminate between infected and uninfected females and prefer the latter. Although there have been many investigations of male‐killer infection rates in H. axyridis, little is known about the influence of host mating on male‐killer infection dynamics. We investigated copulation rates and changes in infection frequency in a wild population of H. axyridis in western Japan. Almost all infected females collected each year laid fertilized eggs and had therefore mated. Mean infection rates of females collected each year were 13% in 2003, 15% in 2012 and 23% in 2013. Statistical analysis showed that neither the copulation rate nor the infection rate differed significantly among years. These results suggest that the infection rate of H. axyridis with male‐killing Spiroplasma is kept approximately constant and that there is no difference in the chance of mating with infected and uninfected females.     

Aphid consumption and residence time of larvae of flightless lady beetles, Harmonia axyridis (Coleoptera: Coccinellidae), on aphid-infested plants

A flightless strain of the lady beetle, Harmonia axyridis Pallas, has been established, by artificial selection, for effective biological control of aphids. However, this biological control technique is costly because of the high cost of production of the adults. It is, therefore, important to consider the utility of the larvae in biological control. This study examined the potential for larvae of the flightless strain to be used as biological control agents compared with those of the wild-type, in terms of how they behave on aphid-infested plants. We found no differences between the strains with regard to residence time and aphid consumption on the plant for the first, second, and fourth instars, and at low and high aphid densities. In contrast, however, aphid consumption by third-instar larvae of the flightless strain tended to be lower, although they remained on the plant longer than third-instar larvae of the wild-type. Consequently, our results suggest the flightless strain does not differ from the wild-type in its ability to control aphids across all larval stages, except for the third instars. On the basis of these results, and in conjunction with the results from previous studies, we discuss the appropriateness of different larval instars for release.     

Population growth of Aphis gossypii and Myzus persicae (Hemiptera: Aphididae) in the presence of Linepithema humile and Tapinoma sessile (Hymenoptera: Formicidae)

Invasive ant species can have dramatic impacts on native ants, through direct predation and by usurping common resources. Most invasive ants and many native ants use honeydew, produced by phloem-sucking hemipterans. Because colonies of invasive ants can become very large after establishment, these ants may facilitate greater hemipteran trophobiont population growth compared with their sympatric native ant counterparts. We examined the population growth of an aphid mutualist, Aphis gossypii, and a nonmutualist, Myzus persicae, exposed to two Dolichoderine ants, Linepithema humile, a globally widespread invasive species, and Tapinoma sessile, a widespread co-occurring native ant, in North America in an enemy-free laboratory study. L. humile worker foraging activity was at least twice that of T. sessile, and populations of the myrmecophile, A. gossypii, were greater when exposed to L. humile than T. sessile, possibly caused, in part, by more frequent encounters with L. humile. L. humile ignored M. persicae when A. gossypii was absent, whereas T. sessile preyed on it. Both ant species preyed on M. persicae when A. gossypii was also present. This suggested that both ants may assess nutritional gains from aphid species (i.e., honeydew versus body tissue), eliminating less productive aphids competing for host plant space. Through their impact on populations of hemipteran mutualists, we suggest that colonies of L. humile and perhaps other invasive ants may acquire more honeydew than native ants, thereby fueling colony growth that leads to numerical dominance and widespread success in introduced environments.     

Effect of alfalfa habitat change on dispersal behavior of Harmonia axyridis Pallas and Hippodamia variegata Goeze (Coleoptera: Coccinellidae)

An alfalfa grassland landscape was divided into 36 plots during 2014 season in Yuanzhou District of Guyuan city in Ningxia Hui Autonomous Region of P.R. China. In these three times repeated experiments, treatment of habitat fragmentation was divided into two grades: continuity (H = 0.0) and fragmentation (H = 1.0), and their habitat loss rate was divided into 6 grades: 90, 80, 70, 50, 20 and 0%. Considering habitat loss 0% of the plot as the reference, as a result of sweeping net and visual methods, results showed that the flight distance of the Harmonia axyridis was much higher than that of the Hippodamia variegata both in fragmentation and continuous habitats. In the alfalfa landscape, H. axyridis moved more by flying, while H. variegata moved more by crawling. There was a threshold effect that affected the movement behavior of lady beetles between 70 and 100% of the habitat loss. The population of the two species of lady beetles in the continuous habitat was higher than that in the habitat fragmentation and was more obvious in 70 and 50% plots of the habitat loss. The dispersal rate of the two species of lady beetles inside plots was higher than outside of the experimental plots, which reflected the low diffusivity of the ladybeetle. Habitat fragmentation affected the dispersal, migration and construction of two species of lady beetles on different spatial scales by disturbing the spread of natural enemies and searching for prey.     

Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae), a predacious ladybird new in Australia

Abstract The originally Palaearctic but now widespread coccinellid Hippodamia variegata was recorded for the first time in Australia at Gatton, Queensland, in November 2000 . Since then, it has been recorded preying on 12 different aphid species and a psyllid, on a variety of crops, weeds and ornamental plants. Within a year of its discovery, it had been recorded within a distance of 800 km north and south and 500 km west of the original collection point. Collection and observation records are presented.     

Foraging behavior of the coccinellid Nephus includens (Coleoptera: Coccinellidae) in response to Aphis gossypii (Hemiptera: Aphididae) with particular emphasis on larval parasitism

This study assessed the effect of parasitism of Nephus includens (Col.: Coccinellidae) larvae by Homalotylus flaminius (Hym.: Encyrtidae) on the predation rates of the predator on the cotton aphid, Aphis gossypii Glover (Hem.: Aphididae) by deriving functional responses for second- and fourth-instar predators at prey densities ranging from 10 to 80 aphids per arena. The relationship between the functional and numerical responses of adult females of N. includens also was determined for prey densities ranging from 10 to 140 aphids per arena. Predation rates of unparasitized and parasitized second-instar N. includens were both fit by a type II functional response model with parameters as follows: unparasitized (a = 0.0768 hours(-1) and T(h) = 0.975 h) and parasitized (a = 0.0787 hours(-1) and T(h) = 0.8823 hours). Predation rates of unparasitized and parasitized fourth-instar N. includens were fit by type III and II models, respectively, with the following parameters: unparasitized (b = 0.1702 hours(-1) and T(h) = 0.2369 hours) and parasitized (a = 0.038 h(-1) and T(h) = 0.539 h). The unparasitized fourth-instar was the most voracious stage, having the highest attack rate and lowest handling time. Considering these attributes, it would seem to be the most effective stage of this predator against A. gossypii. Adult female lady beetles (N. includens) showed a type III functional response and their numerical response increase to prey density was curvilinearly related to prey density, with the highest number of eggs being produced at highest prey densities. The maximum saturation level for both predation and egg production for adult females occurred at a prey density of 120 aphids. Thus, a ratio 1:120 (predator:prey) should be used when releasing this species for augmentative biological control. Release of either fourth-instar or adult stage N. includens should be minimized the potentially negative effect of parasitism by H. flaminius on early developmental stages, and hence increase its efficiency in biocontrol programs.     

Feeding by the coccinellid Psyllobora rufosignata (Coleoptera: Coccinellidae) on the Asian grapevine leaf rust fungus Phakopsora euvitis (Basidiomycota: Uredinales)

Feeding by the coccinellid Psyllobora rufosignata on the rust fungus Phakopsora euvitis was confirmed by observing uredospores of P. euvitis in gut contents of Ps. rufosignata. This is the first record of Ps. rufosignata feeding on P. euvitis and the first record of any Halyziini species feeding on any species of Basidiomycota.