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.     

Harmonia axyridis in Great Britain: analysis of the spread and distribution of a non-native coccinellid

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe H. axyridis is considered to be an invasive alien species. Although not known to have been deliberately introduced to Great Britain, it was first recorded there in 2004, in south-east England. Harmonia axyridis arrived in Great Britain by various means, primarily from mainland Europe, but also from Canada. Extensive national and local media coverage, and a dedicated website (), facilitated public involvement in recording H. axyridis in Great Britain; in excess of 4,000 verified records of the species were received between 2004 and 2006. Through detailed mapping, the objective of our study was to quantify and analyse the spread of H. axyridis in its early stages of invasion in Great Britain. Our data shows that between 2004 and 2006, the species spread north through Great Britain at the rate of 58 km year^-1 and west at the rate of 144.5 km year^-1. In England H. axyridis spread north to Yorkshire and west to Cornwall by the end of 2006, and also reached Wales. Recorded occurrence (of one or more H. axyridis individuals at larval, pupal and/or adult stage) in 10 km squares in Great Britain was: 2004—51; 2005—149; 2006—447. Records of juvenile H. axyridis extend through most of the recorded British range of the species, and we present evidence of bi-voltinism in the population in 2006.     

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.     

Harmonia axyridis in Europe: spread and distribution of a non-native coccinellid

Native to Asia, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is considered an invasive alien ladybird in Europe and North America, where it was widely introduced as a biological control agent of aphids and coccids. In Europe, H. axyridis was sold by various biological control companies from 1995 in France, Belgium and the Netherlands, and was also intentionally released in at least nine other countries. It has spread very rapidly, particularly since 2002, and is now regarded as established in thirteen European countries. The established range extends from Denmark in the north to southern France in the south, and from Czech Republic in the east to Great Britain in the west. In this paper we map the spread and distribution of H. axyridis in Europe, and examine the situation on a country-by-country basis. We report first records of the species in five countries; Spain, Sweden, Denmark, Czech Republic and Italy; and first evidence of H. axyridis establishment in the latter three countries. Despite releases of H. axyridis in Portugal, Spain and Greece, there is little evidence of establishment in southern Europe. It is predicted that the spread and increase within Europe will continue and that H. axyridis will become one of the most widely distributed coccinellids in the continent.     

Estimation of mortality by entomophages on exotic Harmonia axyridis versus native Adalia bipunctata in semi-field conditions in northern Italy

A semi-field experiment was carried out in two peach orchards in northern Italy to assess mortality due to predators and parasitoids on the exotic coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in comparison with the native coccinellid Adalia bipunctata L. (Coleoptera: Coccinellidae). The experiments were conducted in cages to avoid the possible escape of the exotic ladybird (not yet established in Italy). Two kinds of cage experiments were included: ‘exclusion cages’ (access by walking predators impeded) and ‘free cages’ (walking predators free to enter). The cages, containing all the stages of the two ladybird species, were placed in two localities and left for 24 h. All ladybird stages used for the semi-field experiments came from a laboratory rearing. The eggs of H. axyridis experienced less mortality than those of A. bipunctata. The ant workers were the most frequent predators in ‘free cages’ but A. bipunctata cannibalism on eggs was also detected. Larvae of both coccinellid species were predated equally but larval predation of L1 and L2 was higher in comparison to predation of L3 and L4. Pupae and adults of both exotic and native ladybirds were never attacked by predators. Predation on younger larval stages was higher in the ‘free cages’ in comparison with ‘exclusion cages’. No ladybird parasitisation was observed. The ‘free cage’ technique seems to provide a standardised and realistic estimation of predation impact but more studies are needed to evaluate ladybird parasitisation in semi-field conditions.     

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.     

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.     

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 responses of aphid predators on apple to the invasion of an exotic species, Harmonia axyridis

The effects of the invasion ofan exotic predator, Harmonia axyridis(Pallas) (Coleoptera: Coccinellidae), wereinvestigated using three experiments on theecology of aphid predators on apple. Oneexperiment, 1992, was collected prior to theH. axyridis invasion, and two others,1996 to 1997 and 1999 to 2000, were collectedafter the invasion. Except for one year, 1999,H. axyridis was the dominant coccinellid,replacing the formerly dominant Coccinellaseptempunctata L. (Coleoptera: Coccinellidae),another exotic species. The dominance of H. axyridis was greater among larvae thanamong adults. There was no apparent effect ofthe H. axyridis invasion on abundance ofthe predator, Aphidoletes aphidimyza(Rondani) (Diptera: Cecidomyiidae) and apossible positive effect on the abundance ofchrysopids (Neuroptera: Chrysopidae). Principal component analysis indicated thatalthough individual species were affected, theoverall effect of H. axyridis invasion onthe predator guild as a whole was negligible. The data indicate that the interaction betweenthe two exotic species, H. axyridis andC. septempunctata, may be allowing nativecoccinellids to become more abundant on applethan when C. septempunctata was thedominant coccinellid.     

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.     

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

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

Intraguild predation of immature stages of British and Japanese coccinellids by the invasive ladybird Harmonia axyridis

Declines in native aphidophages in North America have been linked to intraguild predation (IGP) by the invasive coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). It is feared that many British species will face a similar fate following the recent establishment of H. axyridis in the UK. Meanwhile, H. axyridis exists in apparent ecological equilibrium with other members of its guild in Japan. The impact of H. axyridis on British coccinellids is uncertain but intraguild predatory interactions do occur, particularly amongst immature stages. This study investigates IGP between immature stages of H. axyridis and various British and Japanese coccinellids. The only asymmetric IG predator of H. axyridis at first instar was Anatis ocellata (Linnaeus). Harmonia axyridis engaged in symmetric IGP with Coccinella septempunctata Linnaeus, Calvia quatuordecimguttata (Linnaeus), Harmonia quadripunctata (Pontoppidan) and Eocaria muiri Timberlake, but was the asymmetric IG predator of all other species studied. The level of IGP was high between fourth instar larvae, and frequently biased towards H. axyridis, except in the case of A. ocellata, which again was the only IG predator of H. axyridis. In interactions between fourth instar larvae and pre-pupae, IGP was unidirectional towards H. axyridis for all species except A. ocellata, which acted as both IG predator and IG prey. Pupae were better protected against IGP than pre-pupae but most species were still susceptible to attack by H. axyridis, although IGP was symmetric with A. ocellata, and H. quadripunctata pupae were never attacked. The differences in susceptibility of the various species and developmental stages to IGP by H. axyridis are discussed in relation to physical defence structures. We find no evidence that Japanese species have superior defences to British ones and suggest that behavioural strategies may enable co-existence in the native range. We discuss the relevance of IGP by H. axyridis to the species it is likely to encounter in Britain.     

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.     

Influence of six aphid prey species on development and reproduction of a ladybird beetle, Coccinella septempunctata

Pre-imaginal development, immaturesurvival, and reproduction of a ladybird, Coccinella septempunctata Linnaeus, werestudied in response to six aphid species, Aphiscraccivora Koch, Aphis gossypii Glover,Aphis nerii Boyer de Fonscolombe,Lipaphis erysimi (Kaltenbach), Myzuspersicae (Sulzer) and Uroleuconcompositae (Theobald) to quantify theirrelative suitability as prey. Pre-adultdevelopment was shortest (13.93 ± 0.12 days)when fed on L. erysimi and longest(22.85 ± 0.10 days) on A. nerii. Immaturesurvival, adult emergence, growth index,relative growth rate, development rate, maleand female longevity, oviposition period,fecundity and hatching percent were maximal, i.e. 73.47 ± 0.89%, 90.07 ± 1.43%,8.62 ± 0.23, 1.52 ± 0.02, 0.07,81.10 ± 1.26 days, 85.70 ± 1.45 days,69.80 ± 1.32 days, 1764.10 ± 8.46,and 87.88 ± 1.05, respectively when C.septempunctata were fed on L. erysimi.The same parameters were minimal, i.e.43.86 ± 1.33%, 71.65 ± 2.75%,2.02 ± 0.08, 0.49 ± 0.02, 0.04,44.40 ± 1.39 days, 53.50 ± 1.00 days,16.40 ± 0.60 days, 203.20 ± 11.83, and48.68 ± 2.06, respectively on A. nerii. Theweights of different ladybird life stages weremaximal after feeding on L. erysimi and minimalon A. nerii. Regression analyses of thedata revealed linear relationships betweendevelopment rate and weight of adult; dailyprey consumption and relative growth rate; logweight of adult male and female; and longevityand fecundity of female. On the basis of thesefindings, the order of suitability of aphidspecies for C. septempunctata is L. erysimi >M. persicae > A. craccivora > A. gossypii >U. compositae > A. nerii. Thus, the presentinformation can be utilized for the massrearing of C. septempunctata by supplyingthe best food and can also help in theprediction of the relative abundance of theladybird on different aphid infestations in thefields.     

Expansion of the geographical distribution of an exotic ladybird beetle, Adalia bipunctata (Coleoptera: Coccinellidae), and its interspecific relationships with native ladybird beetles in Japan

The first record of the exotic ladybird beetle, Adalia bipunctata (Coleoptera; Coccinellidae), in Japan was in 1993 at Osaka Nanko Central Park. Since that time, studies on the life history and geographical distribution of A. bipunctata have been ongoing, and its establishment in the Osaka Nanko area has been confirmed. A. bipunctata is a predacious ladybird beetle and a member of a guild that overlaps in habitat and prey with that of native ladybird beetles such as Harmonia axyridis and Menochilas sexmaculatus. We investigated the distribution of A. bipunctata and its interspecific relationships with native predacious ladybird beetles. In some areas, A. bipunctata was dominant in interspecific relationships with native ladybird beetles. For the first 10 years after A. bipunctata was discovered, it occurred only in the Osaka Nanko area, but the present geographical distribution indicates that it has expanded its range. Though the population density of this species was highest at the area recorded first, and tended to decrease in inverse proportion to the distance from Osaka Nanko Central Park, a satellite occurrence was observed in a remote area. The numbers of aphid and tree species (leaf shelter for aestivation and over-wintering) utilized by A. bipunctata have recently increased. Such increases will cause the rate of distribution of A. bipunctata to accelerate. Interspecific competition between H. axyridis and A. bipunctata, which occurs earlier than H. axyridis, may be avoided by desynchronization of the occurrence seasons, and another common predacious ladybird beetle, Coccinella septempunctata, may escape interspecific competition by habitat segregation.     

Predicting the potential geographical distribution of the harlequin ladybird, Harmonia axyridis, using the CLIMEX model

Harmonia axyridis (Pallas, 1773) (Coleoptera: Coccinellidae) is a ladybird beetle native to temperate and subtropical parts of Asia. Since 1916 populations of this species have been introduced throughout the world, either deliberately, or by accident through international transport. Harmonia axyridis was originally released as a classical biological control agent of aphid and coccid pests in orchards and forests, but since 1994 it is also available as a commercial product for augmentative control in field and greenhouse crops. It is a very voracious and effective natural enemy of aphids, psyllids and coccids in various agricultural and horticultural habitats and forests. During the past 20 years, however, it has successfully invaded non-target habitats in North America (since 1988), Europe (1999) and South America (2001) respectively in a short period of time, attacking a wide range of non-pest species in different insect orders. Becoming part of the agricultural commercial pathway, it is prone to being introduced into large areas across the world by accident. We use the CLIMEX programme (v2) to predict the potential geographical distribution of H. axyridis by means of matching the climate of its region of origin with other regions in the world and taking in account biological characteristics of the species. Establishment and spread seem likely in many regions across the world, including those areas which H. axyridis has already invaded (temperate Europe, North America). Based on the CLIMEX prediction a large part of Mediterranean Europe, South America, Africa, Australia and New Zealand seem highly suitable for long-term survival of H. axyridis as well. In addition we evaluate CLIMEX as a strategic tool for estimating establishment potential as part of an environmental risk assessment procedure for biological control agents we discuss biological and ecological aspects necessary to fine-tune its establishment and spread in areas after it has been introduced.     

Mechanisms of prey finding in the aphidophagous ladybird beetle,Harmonia axyridis [Coleoptera: Coccinellidae]

The mechanism of prey finding in adultHarmonia axyridis was investigated by presenting beetles with small gauze or polyethylene bags containing either aphidinfested leaves or healthy leaves alone together with empty control bags. Beetles were attracted to bags by olfactory and visual cues suggesting the presence of aphids, including the odor of aphid-infested leaves, the odor of aphids, the odor of healthy leaves, and the sight of leaves. Particularly effective in prolonging beetles's stay on bags visited was the odor of aphid-infested leaves. It is suggested that adults ofH. axyridis use olfactory and visual cues to detect prey before actual contact occurs. Use of such cues seems more advantageous than random searching, as has been reported in many previous studies of coccinellid foraging.

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Résumé Le mécanisme de découverte de la proie chez l'adulte d'Harmonia axyridis est étudié en donnant aux Coccinelles de petits sacs en gaze ou en polyéthylène contenant soit des feuilles infestées de pucerons, soit des feuilles saines seules, avec des sacs vides comme témoins. Les coccinelles sont attirées vers les sacs par des indicateurs olfactifs ou visuels suggérant la présence de pucerons, incluant l'odeur des feuilles infestées de pucerons, l'odeur des pucerons, l'odeur des feuilles saines et la vue des feuilles. L'odeur des feuilles infestées de pucerons est particulièrement efficace en prolongeant le séjour des Coccinelles sur le sac visité. Il est suggéré que les adultes d'H. axyridis utilisent des indicateurs olfactifs et visuels pour détecter leur proie avant que s'établisse le véritable contact. L'emploi de tels indicateurs semble plus avantageux que la recherche au hasard, comme cela a été indiqué dans de nombreuses études antérieures sur la quête alimentaire des Coccinelles.

     

Intraguild predation involving Harmonia axyridis: a review of current knowledge and future perspectives

As an effective generalist predator of aphids and other hemipteran pests H. axyridis has been a successful biological control agent. However, the very functional traits that have contributed to its success in this regard also implicate it as an intraguild predator that poses a significant risk not only to the diversity of other natural enemies of Hemiptera (and their associated ecosystem services), but to biodiversity more widely. In this paper we will specifically review the existing data on intraguild predation involving H. axyridis, and consider the strength and symmetry of such interactions both within its native guild and within exotic guilds where it has established as an invasive alien. We will use these studies to interpret the observed population declines in predator diversity in the field, predict species at risk in regions not yet invaded and consider implications for resulting ecosystem services. We will also indicate gaps in our knowledge that require further study in order to identify opportunities for mitigation.     

Susceptibility of a native and an exotic lady beetle (Coleoptera: Coccinellidae) to Beauveria bassiana

The exotic multicolored Asian lady beetle, Harmonia axyridis, became established and recently spread across much of North America and southern Canada. In a habitat now used by both the invading H. axyridis and a native lady beetle, Olla v-nigrum, we discovered that the native lady beetle was commonly infected by Beauveria bassiana; whereas, the exotic H. axyridis, was not. Laboratory assays revealed that B. bassiana isolates collected from naturally infected O. v-nigrum were pathogenic to adult O. v-nigrum but not to adult H. axyridis. In contrast, the GHA strain of B. bassiana was not significantly pathogenic to O. v-nigrum nor H. axyridis. Late-season field collections revealed significantly higher B. bassiana infection of O. v-nigrum than H. axyridis. Our results lead us to hypothesize that low susceptibility of H. axyridis to B. bassiana (found to infect O. v-nigrum) may provide an intraguild advantage to H. axyridis over O. v-nigrum; this may also occur with other species of native lady beetles and other endemic entomopathogens in different habitats and regions.     

Effects of augmentative releases of the coccinellid, Adalia bipunctata, and of insecticide treatments in autumn on the spring population of aphids of the genus Dysaphis in apple orchards

The impact of augmentative releases of indigenous predators and insecticide applications to control the autumn aphid forms of the genus Dysaphis (Homoptera: Aphididae), major pest insects on apple trees, was assessed in one-year field experiments in Switzerland. Eggs and larvae of the two-spot ladybird beetle Adalia bipunctata (L.) (Coleoptera: Coccinellidae) were released on 4-year old apple trees in various numbers at five different dates in autumn 1998 when sexuales of the aphids were present. Additionally, Pyrethrum HP was sprayed at the same five dates to compare the effectiveness of these augmentative releases to a commonly applied insecticide. Augmentative releases of larvae before mid-October significantly prevented the deposition of overwintering eggs by aphids of the genus Dysaphis and consequently reduced the number of hatched fundatrices in spring, 1999. There was a significant negative functional response among the number of released coccinellid larvae in autumn 1998 and the number of observed fundatrices on apple trees in spring 1999. Applications of Pyrethrum HP before mid-October were more effective than augmentative releases of larvae of A. bipunctata. The release of coccinellid eggs had no impact on the number of fundatrices of the genus Dysaphis in the next spring because they did not hatch due to bad weather conditions. The weather conditions in autumn seemed to have an impact on the autumn migration of the winged aphids back to their primary host. The prevention of egg deposition of aphids in autumn is a promising control strategy and deserves further exploration for practical use.