Native Ladybird Research Articles

Landscape change and alien invasions drive shifts in native lady beetle communities over a century.

Understanding causes of insect population declines is essential for the development of successful conservation plans, but data limitations restrict assessment across spatial and temporal scales. Museum records represent a source of historical data that can be leveraged to investigate temporal trends in insect communities. Native lady beetle decline has been attributed to competition with established alien species and landscape change, but the relative importance of these drivers is difficult to measure with short-term field-based studies. We assessed distribution patterns for native lady beetles over 12 decades using museum records, and evaluated the relative importance of alien species and landscape change as factors contributing to changes in communities. We compiled occurrence records for 28 lady beetle species collected in Ohio, USA, from 1900 to 2018. Taxonomic beta-diversity was used to evaluate changes in lady beetle community composition over time. To evaluate the relative influence of temporal, spatial, landscape, and community factors on the captures of native species, we constructed negative binomial generalized additive models. We report evidence of declines in captures for several native species. Importantly, the timing, severity, and drivers of these documented declines were species-specific. Land cover change was associated with declines in captures, particularly for Coccinella novemnotata which declined prior to the arrival of alien species. Following the establishment and spread of alien lady beetles, processes of species loss/gain and turnover shifted communities toward the dominance of a few alien species beginning in the 1980s. Because factors associated with declines in captures were highly species-specific, this emphasizes that mechanisms driving population losses cannot be generalized even among closely related native species. These findings also indicate the importance of museum holdings and the analysis of species-level data when studying temporal trends in insect populations.

Niche overlap with an exotic competitor mediates the abundant niche‐centre relationship for a native lady beetle

AbstractAimThe abundant niche‐centre hypothesis has been used to describe patterns of species abundance relative to position in ecological niche space. Such relationships, however, are inconsistently recovered and may be obscured due to non‐equilibrium distributions, such as those caused novel interactions with exotic species. Here, we explore patterns of fitness for the nine‐spotted lady (Coccinella novemnotata) following the introduction of an exotic competitor, the seven‐spotted lady beetle (C. septempunctata). We examine how niche overlap between species may be causing a range reduction for C. novemnotata through a modified abundant niche‐centre relationship.LocationNorth America.MethodsWe applied a time‐oriented view of the abundant niche‐centre relationship for C. novemnotata, both before and after the introduction of C. septempunctata. We tested for both niche overlap and correlation between C. novemnotata body size (i.e. fitness) and the distance from its niche centroid relative to the invasion status of C. septempunctata.ResultsThe current niche space currently occupied by C. novemnotata is equivalent to its historic niche but has shifted to avoid overlap with C. septempunctata. A historic abundant niche‐centre relationship with larger individuals closer to the niche centroid was observed for C. novemnotata. However, this pattern changed to have smaller beetles near the niche centroid when the current distribution was considered.Main ConclusionsFitness patterns displayed by C. novemnotata are effectively described by the abundant niche‐centre hypothesis. However, this relationship was modified with the introduction of C. septempunctata. Coccinella novemnotata is not excluded from previously suitable areas that correspond to niche space nearest to its niche centroid. Abundant niche‐centre relationships may provide strong predictions of species' fitness across a species geographic range; however, these relationships may be obscured by novel pressures resulting in non‐equilibrium states.

Dietary experience alters predatory behavior of two ladybird species on tomato potato psyllid.

The tomato potato psyllid, Bactericera cockerelli, is an invasive pest in Australia, which can cause severe economic loss in the production of Solanaceous crops. As an invasive pest, B. cockerelli may also modify biotic interactions in Australian agricultural and native ecosystems. Resident generalist predators in an area may have the ability to utilize invasive pest species as prey but this will depend on their specific predatory behavior. The extent to which generalist predators learn from their previous dietary experience (i.e., whether they have used a particular species as prey before) and how this impacts subsequent prey choice will influence predator and prey population dynamics after invasion. In this study, one nonnative resident ladybird, Hippodamia variegata, and one native ladybird, Coccinella transversalis, were investigated. Dietary experience with B. cockerelli as a prey species significantly increased preference for the psyllid in a short term (6 h) Petri dish study where a choice of prey was given. Greater suppression of B. cockerelli populations by experienced ladybirds was also observed on glasshouse grown tomato plants. This was presumably due to altered prey recognition by experience. The result of this study suggest the potential to improve the impact of biological control agents on invasive pests by providing early life experience consuming the target species. It may prove valuable for developing improved augmentative release strategies for ladybirds to manage specific insect pest species.

Apparent niche differences between native and exotic lady beetles

Generalist predators can enhance biological control by filling unique niches and complementing other natural enemies, or disrupt it by preying upon other predators and parasitoids, complicating their net value for biological control. We searched for evidence of complementarity and/or interference between native (primarily Eriopis chilensis and E. eschscholtzi) and exotic (primarily Hippodamia variegata and Harmonia axyridis) lady beetles attacking aphids in alfalfa (Medicago sativa) fields near Santiago, Chile. Across 60 fields and three growing seasons, exotic lady beetle adults and larvae reached peak abundance relatively earlier in the growing season than the natives. Furthermore, exotic lady beetle peaks aligned more closely with aphid peak abundance than did native lady beetle peaks. Abundances of native lady beetle adults and larvae were weakly positively correlated with those of exotics, and peak aphid densities were generally lowest in fields where native and non-native lady beetle adults were similarly abundant. Therefore, we did not see evidence of strong interference between exotics and natives. Natural-agricultural habitat edges reduced native, but not exotic, larval lady beetle lag behind aphid peaks and was associated with greater lady beetle and lower aphid densities. Moreover, farms embedded in landscapes with greater natural habitat cover supported more exotic, but not native, lady beetles, and increased the lag between larval lady beetles and aphids for natives but not exotics. These findings support the idea that farms embedded within mosaics of natural and agricultural habitats may provide greater interaction potential between native lady beetles and aphid pests on farms as they move across the landscape, leading to greater top-down control. In contrast, exotic lady beetle lags were unaffected by landscape composition or configuration, but their greater densities in farms surrounded by more natural habitats might reflect a 'concentration effect' whereby exotic lady beetle abundance becomes concentrated in the agricultural habitats that they prefer. Our findings suggest that some lady beetles introduced for biocontrol may be better adapted to intensively managed habitats than natives, possibly leading to spatiotemporal complementarity between native and exotic lady beetles.

Establishment and spread of the invasive ladybird Harmonia axyridis (Coleoptera: Coccinellidae) in Greece: based on contributions from citizen scientists

Harmonia axyridis (Pallas, 1773), also known as the harlequin ladybird, is an invasive non-native species intentionally introduced to many countries as a biological control agent of agricultural pests. In Greece, H. axyridis was first introduced as a biological control agent in 1994, with releases taking place between 1994 and 2000. For many years there was no evidence to indicate that H. axyridis had established self-sustaining populations. In 2008, a citizen science campaign was initiated aimed at raising awareness regarding the invasive status of H. axyridis to farmers and agronomists. The campaign did not yield results, and it was discontinued in 2011. During this study, the distribution, phenology, and presence of H. axyridis in different habitat types and protected areas in Greece are investigated, using both citizen science data and literature records. Records from iΝaturalist, the Alientoma database and social media examined herein demonstrate that H. axyridis has been established in Greece since 2010. Harmonia axyridis is currently present in 13 administrative districts of Greece, most of them at a considerable distance from the initial release sites. The harlequin ladybird is present in urban and agricultural habitats as well as seventeen NATURA 2000 sites. The adverse socioeconomic and environmental impacts of H. axyridis are briefly discussed alongside suggestions for management activities. Based on our findings, we propose the establishment of a national monitoring scheme for H. axyridis and native ladybirds that will also encourage public participation in recording ladybird observations and provide information on the distribution, spread and impact of this invasive non-native species.

Development and reproduction of a native generalist predator, Coccinella transversalis (Coleoptera: Coccinellidae), on the tomato potato psyllid, Bactericera cockerelli, with a greenhouse assay of biocontrol potential

The tomato potato psyllid (Bactericera cockerelli Šulc (Hemiptera: Triozidae)) can cause significant losses to Solanaceous crop via direct feeding and by transmitting the bacterium Candidatus Liberibacter solanacearum. In the present study, an Australian native ladybird beetle, Coccinella transversalis Fabricius, was examined as a potential biological control agent of B. cockerelli. An age-stage, two sex life table of C. transversalis was constructed, demonstrating that C. transversalis could survive, develop, and reproduce successfully when fed B. cockerelli as a monotypic diet. Predation on B. cockerelli by C. transversalis third and fourth instar larvae and adults fit the Holling 'type II' disc equation. In a glasshouse trial using B. cockerelli-infested tomato plants, C. transversalis larvae reduced the population growth of B. cockerelli larvae by 66% and adults by 59% over one month, reducing loss of leaf chlorophyll and increasing plant biomass relative to controls. The results of the present study were compared with a previous study for the resident ladybird beetle species against B. cockerelli. Whereas C. transversalis suppressed B. cockerelli populations, the impact was less than that of the introduced ladybird, Hippodamia variegata Goeze, which showed higher voracity in previous work. However, B. cockerelli is a suitable prey for C. transversalis and this species should aid in its biological control in agricultural ecosystems in Australia.

The Biology and Ecology of Parasitoid Wasps of Predatory Arthropods.

Parasitoid wasps are important components of insect food chains and have played a central role in biological control programs for over a century. Although the vast majority of parasitoids exploit insect herbivores as hosts, others parasitize predatory insects and arthropods, such as ladybird beetles, hoverflies, lacewings, ground beetles, and spiders, or are hyperparasitoids. Much of the research on the biology and ecology of parasitoids of predators has focused on ladybird beetles, whose parasitoids may interfere with the control of insect pests like aphids by reducing ladybird abundance. Alternatively, parasitoids of the invasive ladybird Harmonia axyridis may reduce its harmful impact on native ladybird populations. Different life stages of predatory insects and spiders are susceptible to parasitism to different degrees. Many parasitoids of predators exhibit intricate physiological interrelationships with their hosts, adaptively manipulating host behavior, biology, and ecology in ways that increase parasitoid survival and fitness.

A roadmap for ladybird conservation and recovery.

Ladybirds (Coleoptera: Coccinellidae) provide services that are critical to food production, and they fulfill an ecological role as a food source for predators. The richness, abundance, and distribution of ladybirds, however, are compromised by many anthropogenic threats. Meanwhile, a lack of knowledge of the conservation status of most species and the factors driving their population dynamics hinders the development and implementation of conservation strategies for ladybirds. We conducted a review of the literature on the ecology, diversity, and conservation of ladybirds to identify their key ecological threats. Ladybird populations are most affected by climate factors, landscape composition, and biological invasions. We suggest mitigating actions for ladybird conservation and recovery. Short-term actions include citizen science programs and education, protective measures for habitat recovery and threatened species, prevention of the introduction of non-native species, and the maintenance and restoration of natural areas and landscape heterogeneity. Mid-term actions involve the analysis of data from monitoring programs and insect collections to disentangle the effect of different threats to ladybird populations, understand habitat use by taxa on which there is limited knowledge, and quantify temporal trends of abundance, diversity, and biomass along a management-intensity gradient. Long-term actions include the development of a worldwide monitoring program based on standardized sampling to fill data gaps, increase explanatory power, streamline analyses, and facilitate global collaborations.

Urbanization Negatively Affects the Species Composition of Native Ladybirds in Central Europe

We evaluated the differences in frequency (percentage of sampling sessions where the species was recorded) and abundance (mean number of individuals per 100 sweeps) of native species of ladybirds (Coleoptera: Coccinellidae) in the center of Prague, Central Europe, and in its outskirts. Between 2016 and 2019, coccinellids were sampled on Tilia stands using a standardized sweeping method. Twenty-one species were identified in the total sample of 2,761 adults that were collected in 504 sampling sessions. Species richness was significantly higher in the outskirts than in the center, where seven species were absent and two species were found only once. In contrast, only five species were more abundant in the center than in the outskirts. Communities of native coccinellids were poorer in species, and most species were less abundant in the center than in the outskirts.

<i>HARMONIA AXYRIDIS</i> (PALLAS, 1773) (COLEOPTERA: COCCINELLIDAE) IN MOSCOW CITY AND YALTA CITY: ITS COMPETITION WITH NATIVE SPECIES OF COCCINELLIDS AND THE INFLUENCE OF PARASITOIDS ON ITS NUMBER

The Asian ladybird Harmonia axyridis is an invasive species that is rapidly spreading around the world. Occupying new territories, H. axyridis often causes a decrease in the biodiversity of native species of coccinellids, competing with them for food resources. A comparison of H. axyridis with six species of ladybirds was made in terms of the ratio of the number of individuals and the degree of infestation with parasitoids. In Moscow city, the most widespread species of ladybirds was Adalia bipunctata (67.1%), H. axyridis ranks second (16.5%). The total infestation of H. axyridis pupae with parasitoids is 1.8% in Moscow city and 12.2% in Yalta city. Infestation with parasitoids of other ladybird species in Moscow city is much higher: from 11.0% to 36.4%. Thus, parasitoids effectively regulate the number of native ladybird species, but have an insignificant effect on the number of the invasive species H. axyridis .

Aphid Species and Feeding Location on Canola Influences the Impact of Glucosinolates on a Native Lady Beetle Predator.

Aphids that attack canola (Brassica napus L.) exhibit feeding preferences for different parts of canola plants, which may be associated with brassica-specific glucosinolates. However, this idea remains untested. Furthermore, canola aphid species employ different strategies for tolerating glucosinolates. While the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), excretes glucosinolates, the cabbage aphid Brevicoryne brassicae (L.) (Hemiptera: Aphididae) sequesters them. Given the different detoxification mechanisms, we predicted that both aphid species and aphid feeding location would affect prey suitability for larvae of the predator, Hippodamia convergens (Guérin-Méneville) (Coleoptera: Coccinellidae). We hypothesized that aphids, specifically glucosinolate-sequestering cabbage aphid, reared on reproductive structures that harbor higher glucosinolates concentrations would have greater negative effects on predators than those reared on vegetative structures which have lower levels of glucosinolates, and that the impact of aphid feeding location would vary depending on the prey detoxification mechanism. To test these predictions, we conducted experiments to compare 1) glucosinolates profiles between B. brassicae and M. persicae reared on reproductive and vegetative canola structures, 2) aphid population growth on each structure, and 3) their subsequent impact on fitness traits of H. convergens. Results indicate that the population growth of both aphids was greater on reproductive structures, with B. brassicae having the highest population growth. B. brassicae reared on reproductive structures had the highest concentrations of glucosinolates, and the greatest adverse effects on H. convergens. These findings suggest that both aphid-prey species and feeding location on canola could influence populations of this predator and, thus, its potential for biological control of canola aphids.

Exotic predators are not better biocontrol agents: the harlequin ladybird is not the most voracious in Mexico.

The use of exotic species for pest biocontrol has been a common pathway for introduction and dispersal of invasive species that may have undesired outcomes. Biocontrol agents are believed to be a less damaging alternative than pesticides, but some species may also prey on or parasitize native species or outcompete them for resources. The harlequin ladybird (Harmonia axyridis) is a well-known biocontrol agent originally from Asia that has established invasive populations in 59 countries around the globe. Harlequin ladybirds are generalist predators that in addition to pests prey on an array of different species including other coccinelids’ eggs and larvae. In Mexico, native ladybirds that share ecological requirements with harlequin ladybirds are at risk of being outcompeted and predated upon. The aim of our study was to compare the foraging efficiency of harlequin ladybirds against three species of native coccinelids when preying on aphids. We investigated the foraging behaviour of ladybirds alone and in pairs with a conspecific, a native heterospecific or an exotic heterospecific. We found that the native Cycloneda sanguinea was the species that consumed the most aphids, while Hippodamia convergens was the fastest to find and consume each aphid. Harlequin ladybirds and H. convergens consumed the same number of aphids while P. vittigera consumed less. Conspecific competition was stronger than heterospecific competition. We discuss the suitability of using the exotic invasive harlequin ladybird for aphid biocontrol in comparison with native coccinelids.

Does the non-native Harlequin ladybird disrupt the feeding behaviour of the native two-spot ladybird?

Since its arrival in 2004, the non-native Harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae) has rapidly spread throughout Britain, and it is now the most common coccinellid in England. There have since been concerns about the detrimental effects it may have on native coccinellids because there is a strong correlation between the arrival of H. axyridis and the decline in native species, including the two-spot ladybird, Adalia bipunctata. However, there have been few studies of the behavioural interactions between these two species, which occupy a high-niche overlap. This study investigated if the presence of H. axyridis impacts the feeding behaviour of A. bipunctata through direct competition for aphid prey. Foraging and interactive behaviour of A. bipunctata and H. axyridis were investigated within microcosms. Adalia bipuncata exhibited a similar consumption rate and time in the presence of H. axridis, yet H. axyridis consumed 3.5 times more prey items and were seven times faster compared to A. bipuncata. Observations showed that H. axyridis does not directly disrupt the feeding behaviour of A. bipunctata, but rather indirectly excludes the native species through being a superior competitor for prey items. Results indicate that the decline in native coccinellid species may be a consequence of H. axyridis competitive advantage, but that the concept of coexistence should not be dismissed.

Interactions of ants with native and invasive lady beetles and the role of chemical cues in intraguild interference

The predator-predator naïveté hypothesis suggests that non-native predators benefit from being unknown to native predators, resulting in reduced intraguild interference with native predators. This novelty advantage should depend on the ability of native predators to recognize cues of non-native predators. Here, we compared ant aggression and lady beetle reaction in four native and the invasive lady beetle species Harmonia axyridis. In addition, we tested whether lady beetle cuticular hydrocarbons (CHCs) are involved in species recognition, which might explain naïveté if the invasive species has a specific CHC profile. To this end, we conducted behavioral assays confronting two native ant species with both living lady beetles and lady beetle elytra bearing or lacking CHCs of different lady beetle species. Finally, we characterized CHC profiles of the lady beetles using GC–MS. In general, the aggression of Lasius niger was more frequent than that of Myrmica rubra and L. niger aggression was more frequent towards most native lady beetle species compared to H. axyridis. The removal of CHCs from lady beetle elytra reduced aggression of both ant species. If CHCs of respective lady beetle species were added on cue-free elytra, natural strength of L. niger aggression could be restored. CHC analyses revealed a distinct cue composition for each lady beetle species. Our experiments demonstrate that the presence of chemical cues on the surface of lady beetles contribute to the strength of ant aggression against lady beetles. Reduced aggression of L. niger towards H. axyridis and reduced avoidance behavior in H. axyridis compared to the equally voracious C. septempunctata might improve the invasive lady beetle’s access to ant-tended aphids.

Community science data suggests that urbanization and forest habitat loss threaten aphidophagous native lady beetles.

Community scientists have illustrated rapid declines of several aphidophagous lady beetle (Coccinellidae) species. These declines coincide with the establishment of alien coccinellids. We established the Buckeye Lady Beetle Blitz program to measure the seasonal occupancy of coccinellids within gardens across a wide range of landscape contexts. Following the Habitat Compression Hypothesis, we predicted that gardens within agricultural landscapes would be alien‐dominated, whereas captures of natives would be higher within landscapes encompassing a high concentration of natural habitat.Within the state of Ohio, USA, community scientists collected lady beetles for a 7‐day period across 4 years in June and August using yellow sticky card traps. All identifications were verified by professional scientists and beetles were classified by three traits: status (alien or native), mean body length, and primary diet. We compared the relative abundance and diversity of coccinellids seasonally and determined if the distribution of beetles by size, status, and diet was related to landscape features.Alien species dominated the aphidophagous fauna. Native aphidophagous coccinellid abundance was positively correlated with forest habitat while alien species were more common when gardens were embedded within agricultural landscapes. Urbanization was negatively associated with both aphidophagous alien and native coccinellids. Synthesis and Applications: Our census of native coccinellid species within residential gardens—a widespread and understudied habitat—was enabled by volunteers. These data will serve as an important baseline to track future changes within coccinellid communities within this region. We found that native coccinellid species richness and native aphidophagous coccinellid abundance in gardens were positively associated with forest habitat at a landscape scale of 2 km. However, our understanding of when and why (overwintering, summer foraging, or both) forest habitats are important remains unclear. Our findings highlight the need to understand how declining aphidophagous native species utilize forest habitats as a conservation priority.

Long Term Monitoring in Switzerland Reveals That Adalia bipunctata Strongly Declines in Response to Harmonia axyridis Invasion.

Simple SummaryThe harlequin ladybird, Harmonia axyridis, is an Asian species that has invaded Europe and other continents, where it is suspected to cause the decline of native ladybirds through competition and predation. In north-western Switzerland, ladybirds were monitored for 11 years in four habitats (broadleaved hedges, meadows, pine and spruce stands) to assess the decline of native ladybirds following the invasion of the harlequin ladybird. These surveys showed that, on broadleaved hedges, the harlequin ladybird quickly became the most abundant species, representing 60–80% of all specimens collected in this habitat. One species, the two-spot ladybird, Adalia bipunctata, almost disappeared during this period, whereas it was the most abundant ladybird on broadleaved trees and shrubs when this study started. The other native species did not show any clear sign of decline. The harlequin ladybird was the second most abundant species in pine stands, and was not abundant in meadows and in spruce stands. The total number of ladybirds feeding on aphids did not decline during this period, suggesting that the arrival of the harlequin ladybird did not affect the predation pressure on aphids. Nevertheless, the severe decline of the two-spot ladybird deserves further investigations.A long-term monitoring was conducted at 40 sites in four different habitats in north-western Switzerland to observe changes in populations of native ladybirds, following the invasion of the Asian harlequin ladybird, Harmonia axyridis. From 2006 to 2017, the same trees and meadows were sampled at least seven times per year using standard protocols. On 15 broadleaved hedges, H. axyridis quickly became the dominant species, representing 60 to 80% of adult ladybirds collected. It was second in abundance at five pine (Pinus sylvestris) stands and was a minor component of the ladybird complex at five spruce (Picea abies) stands and in 15 meadows. This survey revealed the severe decline of Adalia bipunctata, which was the most abundant native ladybird on broadleaved trees in 2006–2009 and has almost disappeared since 2010. So far, other native ladybirds do not seem to decline significantly, including species occupying the same ecological niches as H. axyridis. The total number of aphidophagous ladybirds did not decline either, suggesting that the biological control function of ladybirds on aphids living in these habitats has not been affected by the arrival of H. axyridis. Recommendations are given to further assess the impact of H. axyridis on native ladybirds and aphids.

Mortality of native and invasive ladybirds co-infected by ectoparasitic and entomopathogenic fungi.

Harmonia axyridis is an invasive alien ladybird in North America and Europe. Studies show that multiple natural enemies are using Ha. axyridis as a new host. However, thus far, no research has been undertaken to study the effects of simultaneous infection by multiple natural enemies on Ha. axyridis. We hypothesized that high thallus densities of the ectoparasitic fungus Hesperomyces virescens on a ladybird weaken the host’s defenses, thereby making it more susceptible to infection by other natural enemies. We examined mortality of the North American-native Olla v-nigrum and Ha. axyridis co-infected with He. virescens and an entomopathogenic fungus—either Beauveria bassiana or Metarhizium brunneum. Laboratory assays revealed that He. virescens-infected O. v-nigrum individuals are more susceptible to entomopathogenic fungi, but Ha. axyridis does not suffer the same effects. This is in line with the enemy release hypothesis, which predicts that invasive alien species in new geographic areas experience reduced regulatory effects from natural enemies compared to native species. Considering our results, we can ask how He. virescens affects survival when confronted by other pathogens that previously had little impact on Ha. axyridis.

Comparison of native and non-native predator consumption rates and prey avoidance behavior in North America and Europe.

Novel predator–prey interactions can contribute to the invasion success of non‐native predators. For example, native prey can fail to recognize and avoid non‐native predators due to a lack of co‐evolutionary history and cue dissimilarity with native predators. This might result in a competitive advantage for non‐native predators. Numerous lady beetle species were globally redistributed as biological control agents against aphids, resulting in novel predator–prey interactions. Here, we investigated the strength of avoidance behavior of the pea aphid (Acyrthosiphon pisum) toward chemical cues of native lady beetles and non‐native Asian Harmonia axyridis and European Coccinella septempunctata and Hippodamia variegata in North America, hypothesizing that cues of non‐native lady beetles induce weaker avoidance behavior than cues of co‐evolved native lady beetles. Additionally, we compared aphid consumption of lady beetles, examining potential predation advantages of non‐native lady beetles. Finally, we compared cue avoidance behavior between North American and European pea aphid populations and aphid consumption of native and non‐native lady beetles in North America and Europe. In North America, pea aphids avoided chemical cues of all ladybeetle species tested, regardless of their origin. In contrast to pea aphids in North America, European pea aphids did not avoid cues of the non‐native H. axyridis. The non‐native H. axyridis and C. septempunctata were among the largest and most voracious lady beetle species tested, on both continents. Consequently, in North America non‐native lady beetle species might have a competitive advantage on shared food resources due to their relatively large body size, compared to several native American lady beetle species. In Europe, however, non‐native H. axyridis might benefit from missing aphid cue avoidance as well as a large body size. The co‐evolutionary time gap between the European and North American invasion of H. axyridis likely explains the intercontinental differences in cue avoidance behavior and might indicate evolution in aphids toward non‐native predators.

Predation and avoidance behaviour in aphid‐ladybird interactions of native and invasive ladybirds in Europe

While detrimental effects of invasive predators on native species are well documented, we often lack a mechanistic understanding of the invasion success. Lack of prey avoidance behaviour can lead to higher consumption rates by invasive predators compared to native predators. This competitive advantage is expected to contribute to the invasion success of non‐native predators. We compared aphid consumption and cue avoidance behaviour of aphids between four native ladybird species (Coccinella septempunctata, Adalia bipunctata, Propylea quatuordecimpunctata, and Hippodamia variegata) and the invasive Asian ladybird Harmonia axyridis. The invasive H. axyridis and the native C. septempunctata consumed more aphids than the three smaller native ladybird species. In line with our expectations, aphids avoided leaves bearing cues of most native ladybird species but not of the invasive H. axyridis. Our results indicate that body size rather than ladybird origin determined aphid predation rates. The lack of aphid avoidance behaviour towards cues of H. axyridis indicates that they were not able to recognise the chemical cues of the invasive predator. Relatively large body size and the absence of cue avoidance in aphids might benefit the invasive H. axyridis, particularly in comparison to smaller native ladybird species. The absence of avoidance behaviour in aphids might lead to even higher predation rates of H. axyridis under more natural conditions.

Interactions between exotic and native lady beetle species stabilize community abundance.

A 23-year time-series of abundance for 13 lady beetle species (Coccinellidae) was used to investigate community stability. The community exhibited persistence in ten habitats, no overall trend in abundance, and low temporal variability quantified as Population variability (PV) = 0.33 on a scale from 0 to 1 that declined to 0.16 in the past 8years. This high level of stability occurred as exotic lady beetles disrupted populations of the native species. For hypothetical communities of pairs of species (with randomly generated annual abundances in the range for lady beetles), PV increased linearly with the correlation coefficients between individual time series, illustrating a "portfolio effect". PV for the real community and the negative correlation between the abundance of exotics and natives fit this relationship precisely. A gradual decline of natives matched by an equal gradual rise in the abundance of exotics contributed to the negative correlation that stabilized the community. The abundance of the dominant species, an exotic, was negatively correlated with other exotics and most natives, and its stability increased over time, helping to stabilize the community. The community was most stable in habitats where beetle abundance was high (crops, particularly perennial crops) and, unexpectedly, was least stable in habitats with high diversity and stability of vegetation cover (forests). These data are consistent with the hypothesis that competition between exotic and native species, with release from competition for natives in some years, stabilized the abundance of this community. Stability may not last if populations of native species continue declining.