Suitability of exotic and native lady beetle eggs (Coleoptera: Coccinellidae) for development of lady beetle larvae

Predation upon lady beetle (Coleoptera: Coccinellidae) eggs in the field is most often instances of egg cannibalism by larvae or adults while the majority of the remaining predation events upon coccinellid eggs is done by other species of Coccinellidae. Thus the recent introduction and establishment of Harmonia axyridis in the US could negatively affect native species of Coccinellidae via egg predation. However, little is known regarding the suitability of interspecific coccinellid eggs as a food source for larval development. In this study, it was found that native first or third instar Coleomegilla maculata and Olla v-nigrum larvae were incapable of surviving to the adult stage when provided solely exotic H. axyridis eggs. In stark contrast, H. axyridis larvae survived equally well when cannibalizing eggs or eating eggs of either native species. When C. maculata and O. v-nigrum were grouped as ‘native’ and compared with the exotic H. axyridis, more native eggs were attacked than exotic eggs and a higher percentage of eggs was attacked by H. axyridis larvae. Native and exotic larvae attacked a similar percentage of native eggs but native larvae attacked significantly fewer exotic eggs than did exotic larvae. These data suggest that H. axyridis may prey upon the eggs of these native species, when encountered in the field, compared with the likelihood of the native species preying upon H. axyridis eggs. Therefore, eggs of the native species C. maculata and O. v-nigrum will continue to be subjected to cannibalism and also to possible predation by other native species and the exotic H. axyridis.     

Overwintering, phenology and fecundity of Harmonia axyridis in comparison with native coccinellid species in Italy

Classical biological control is generallyunderstood as an environmentally safe practiceof insect pest management. However, questionshave been raised about possible negativeeffects for native species. As part of a riskassessment study a semi-field trial was carriedout in northern Italy in order to compare theoverwintering of the native species Propylea quatuordecimpunctata (L.) and Adalia bipunctata (L.) (Coleoptera:Coccinellidae) with the exotic Harmoniaaxyridis (Pallas) (Coleoptera: Coccinellidae).All the experiments were carried out from April1998 to April 2001. Harmonia axyridis wasable to overwinter successfully in northernItaly. Fecundity, oviposition rate, longevityand rate of increase of overwintering femaleswere calculated and the phenology of the threespecies was studied. Harmonia axyridisoverwintering mortality (31.9%) was lower incomparison with mortality of native species(68.9% for P. quatuordecimpunctata and61.3% for A. bipunctata). As aconsequence, post-overwintering rate ofincrease of the exotic species was higher.Longevity of overwintered females was similaramong the species. Mean fecundity of H. axyridis (783.8 eggs per female) was slightlyhigher than A. bipunctata (720.2 eggs perfemale), and much higher than P. quatuordecimpunctata (193.7 eggs per female). H. axyridis and A. bipunctatacompleted four generations and P. quatuordecimpunctata three generations in a year. The results show that H. axyridisappears to posses a high potential for establishment in Italy.     

Interactions between the fungal pathogen Beauveria bassiana and three species of coccinellid: Harmonia axyridis, Coccinella septempunctata and Adalia bipunctata

Harmonia axyridis is a predatory coccinellid, native to central and eastern Asia. It has been available in many countries for use as a biological control agent of pest aphids and scale insects. In many of these countries, including the USA, H. axyridis has established. It is now considered an invasive alien species for a number of reasons, including its impact on functional biodiversity. Beauveria bassiana is known to be a natural mortality agent of overwintering coccinellids and is a potential candidate for the biological control of H. axyridis. In this paper we compare the susceptibility of three species of coccinellid, H. axyridis (cultures derived from Japan and UK), Coccinella septempunctata and Adalia bipunctata to infection by B. bassiana (commercial strain GHA) after exposure at three doses (10⁵, 10⁷, 10⁹ conidia ml⁻¹). The two subpopulations of H. axyridis (Japan and UK) were more resistant to B. bassiana infection than either A. bipunctata or C. septempunctata. This is exemplified by the median lethal doses at 10 days post-inoculation (LD₅₀) of 10^6.2, 10^6.0, 10^8.3, 10^9.6 conidia ml⁻¹ for A. bipunctata, C. septempunctata, H. axyridis (Japan) and H. axyridis (UK), respectively. Only doses of 10⁹ conidia ml⁻¹ resulted in mortality of H. axyridis, in contrast, 80% of C. septempunctata and 70% of A. bipunctata exposed to 10⁷ conidia ml⁻¹ of B. bassiana succumbed to infection. The fecundity (cumulative mean egg production over 10 days) of A. bipunctata and H. axyridis (UK) was also assessed. The fecundity of C. septempunctata could not be assessed because this species requires diapause prior to the onset of reproduction and these studies were on beetles that had recently eclosed (2–8 weeks). Harmonia axyridis (Japan) produced no eggs in most treatments including the control and so was excluded from analysis. High dose (10⁹ conidia ml⁻¹) inoculation reduced the fecundity of A. bipunctata to zero but egg production was similar for individuals inoculated with doses of 10⁵, 10⁷ conidia ml⁻¹ and control individuals. In contrast, all doses of B. basssiana reduced H. axyridis (UK) egg production dramatically. We discuss these results in relation to the potential for control of H. axyridis using B. bassiana.     

Laboratory study of cannibalism and interspecific predation in ladybirds

Abstract.

• 1 In the absence of aphids, adult females of Adalia bipunctata (L.) showed a greater reluctance to eat eggs than males.
• 2 Eggs and young larvae were more vulnerable to cannibalism than older larvae and starved larvae were more vulnerable than well-fed larvae.
• 3 Both egg and larval cannibalism is inversely related to the abundance of aphids.
• 4 Eggs are a better food, in terms of larval growth and survival, than aphids.
• 5 In the absence of aphids interspecific predation occurred, but not equally, between the coccinellids A.bipunctata, A.decempunctata (L.), Coccinella septempunctata L. and C.undecempunctata L.
• 6 Larvae and adults of A. bipunctata and C.septempunctata were reluctant to eat conspecific eggs painted with a water extract of the other species' eggs and larvae of C. septempunctata were more likely to die after eating a few eggs of A.bipunctata than vice versa.
• 7 These results indicate that cannibalism occurs mainly when aphid prey is scarce and is adaptive in that it improves the chances of survival, and coccinellids, to varying degrees, are defended against interspecific predation.

     

Do microsporidia function as “biological weapon” for Harmonia axyridis under natural conditions?

Invasive alien species, such as the multicoloured Asian ladybird Harmonia axyridis, are often regarded as major drivers of biodiversity loss. Therefore understanding which characteristics or mechanisms contribute to their invasive success is important. Here the role of symbiotic microsporidia in the hemolymph of H. axyridis was investigated in the context of intraguild predation between wild‐caught H. axyridis and the native ladybird species Coccinella septempunctata. The microsporidia were recently discussed to contribute to the unpalatability of Harmonia for other coccinellids during intraguild predation and to function as “biological weapons”. In the present study, visual detection of microsporidia in hemolymph samples revealed that 73.5 % of H. axyridis were infected. Intraguild predation experiments between larvae of the two species showed a significant competitive advantage for H. axyridis, even against larger larvae of C. septempunctata. Adult C. septempunctata always killed and fed on H. axyridis larvae. However only 11.4 % (4 of 47) of C. septempunctata that fed on infected H. axyridis died within 4 months. In contrast to previous studies this suggests that microsporidia or harmonine, the chemical defense compound of H. axyridis, do not lead to death of C. septempunctata preying on larvae of H. axyridis. Instead our results support the idea that competitive advantage during intraguild predation greatly facilitates the success of H. axyridis and that this may help this highly invasive species to outcompete native species. The impact of microsporidia on Harmonia itself as well as on interspecific interactions require further studies.     

三种瓢虫集团内捕食卵对初龄幼虫生长发育的影响

种内和种间捕食卵现象常见于对食蚜瓢虫的研究报道中。源自亚洲、入侵北美和欧洲的异色瓢虫(Harmonia axyridis)由于卵含有生物碱而被认为可能是威胁本土瓢虫生存的原因,但该推测只有在明确异色瓢虫在原产地是否对本土瓢虫具有类似负面影响后,才能确证。通过饲喂实验,旨在明确异色瓢虫在原产地是否对本土七星瓢虫(Coccinella septempunctata)和龟纹瓢虫(Propylea japonica)具有食卵负面作用,为揭示异色瓢虫入侵机理提供依据。实验中用蚜虫(对照)和3种瓢虫卵对瓢虫初孵1龄幼虫进行饲喂处理,观察幼虫生长发育指标。对瓢虫初龄幼虫存活表现的分析结果表明:(1)龟纹瓢虫初孵幼虫取食同种卵和七星瓢虫卵后存活至2龄的比率均高于取食异色瓢虫卵的近7倍;(2)七星瓢虫幼虫取食异色瓢虫卵未存活到2龄,但取食其他瓢虫卵和蚜虫后85%以上个体存活至2龄;(3)异色瓢虫初孵幼虫取食同种和其他2种瓢虫卵后有90%以上存活至2龄。龟纹瓢虫和七星瓢虫1龄幼虫取食异种卵比同种卵后的发育历期显著延长,但异色瓢虫并未表现出差异。龟纹瓢虫和异色瓢虫1龄幼虫取食同种和异种卵后的体增重相同,但七星瓢虫取食龟纹瓢虫卵后体增重比取食同种卵减小。研究结果说明,异色瓢虫卵对七星瓢虫和龟纹瓢虫初孵幼虫的生存具有负面影响,反之不然。所以,取食异色瓢虫卵本身并非是导致北美和欧洲本土瓢虫数量减少的原因。     

Why do ladybirds (Coleoptera: Coccinellidae) cannibalize?

Cannibalism of eggs by larvae ofAdalia bipunctata, an aphidophagous species of ladybirds, is important for survival when aphids are scarce. Ladybirds survive longer by eating eggs of their own species rather than aphids. Since it costs less, in terms of larval growth, to eat eggs rather than aphids, cannibalism has a strong advantage under conditions of prey scarcity.     

Influence of the prey aphid Uroleucon nigrotuberculatum parasitizing Solidago canadensis on the larval and adult survivorship of the predatory ladybird beetle Harmonia axyridis

The ladybird beetle Harmonia axyridis is an aphidophagous natural enemy in Japan. The alien plant Solidago canadensis is widespread in Japanese agricultural landscapes, and the alien aphid Uroleucon nigrotuberculatum parasitizes the plant. We examined the quality of the aphid as food for all developmental stages of H. axyridis. First-instar larvae fed the aphid died within 2 days. Larvae at later developmental stages survived for significantly longer periods but died before pupation. Adults fed the aphid also died within 10 days. These results suggest that U. nigrotuberculatum parasitizing S. canadensis is unsuitable for H. axyridis during the beetle’s entire life cycle.     

Negative dietary effects of Colorado potato beetle eggs for the larvae of native and introduced ladybird beetles

Aphidophagous ladybird beetles (Coleoptera: Coccinellidae) are attracted to and feed heavily on aphids, but many species will also feed opportunistically on other prey that they encounter. In potatoes (Solanum tuberosum L.) in Washington State, USA, coccinellids feed on both green peach aphids (“GPA,” Myzus persicae Sulzer) and eggs of the Colorado potato beetle (“CPB,” Leptinotarsa decemlineata Say). The guild of aphidophagous ladybirds includes two native species, Hippodamia convergens Guérin-Méneville and Coccinella transversoguttata Brown. Recently, an introduced species, Coccinella septempunctata L., has invaded and apparently displaced its native congener. A second exotic, Harmonia axyridis Pallas, has colonized the area and is becoming more abundant. We compared larval development of each species on a monotypic diet of GPA, a monotypic diet of CPB eggs, or a mixed diet of both GPA and CPB eggs. Our goal was to answer two questions: (1) do larvae of the four ladybird species benefit from including CPB eggs in their diet and (2) do the four ladybird species differ in their ability to utilize CPB eggs as prey? No larva of any species completed development on a pure diet of CPB eggs, and survivorship was highest for all species when they fed on a pure diet of GPA. One native species, H. convergens, and one exotic species, H. axyridis, exhibited significantly lower survivorship on a mixed diet of both CPB eggs and GPA, compared to a pure GPA diet; H. axyridis also took longer to develop from egg to adult when both prey were provided. Survivorship of the two Coccinella spp. was not altered by the inclusion of CPB eggs with GPA, although CPB eggs lengthened the development time of C. transversoguttata. Adult size was not consistently affected by diet for any of the coccinellids. Overall, no ladybird species benefited from the inclusion of potato beetle eggs in its diet. The two Coccinella species responded similarly to the inclusion of CPB eggs, and so we would not expect any difference in the success of coccinellid larval development in potato fields following the replacement of C. transversoguttata by C. septempunctata. Hippodamia convergens and H. axyridis, the two species whose survivorship was depressed by combining CPB egg and aphid prey, were also the two species that consumed the greatest number of CPB eggs during successful larval development. A comparison of total egg consumption by each species cohort suggested that displacement of the other species by H. axyridis would not alter CPB biological control, because the higher per capita feeding rate by H. axyridis larvae compensated for individuals’ greater mortality risk on a diet including CPB eggs.     

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.     

Effects of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae), on three non-target coccinellids

A microsporidium from Hippodamia convergens was transmitted horizontally to three non-target coccinellid hosts (Adalia bipunctata L., Coccinella septempunctata L. and Harmonia axyridis Pallas) under laboratory conditions. For all species examined, microsporidia-infected larvae took significantly longer to develop than did uninfected larvae but the microsporidium had no effect on larval mortality. Adult sex ratios of uninfected and microsporidia-infected adults were about 1:1 (♀:♂) and did not differ significantly. At the end of a 90-day trial, microsporidia-infected H. convergens produced significantly fewer eggs and did not live as long as uninfected individuals. Differences in fecundity and longevity were not observed for the three non-target coccinellids that were examined. Mean spore counts from smear preparations of microsporidia-infected A. bipunctata did not differ significantly from H. convergens, suggesting that A. bipunctata (a native coccinellid) is a suitable host for the microsporidium but infection was lighter in C. septempunctata and H. axyridis (introduced species). Vertical transmission of the pathogen was observed during the 90-day trial by examining eggs and larvae that were produced by microsporidia-infected adults. For all species examined, 100% vertical transmission of the pathogen was eventually observed. Three eugregarines were found in two adult A. bipunctata: Gregarine A trophozoites are similar in size to those of Gregarina katherina Watson (described earlier from Coccinella spp.), Gregarine B trophozoites are similar in size to those of Gregarine A but are morphology distinct, and Gregarine C trophozoites are similar in size to G. barbarara Watson (described earlier from A. bipunctata).     

Development and application of molecular gut‐content analysis to detect aphid and coccinellid predation by Harmonia axyridis (Coleoptera: Coccinellidae) in Italy

Despite their positive effect in reducing pest populations, exotic generalist predators sometimes become invasive and contribute to the displacement of indigenous species in the same trophic level. Although laboratory experiments have linked intraguild predation (IGP) to these interactions, field evidence and quantification of IGP are still lacking for most systems. The recent establishment of the exotic Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Italy raises concern about the detrimental effect that the ladybird could have on native coccinellids. Here we assessed, under laboratory conditions, the acceptability and suitability of eggs of 2 native ladybirds, Adalia bipunctata L. and Oenopia conglobata (L.), as prey items for H. axyridis larvae. Then we developed primers for molecular gut‐content analysis to detect predation by H. axyridis on the 2 ladybirds and on the aphid Eucallipterus tiliae L. Species‐specific 16S primers were developed for the 3 species and laboratory feeding trials were conducted to quantify the rate of prey DNA breakdown in the gut of H. axyridis. Moreover, to field evaluate primers, H. axyridis 4th instars (n = 132) were systematically collected from linden trees in northern Italy and screened for the presence of prey DNA. Seventy‐three percent and 7% of field collected H. axyridis were positive for aphid and coccinellid DNA, respectively. Predation upon aphid and A. bipunctata was lower than predicted if density dependent consumption was expected, while predation upon O. conglobata was significantly higher. Here, we provided the first evidence of IGP among feral populations of H. axyridis and indigenous ladybird beetles, occurring in Italy.     

Habitat or prey specialization in predatory ladybird beetles: a case study of two closely related species

This study examines the niche and diet breadth of two closely related sympatric aphidophagous ladybirds: Adalia decempunctata and A. bipunctata. The degree of habitat specialization of these species is investigated, and its effect on life history traits of females is explored. The importance of prey quality in determining the diet breadth is also examined. The niches occupied by these species in three countries, the UK, Belgium and southern France, are similar: A. decempunctata is an arboreal habitat specialist with a narrower set of prey than A. bipunctata, which is commonly found on several types of vegetation. The niches of the two species overlap on trees. Experiments indicate that habitat specialization has resulted in A. decempunctata investing more in each of its offspring than A. bipunctata. A. decempunctata females lay, relative to their body size, heavier eggs than those of the more generalist A. bipunctata, which results in A. decempunctata having bigger larvae. In addition, A. decempunctata larvae are better at surviving starvation than A. bipunctata larvae. In contrast to the expected pattern in food specialization, our study failed to demonstrate a better efficiency of the specialist when fed its usual prey and a detrimental effect when fed on prey that it is unlikely to encounter in the field. The reproductive performance of the specialist ladybird was better when fed an aphid that it was unlikely to regularly feed on in the field. Therefore, the narrow diet of the specialist ladybird is most likely a consequence of it occupying a narrow habitat rather than the quality of the prey. Although further studies on specialization in predatory insects are needed, the results indicate that unlike the role of plant quality in host specialization in herbivorous insects, prey quality has not been the main determinant of ecological specialization in these predatory insects.     

Interactions between the parasitoid wasp Dinocampus coccinellae and two species of coccinellid from Japan and Britain

The establishment, spread and increase of the invasive coccinellid Harmonia axyridis Pallas in North America and Britain have coincided with declines in native ladybirds. In Britain, this pattern was predicted soon after H. axyridis was first recorded. However, predictions of the impact that H. axyridis may have on the parasites, parasitoids and pathogens of native coccinellids have been less certain, largely because of uncertainty over whether H. axyridis would become an alternate host for such agents. Dinocampus coccinellae (Schrank) is a braconid parasitoid of ladybirds of the sub-family Coccinellinae. In Japan, this wasp uses both Coccinella septempunctata brucki Mulsant and H. axyridis as hosts, but successfully parasitizes a higher proportion of the former species. Data are here presented that indicate the same is true in Britain, but to a greater extent. This study investigates the interactions of D. coccinellae with C. septempunctata L. and H. axyridis from Japan and Britain. We show that coccinellid activity affects encounter rates with the wasp, with H. axyridis being more active than C. septempunctata in the tests. Escalation rates from investigation to ovipositional attack were higher for C. septempunctata than H. axyridis for both Japanese and British wasps. Wasp emergence was higher, relative to ovipositional attack, from C. septempunctata than from H. axyridis. Wasps did not discriminate between Japanese and British ladybirds of the same species. British wasp eclosion rates were higher from Japanese than from British H. axyridis. We consider both the effect D. coccinellae may have on H. axyridis population demography, and the effect H. axyridis may have on D. coccinellae in Europe.     

Intraguild predation and successful invasion by introduced ladybird beetles

Introductions of two ladybird beetle (Coleoptera: Coccinellidae) species, Coccinella septempunctata and Harmonia axyridis, into North America for aphid biocontrol have been followed by declines in native species. We examined intraguild predation (IGP) between larvae of these two exotic species and larvae of the two most abundant native coccinellids in eastern Washington State, C. transversoguttata and Hippodamia convergens. In pairings between the two native species in laboratory microcosms containing pea (Pisum sativum) plants, neither native had a clear advantage over the other in IGP. When the natives were paired with either Harmonia axyridis or C. septempunctata, the natives were more frequently the victims than perpetrators of IGP. In contrast, in pairings between the exotic species, neither had an IGP advantage, although overall rates of IGP between these two species were very high. Adding alternative prey (aphids) to microcosms did not alter the frequency and patterns of relative IGP among the coccinellid species. In observations of encounters between larvae, the introduced H. axyridis frequently survived multiple encounters with the native C. transversoguttata, whereas the native rarely survived a single encounter with H. axyridis. Our results suggest that larvae of the native species face increased IGP following invasion by C. septempunctata and H. axyridis, which may be contributing to the speed with which these exotic ladybird beetles displace the natives following invasion.     

Ladybird population dynamics in potato: comparison of native species with an invasive species, Harmonia axyridis

Following the detection of the harlequin ladybird, Harmonia axyridis, in 2003 in potato crops in Belgium, a study was carried out between 2004 and 2006 on the phenology of this species compared to native species in potato. The results confirmed the success of H. axyridis, with high population levels in 2004 and 2005. In 2006, aphid populations were very low and no H. axyridis larvae were sampled in potato, but the indigenous species Coccinella septempunctata and Propylea quatuordecimpunctata were detected. A species by species comparison of the date of first larvae detection, the larvae population peak, and the difference between this peak and the aphid population peak was performed. Results showed a clear correlation between C. septempunctata and P. quatuordecimpunctata and potato aphids, with a delay of 3.5 and 6.5 days between the aphid and ladybird population peaks for the two native species. H. axyridis arrived 7–8 days after the two indigenous species and the larval peak population occurred 15.8 days after the aphid population peak. This meant that H. axyridis had to complete its larval development with very low aphid populations or even with no aphids at all. The reason for its late arrival and the possible food resources used by H. axyridis larvae are discussed.     

Dispersal potential of native and exotic predatory ladybirds as measured by a computer-monitored flight mill

The performance of three species of predatory ladybirds was compared in a flight mill and the effect of diet on their flight parameters was tested. The invasive ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) outperformed Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) and Adalia bipunctata (L.) (Coleoptera: Coccinellidae) in terms of flight distance, duration and velocity. Harmonia axyridis flew at least two times further, needed three times less breaks and flew two times faster than C. montrouzieri and A. bipunctata fed the same diet. Ladybirds reared on eggs of Ephestia kuehniella (Zeller) (Lepidoptera: Pyralidae) performed better than their counterparts reared on natural prey (aphids for H. axyridis and A. bipunctata, mealybugs for C. montrouzieri). The findings of this study indicate that comparative flight studies can be useful to identify candidate biocontrol agents with pronounced dispersal abilities and thus can yield significant evidence to be used in an environmental risk assessment. However, it also demonstrates that variability related to mass rearing conditions should not be ignored when standardizing a risk assessment procedure for candidate biocontrol agents.     

Toxicity of the aphid Aulacorthum magnoliae to the predator Harmonia axyridis (Coleoptera: Coccinellidae) and genetic variance in the assimilation of the toxic aphids in H. axyridis larvae

Larvae of the ladybird Harmonia axyridis are reported to suffer high mortality when they are fed with the aphid Aulacorthum magnoliae associated with the elder Sambucus sieboldiana. In the present study we first examined whether aphid toxicity to H. axyridis was altered when the aphids were reared on different host plants, and second whether some ladybird families could adapt specially to the toxic aphids. Ladybird larvae from each egg batch were divided into two groups; one group was fed with A. magnoliae reared on potato, and the other group was fed with A. magnoliae reared on elder. The ladybird larvae fed with elder aphids suffered significantly higher mortality and grew more slowly than did larvae fed with potato aphids. This result indicates that A. magnoliae aphids absorbed toxic substances or their precursors from S. sieboldiana. We suggest that host association of A. magnoliae with the primary host plant S. sieboldiana has been maintained because of the advantage of deterring predation. Significant and positive correlation was detected across H. axyridis sib families between the mean weight of larvae fed with elder aphids and the mean weight of larvae fed with potato aphids. The hypothesis that some ladybird families adapt specially to the toxic aphids was not supported. However, the available evidence showed that a large amount of genetic variance in performance is maintained in a wild population of the ladybird H. axyridis.     

Chemical protection of Calvia quatuordecimguttata eggs against intraguild predation by the invasive ladybird Harmonia axyridis

Intraspecific and interspecific predation of eggs is a well documented phenomenon amongst aphidophagous coccinellids. The invasive species Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is known to be a top intraguild predator and reported to attack the eggs of many coccinellid species both in a laboratory setting and in the wild. A previous laboratory study highlighted that while many species’ eggs were highly palatable to H. axyridis, the eggs of Calvia quatuordecimguttata (Linnaeus) (Coleoptera: Coccinellidae) appeared to be extremely well protected from attack. Here we present the results of behavioural experiments testing the hypothesis that substances on the egg surface are responsible for this protection, and report preliminary results of GC-MS analysis of these compounds. When the coatings of C. quatuordecimguttata eggs were removed using hexane, they became significantly more susceptible to predation by neonate H. axyridis larvae. However, their overall palatability was not affected, in that complete consumption was never or rarely observed. This suggests that the surface compounds are a true indicator of unpalatability in this species. The effect of hexane-washing on already palatable conspecific eggs was also analysed but had no significant effect on the susceptibility of eggs to cannibalism. We conclude that the eggs of at least one European species are effectively protected by surface deterrents from intraguild predation by H. axyridis. This effect might be due to both the diversity and abundance of hydrocarbons present within the egg coating, the presence of alkenes and/or the presence of patches of a red substance on the eggs’ surface, which is thought to belong to the acid group. In conjunction with data on the susceptibility of other immature stages of C. quatuordecimguttata, this finding may indicate a decreased risk of the species falling victim to invasive H. axyridis, despite their coincident habitat ranges.     

The influence of density-dependent factors on larval development in native and invasive populations of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pall.) (Coleoptera,Coccinellidae)

Effects of the number of larvae per Petri dish (1, 5, and 10) on the preimaginal development of individuals of the native (Irkutsk, southern Siberia) and invasive (Sochi, the Northern Caucasus) populations of the multicolored Asian ladybird Harmonia axyridis were investigated in the laboratory. The experiments were conducted under short (12 h) and long (18 h) day conditions; the larvae were fed on the green peach aphid Myzus persicae or on the eggs of the grain moth Sitotroga cerealella. An increase in the number of larvae developed in one Petri dish resulted in a significant decrease in the rate of development in individuals from both populations which fed on aphids. Survival decreased with an increase in the number of larvae developed in one Petri dish fed on both prey species, but only in larvae from the invasive population of H. axyridis. The weight of emerging adults decreased with the number of larvae per dish in individuals from both study populations, but only when fed on aphids. These data suggest that the influence of density-dependent factors on the development of H. axyridis depends significantly on larval prey species. In addition, larvae from the invasive population have somewhat more aggressive interactions with competitors, this possibly having been one of the prerequisites for invasion.