Dropping behaviour of larvae of aphidophagous ladybirds and its effects on incidence of intraguild predation: interactions between the intraguild prey, Adalia bipunctata (L.) and Coccinella septempunctata (L.), and the intraguild predator, Harmonia axyridis Pallas

Abstract.  1. Two experiments were performed in the laboratory to assess the behaviour of dropping from a host plant as a defence against intraguild predation in aphidophagous ladybird larvae.
2. In the first experiment, encounters were observed on bean plants between fourth instars of the intraguild predator species, Harmonia axyridis , and first instars of two other ladybird species, Adalia bipunctata (L.) and Coccinella septempunctata (L.). The percentages of first instars of the latter two species that dropped from the plant in response to attack differed dramatically, with 47.5% of C. septempunctata first instars dropping vs. 0% of A. bipunctata .
3. In the second experiment, first instars of A. bipunctata or C. septempunctata and a fourth instar of H. axyridis were allowed to forage together on bean plants for 3 h. During this time, 44.3% of C. septempunctata larvae dropped from the plant, but less than 2% of A. bipunctata larvae did so. In contrast, 95.0% of A. bipunctata larvae fell victim to intraguild predation by H. axyridis vs. only 54.5% of C. septempunctata larvae.
4. The significance of dropping behaviour of ladybird larvae as a defence against intraguild predation, and the relationship of dropping behaviour to species-specific habitat affinity of ladybirds, is discussed.     

Interactions between introduced and native predatory ladybirds (Coleoptera, Coccinellidae): factors influencing the success of species introductions

Abstract.  1. Why some exotic species invade successfully while others do not is poorly understood. This study focuses on the inability of the exotic predatory ladybird Adalia bipunctata (L.) (Coleoptera, Coccinellidae) to establish and spread in Japan. In particular, this study considers the interactions between A. bipunctata and three native Japanese ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata Linn., and Propylea japonica Mulsant.
2. The number of aphids occurring on the host plants declined dramatically over the course of larval development in the presence of H. axyridis or C. septempunctata . In contrast, many aphids remained at the end of the experiment when only A. bipunctata or P. japonica was present.
3. The survivorship of A. bipunctata from second instar to adult was significantly reduced in the presence of H. axyridis , but not in the presence of C. septempunctata or P. japonica . Most deaths were the result of intra-guild predation when H. axyridis or C. septempunctata larvae were present. Larvae of H. axyridis and C. septempunctata suffered very high mortality in this experiment, but survived better when co-occurring with A. bipunctata than when occurring only with conspecifics. In contrast, P. japonica survived better when occurring with conspecifics than with A. bipunctata .
4. The finding that larvae of the exotic species A. bipunctata died at high rates from intra-guild predation by the native Japanese ladybirds H. axyridis and C. septempunctata may bear on the question of why A. bipunctata has not established more successfully in Japan whereas H. axyridis and C. septempunctata have been very successful in establishing themselves in North America.     

The potential of three native insect predators to control the rosy apple aphid, Dysaphis plantaginea

The potential of three aphidophagous predators, Adalia bipunctata, Aphidoletes aphidimyza, and Episyrphus balteatus to control the rosy apple aphid, Dysaphis plantaginea Pass., a major pest on apple in Europe, was assessed by means of laboratory and field cage experiments in Northern Switzerland. Under laboratory conditions, all three predators efficiently preyed upon D. plantaginea on apple seedlings. The searching success of larvae of A. bipunctata for individual aphids was not dependent on the size of branches of apple trees varying in leaf surface area from 150 cm² to 960 cm². Fifty and 70% of individual aphids were found and killed 6 hours and 48 hours, respectively, after release of single second instar larva of A. bipunctata. In a first field cage experiment in 1996, A. bipunctata, and to a lesser extent E. balteatus, proved to be effective and consistent predators of D. plantaginea during spring conditions, being little affected by cool temperatures and wet weather. In a subsequent field cage experiment in 1997, larvae of A. bipunctata and E. balteatus were released singly and in combination on aphid infested apple seedlings to study interactions between these two promising control agents. Both species had a significant negative effect on aphid population increase. The two species did not significantly interact and thus, their joint effect is best explained by an additive model. Combined releases of the two predator species reduced aphid densities to 5% of the control. This indicates the potential for augmentative releases of these native aphid predators to control D. plantaginea.     

Intraguild predation behaviour of ladybirds in semi-field experiments explains invasion success of Harmonia axyridis

Harmonia axyridis has been introduced as a biological control agent in Europe and the USA. Since its introduction, it has established and spread, and it is now regarded as an invasive alien species. It has been suggested that intraguild predation is especially important for the invasion success of H. axyridis. The aim of this study was to compare the intraguild predation behaviour of three ladybird species (Coccinella septempunctata, Adalia bipunctata, and H. axyridis). Predation behaviour was investigated in semi-field experiments on small lime trees (Tilia platyphyllos). Two fourth-instar larvae placed on a tree rarely made contact during 3-hour observations. When placed together on a single leaf in 23%-43% of the observations at least one contact was made. Of those contacts 0%-27% resulted in an attack. Harmonia axyridis attacked mostly heterospecifics, while A. bipunctata and C. septempunctata attacked heterospecifics as often as conspecifics. In comparison with A. bipunctata and C. septempunctata, H. axyridis was the most successful intraguild predator as it won 86% and 44% of heterospecific battles against A. bipunctata and C. septempunctata respectively, whilst A. bipunctata won none of the heterospecific battles and C. septempunctata won only the heterospecific battles against A. bipunctata. Coccinella septempunctata dropped from a leaf earlier and more often than the other two species but was in some cases able to return to the tree, especially under cloudy conditions. The frequency with which a species dropped did not depend on the species the larva was paired with. The results of these semi-field experiments confirm that H. axyridis is a strong intraguild predator as a consequence of its aggressiveness and good defence against predation from heterospecific species. The fact that H. axyridis is such a strong intraguild predator helps to explain its successful establishment as invasive alien species in Europe and the USA.     

异色瓢虫对与本土蚜虫共存的外来扶桑绵粉蚧的搜寻和捕食行为

[目的]外来入侵物种作为猎物可能影响本土广谱捕食者的搜寻和捕食行为.本研究旨在揭示异色瓢虫Harmonia axyridis对与本土豌豆修尾蚜Megoura japonica混合发生的外来入侵扶桑绵粉蚧Phenacoccus solenopsis的搜寻和捕食行为.[方法]试验前对异色瓢虫设两个饲喂猎物种类处理:用扶桑绵...     

Effect of cannibalism and predation on the larval performance of two ladybird beetles

The relative impact of cannibalism and predation on the development and survival of fourth instar larvae was assessed in two species of aphidophagous ladybird beetles, Coccinella septempunctata and Harmonia axyridis. The effect of eating aphids, conspecific larvae or heterospecific larvae on larval performance differed in the two species: aphids were the best food for C. septempunctata and survival of C. septempunctata larvae was significantly lower when offered heterospecific larvae rather than conspecific larvae or aphids as food, indicating that H. axyridis larvae were not suitable food for C. septempunctata. However, as the different foods did not affect the larval performance of H. axyridis, this species appears to be more polyphagous. Both intraguild predation by the aggressive larvae of H. axyridis and the polyphagous food habit of this species may account for its dominance in ladybird assemblages and its displacing other ladybird beetles in several places in the world.     

Effects of competition,cannibalism and intra‐guild predation on larval development of the European coccinellid Adalia bipunctata and the invasive species Harmonia axyridis

Abstract 1. Resource competition and intra‐guild predation (IGP) are important determinants of the structure of aphidophagous guilds. The likelihood and outcome of IGP is influenced by the density of extra‐guild prey and the characteristics of the species involved. 2. The nature of intra‐guild interactions between alien and indigenous coccinellids often determines the success of coccinellid invasions, as exemplified by Adalia bipunctata in Japan and Harmonia axyridis in North America and Europe. 3. Harmonia axyridis has negatively impinged on indigenous species in its introduced range, and its recent arrival in Britain poses a threat for members of native aphidophagous guilds. IGP and inter‐specific competition between H. axyridis and British coccinellids are predicted to occur. However, the results of such interactions have been little studied. 4. Here we investigate the effects of different diets, designed to mimic possible conditions in the wild, on the survival, development, and adult size of H. axyridis and A. bipunctata. Results demonstrate a skew in the consequences of IGP between the two species: the supplementation of a limited aphid diet with non‐conspecific eggs leads to a significant advantage for H. axyridis in respect of all parameters assessed, but gives no benefit to A. bipunctata. 5. We conclude that IGP of A. bipunctata by H. axyridis will contribute to the spread and increase of H. axyridis in Britain. 6. We further conclude that the skewed nature of IGP between A. bipunctata and H. axyridis at least in part explains the limited spread of A. bipunctata in Japan.     

Interaction of Acronicta rumicis (Lepidoptera: Noctuidae) and its larval parasitoid, Glyptapanteles liparidis (Hymenoptera: Ichneumonidae)

As a result of parasitism by Glyptapanteles liparidis in the first, second, third and fourth instar larvae of Acronicta rumicis, the mortality of each larval stage was found to be 46.67, 90, 71 and 16.67%, respectively. The mortality was highest when G. liparidis parasitized the second and third instar larvae. The difference in mortality between the parasitized group and the control group was 72.14% in the second instar larvae. With regards to the food consumption of the parasitized larvae, the first and second instar larvae consumed 6495.58 ± 646.52 mm² (leaf surface) and 7951.12 ± 4167.36 mm², respectively, while the third and fourth larvae consumed 13 826.77 ± 3396.66 mm² and 18 599.85 mm², respectively, showing that food consumption increased with instar stages of the host larvae. The clutch size of G. liparidis increased in relation to the instar stages of the host: it was 25.25 ± 7.89, 48.65 ± 53.75, 91.09 ± 44.52 and 114 individuals when they were fed with the first, second, third and the fourth instar larvae of the host, respectively.     

Effect of emigration on cannibalism and intraguild predation in aphidophagous ladybirds

Abstract.  1. The incidence and timing of emigration, cannibalism, and intraguild predation of larvae of three aphidophagous ladybirds (Coleoptera: Coccinellidae), Harmonia axyridis Pallas, Coccinella septempunctata brucki Mulsant, and Propylea japonica Mulsant, relative to the presence of prey was determined in the laboratory in single- and mixed-species populations.
2. In single-species populations, 80% of the larvae of C. s. brucki emigrated prior to the extinction of the aphid population and no larvae were lost due to cannibalism; however > 80% of the larvae of the other two species were still present when the aphid became extinct and the losses due to cannibalism for H. axyridis and P. japonica were 25% and 14% respectively. Finally, 28% of the P. japonica larvae completed their development, whereas no larvae of the other two species became adult.
3. In mixed-species populations, mortality of P. japonica attributable to cannibalism or intraguild predation increased greatly to 60%, whereas that of the other two species remained about the same. Consequently, survival of H. axyridis larvae improved and survival of P. japonica worsened; however the survival of C. s. brucki larvae was not affected by the other two species. Early emigration by C. s. brucki larvae may have enabled them to escape intraguild predation by H. axyridis in this system.     

Infectivity of Four Species of Nematodes (Rhabditoidea: Steinernematidae, Heterorhabditidae) to the Asian Longhorn Beetle, Anoplophora glabripennis (Motchulsky) (Coleoptera: Cerambycidae)

Four species of entomopathogenic nematodes, Steinernema carpocapsae , Heterorhabditis bacteriophora , H. indica and H. marelatus , were tested for their ability to kill and reproduce in larvae of the Asian longhorn beetle, Anoplophora glabripennis (Motchulsky). The larvae were permissive to all four species but mortality was higher and production of infective juveniles was greater for S. carpocapsae and H. marelatus . The lethal dosage of H. marelatus was determined to be 19 infective juveniles for second and third instar larvae and 347 infective juveniles for fourth and fifth instar larvae. H. marelatus infective juveniles, applied via sponges to oviposition sites on cut logs, located and killed host larvae within 30 cm galleries and reproduced successfully in several of the larvae.     

Relationships Between Attack and Escape Rates, Cannibalism, and Intraguild Predation in Larvae of Two Predatory Ladybirds

We compared incidences of cannibalism and intraguild (IG) predation (IGP) and quantified attack and escape rates—mechanisms which possibly account for the difference in incidences of these interactions—in laboratory experiments with Harmonia axyridis and Coccinella septempunctata. There was a tendency for H. axyridis to act as an IG predator and C. septempunctata as an IG prey. Cannibalism was also often observed in both species. The incidences of both IGP and cannibalism were different between the species. The average attack rates of C. septempunctata were less than 20%, but those of the more aggressive H. axyridis exceeded 50%. Larvae of both species attacked conspecifics and heterospecifics H. axyridis larvae successfully escaped when attacked by both conspecifics and by heterospecifics, while larvae of C. septempunctata escaped from attacks of conspecifics but not from those of heterospecifics. Thus the aggressive behavior of H. axyridis, in particular, of the third and fourth instars, negatively affects the larval survival of C. septempunctata. It may contribute to the dominance of H. axyridis in ladybird assemblages and its displacement of other ladybird species in several places in the world.     

Larval life history responses to food deprivation in three species of predatory lady beetles (Coleoptera: Coccinellidae)

We studied life history responses of larvae of three coccinellid species, Coleomegilla maculata (DeGeer), Hippodamia convergens Guerin-Meneville, and Harmonia axyridis (Pallas), when deprived of food for different periods of time during the fourth stadium. The coccinellid species did not differ in starvation resistance when larvae were starved throughout the stadium; however, for larvae fed only on day 1 of the stadium, H. convergens had the highest starvation resistance, followed by H. axyridis and then C. maculata. Percentage weight loss of larvae was affected by food deprivation period and coccinellid species. Both C. maculata and H. axyridis lost significantly more weight than H. convergens when starved throughout the fourth stadium. When deprived of food for 4 d of the stadium, C. maculata lost a higher percentage of initial body weight than H. axyridis. Percentage weight loss of H. convergens did not differ from that of C. maculata or H. axyridis. The weight of fourth instars and adults declined in an accelerating pattern as food deprivation period increased. However, food deprivation period had no significant effect on pupal development time for any of the three species or on larval development time for C. maculata and H. convergens. The increase in H. axyridis larval development time as a result of an increase in food deprivation period was curvilinear. Based on this laboratory study, it would seem that H. convergens is better able to cope with acute nutritional stress than either C. maculata or H. axyridis.     

Effect of temperature on the susceptibility of Culiseta longiareolata (Macquart) (Dipt., Culicidae) to two standard strains of biocontrol bacteria

The susceptibility of fourth instar larvae of Culiseta longiareolata (Macq.) to two standard strains of biocontrol bacteria was investigated under laboratory conditions. Larvae were found to be more susceptible to Bacillus sphaericus (Bs-IPS88) than to Bacillus thuringiensis H14 (Bti-IPS82) at 25 ± 1°C at concentrations ranging from 0.003 to 0.004 ppm at 2 and 3 days post-treatment. Statistical regression analysis revealed a significant mortality–concentration relationship for the two bacteria. Larval mortality increased significantly as temperature increased from 20 ± 1°C to 28 ± 1°C.     

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.     

Gypsy moth parasitoids in the declining outbreak in Lithuania

A 3 year study was conducted on the parasitoids of gypsy moth larvae in two reducing outbreak areas in Lithuania. Overall parasitism of 25.0 ± 2.0% in the first post-culmination year was significantly lower than the 36.3 ± 1.4 and 35.2 ± 1.4% parasitism in the two subsequent years. When analysed in terms of the life stage at which the host was collected, the total parasitism over 3 years was constantly increasing from 3.1 ± 0.8 in the first to 72.5 ± 2.9% in the sixth instar. Parasetigena silvestris R.-D. dominated causing 48.7 ± 1.5% parasitism and 16.7 ± 0.6% larval mortality preferably in late instars. Phobocampe disparis Vier. contributed to 21.9 ± 1.2% parasitism and 7.5 ± 0.5% mortality recovering from early instar larvae. Meteorus pulchricornis Wes. parasitized 4.3 ± 0.6% gypsy moth larvae causing 1.5 ± 0.2% mortality and a few Apanteles species provided 2.8 ± 0.6% parasitism and 1.0 ± 0.2% mortality. The gypsy moth in Lithuania was reported to act as host for Rogas sp. (Hym., Braconidae) and Siphona boreata Mes. (Dipt., Tachinidae).     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Potential for intraguild predation and competition among predatory Coccinellidae and Chrysopidae

In a laboratory study two coccinellid species, Coleomegilla maculata (DeGeer) and Harmonia axyridis Pallas, completed preimaginal development on lacewing eggs, Chrysoperla carnea Stephens or pea aphids, Acyrthosiphon pisum (Harris) in similar times. Preimaginal survival on C. carnea eggs was similar to survival on A. pisum for all stages of C. maculata and H. axyridis. Coccinellid adults that developed on C. carnea eggs were smaller than adults reared on A. pisum. Coccinella septempunctata L. did not complete preimaginal development on C. carnea eggs. Chrysoperla carnea preimaginal developmental time was approximately 20 days when fed either C. maculata eggs or A. pisum. Chrysoperla carnea fed C. maculata eggs developed into smaller adults, compared to adults reared as larvae on A. pisum, Ostrinia nubilalis (Hübner) eggs, or A. pisum alternated daily with O. nubilalis eggs. C. carnea did not complete preimaginal development on H. axyridis eggs. Cannibalism occurred more frequently between C. carnea third instars than between C. maculata fourth instars. When a C. carnea third instar was paired with a C. maculata fourth instar, more C. maculata were preyed upon by C. carnea, regardless of the herbivorous prey density. In the field these two predator species may negatively affect each other and reduce their suppression of pest densities.     

Influence of the host plant on occluded virus production and lethal infectivity of a baculovirus

Intra- and inter-specific effects of cotton, soybean, and clover on the time until death of Helicoverpa zea (Boddie) and Heliothis virescens (F.) larvae lethally infected with H. zea nucleopolyhedrovirus (HzSNPV) were evaluated in the laboratory. In the first test, on second instar only, the time until death of lethally infected larvae of both species differed with the plant tissues (vegetative or reproductive) and plant species. The total viral activity produced per larva in LC(50) units (occluded viral bodies (OBs) per larva/LC(50) in OBs/mm(2) of diet surface) was greater from H. virescens larvae fed vegetative than reproductive tissues of all host plants, but from H. zea virus production was greater only when fed vegetative tissue of soybean. In a second test that compared second and fourth instar H. virescens on cotton, total viral activity from larvae treated in both instars was greater when fed vegetative than reproductive tissues. Results of these tests suggest that the ability of host plants to influence baculovirus disease is more complex than previously believed. When examining the epizootic potential of a baculovirus, more attention must be given to the effects of the host plant on the insect-virus interactions.     

Experimental Infection of Nosema sp. Ghose in an Unnatural Host Lesioderma sericorne

ABSTRACT. Various doses of a microsporan parasite, Nosema sp., were fed to third and fourth instar larvae of Lesioderma sericorne that infested different types of stored grains. A spore dose of 3 × 10³ spores/individual resulted in a 39% infection rate, reduction in larval and adult weights, and mean spore concentrations of 1.28 ± 0.2 × 10⁸ spores/larva and 1.1 ± 0.2 × 10⁸ spores/adult. At the above dose, mortality was not well marked (about 35% in larvae and 25% in adults). At 3 × 10⁴ spores/individual, the rate of mortality increases to 80% in larvae and 60% in adults. However, more of the pest population (88% of larvae and 73% of adults) died at a dose of 3 × 10⁵ spores/individual. This dose produced mean spore concentrations of 3.91 ± 0.2 × 10⁸ spores/larva and 2.89 ± 0.2 × 10⁸ spores/adult. Insect death was caused by heavy damage to gut epithelia and fat bodies.