Living with the enemy: parasites and pathogens of the ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis is an invasive alien predator in many countries across the world. The rapid establishment and spread of this species is of concern because of the threat it poses to biodiversity as a generalist predator. Understanding the mechanisms that contribute to the success of this species as an invader is not only intriguing but also critical to our understanding of the processes governing such invasions. The enemy release hypothesis (ERH) could explain the rapid population growth of many invasive alien species. However, empirical evidence in support of the ERH is lacking. An alternative hypothesis that could explain rapid population growth is evolution of increased competitive ability (EICA). Here we provide an overview of the parasites and pathogens of coccinellids with a particular focus on H. axyridis as a host. We examine the differential susceptibility of host species and highlight the resilience of H. axyridis in comparison to other coccinellids. We recognise the paucity and limitations of available information and suggest that studies, within a life-table framework, comparing life history traits of H. axyridis in both the native and introduced ranges are necessary. We predict that H. axyridis could benefit from both enemy release and EICA within the introduced range but require further empirical evidence.     

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

1. 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.
2. 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.
3. 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.
4. 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.
5. 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.

     

Investigating the genetic load of an emblematic invasive species: the case of the invasive harlequin ladybird Harmonia axyridis

Introduction events can lead to admixture between genetically differentiated populations and bottlenecks in population size. These processes can alter the adaptive potential of invasive species by shaping genetic variation, but more importantly, they can also directly affect mean population fitness either increasing it or decreasing it. Which outcome is observed depends on the structure of the genetic load of the species. The ladybird Harmonia axyridis is a good example of invasive species where introduced populations have gone through admixture and bottleneck events. We used laboratory experiments to manipulate the relatedness among H. axyridis parental individuals to assess the possibility for heterosis or outbreeding depression in F₁ generation offspring for two traits related to fitness (lifetime performance and generation time). We found that inter‐populations crosses had no major impact on the lifetime performance of the offspring produced by individuals from either native or invasive populations. Significant outbreeding depression was observed only for crosses between native populations for generation time. The absence of observed heterosis is indicative of a low occurrence of fixed deleterious mutations within both the native and invasive populations of H. axyridis. The observed deterioration of fitness in native inter‐population crosses most likely results from genetic incompatibilities between native genomic backgrounds. We discuss the implications of these results for the structure of genetic load in H. axyridis in the light of the available information regarding the introduction history of this species.     

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.     

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.     

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.     

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.     

Do invasive alien plants benefit more from global environmental change than native plants?

Invasive alien plant species threaten native biodiversity, disrupt ecosystem functions and can cause large economic damage. Plant invasions have been predicted to further increase under ongoing global environmental change. Numerous case studies have compared the performance of invasive and native plant species in response to global environmental change components (i.e. changes in mean levels of precipitation, temperature, atmospheric CO₂ concentration or nitrogen deposition). Individually, these studies usually involve low numbers of species and therefore the results cannot be generalized. Therefore, we performed a phylogenetically controlled meta‐analysis to assess whether there is a general pattern of differences in invasive and native plant performance under each component of global environmental change. We compiled a database of studies that reported performance measures for 74 invasive alien plant species and 117 native plant species in response to one of the above‐mentioned global environmental change components. We found that elevated temperature and CO₂ enrichment increased the performance of invasive alien plants more strongly than was the case for native plants. Invasive alien plants tended to also have a slightly stronger positive response to increased N deposition and increased precipitation than native plants, but these differences were not significant (N deposition: P = 0.051; increased precipitation: P = 0.679). Invasive alien plants tended to have a slightly stronger negative response to decreased precipitation than native plants, although this difference was also not significant (P = 0.060). So while drought could potentially reduce plant invasion, increases in the four other components of global environmental change considered, particularly global warming and atmospheric CO₂ enrichment, may further increase the spread of invasive plants in the future.     

Can Developmental and Behavioral Characteristics of the Multicolored Asian Lady Beetle Harmonia Axyridis Explain its Invasive Success?

Major impacts of biological invasions are widely recognized and underscore the need to understand the relation between life-history traits of invasive species and the invasion process. Growth of juveniles and adult survival of invasive species are key factors in invasion process. Life-history traits that increase juvenile fitness including increased rates of development and behavioral characteristics that facilitate competitive success such as increased predator efficiency and foraging ability may explain invasiveness of a species. Invasion of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) in North America provides an opportunity to investigate life-history traits of juveniles of an invasive species. Here, we evaluate both developmental and behavioral traits that may explain the success of H. axyridis by comparing it to an ecologically similar indigenous species Coleomegilla maculata lengi Timberlake (Coleoptera: Coccinellidae). Three points may contribute to the invasiveness of H. axyridis. First, development of H. axyridis was faster during the 2nd larval instar than C. maculata, a characteristic that may reduce vulnerability at young instars. Second, H. axyridis reached the 4th instar more rapidly than C. maculata. The 4th instar of H. axyridis was also characterized by higher predation efficiency with increased voracity, lethal contact and search efficiency of pea aphids Acyrthosiphon pisum. Finally, surprisingly, a 5th larval instar occured in 33% of the individuals of H. axyridis and was characterized by the same developmental time, but with increased voracity and weight gain compared to 4th larval instars, suggesting an increased fitness of these individuals. These developmental characteristics coupled with increased predation efficiency and behavioral characteristics enhanced the juvenile growth and predatory abilities of this species and may contribute to the invasive ability of H. axyridis.     

Potential spatial interaction of the invasive species Harmonia axyridis (Pallas) with native and endemic coccinellids

Biological invasions represent a serious menace to local species assemblages, mainly due to interspecific relationships such as competition and predation. One important invasive species worldwide is Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), which has invaded many regions of the world, threatening the native and endemic coccinellid assemblages due to negative interspecific interactions. These interactions have been widely studied at a local scale, but have been less studied at regional scales. Our aim was to estimate and analyse the potential spatial interaction associated with the co‐occurrence of H. axyridis with native and endemic species in Chile, considering bioclimatic and land cover variables. First, we created species distribution models (SDM) for H. axyridis, native and endemic coccinellids and six representative coccinellid species using maximum entropy technique. Then, we overlapped each SDM with land cover types to estimate the bioclimatic suitability within each land cover type. Finally, we identified the co‐occurrences of organisms according to the SDM and the land cover types, estimating in what land covers H. axyridis and the other coccinellids are more likely to co‐occur. Our results show that the suitable area for H. axyridis occurs from 30° to 42°S in Chile, while for native and endemic species this area is greater. The six selected species are mainly concentrated in central Chile, but differ in their potential suitable areas; Adalia angulifera Mulsant and Scymnus bicolor (Germain) have the largest range, and Mimoscymnus macula (Germain) has the most restricted one. The highest level of potential spatial interactions with H. axyridis occurs in central Chile, specifically in croplands and scrublands, and the lowest in primary native forest for all the species. Our results provide a spatially explicit baseline for coccinellid conservation and management of this invasive species.     

Alien plant species favoured over congeneric natives under experimental climate warming in temperate Belgian climate

Climate warming and biological invasions by alien species are two key factors threatening the world’s biodiversity. To date, their impact has largely been studied independently, and knowledge on whether climate warming will promote invasions relies strongly on bioclimatic models. We therefore set up a study to experimentally compare responses to warming in native and alien plant species. Ten congeneric species pairs were exposed to ambient and elevated temperature (+3°C) in sunlit, climate-controlled chambers, under optimal water and nutrient supply to avoid interaction with other factors. All species pairs combined, total plant biomass reacted differently to warming in alien versus native species, which could be traced to significantly different root responses. On average, native species became less productive in the warmer climate, whereas their alien counterparts showed no response. The three alien species with the strongest warming response (Lathyrus latifolius, Cerastium tomentosum and Artemisia verlotiorum) are currently non-invasive but all originate from regions with a warmer climate. Still, other alien species that also originate from warmer regions became less or remained equally productive. Structural or ecophysiological acclimation to warming was largely absent, both in native and alien species, apart from light-saturated photosynthetic rate, where warming tended to restrain the native but not the alien species. A difference in the capacity to acclimate photosynthetic rates to the new climate may therefore have caused the contrasting biomass response. Future experiments are needed to ascertain whether climate warming can effectively tip the balance between native and alien competitors.     

Life history of the harlequin ladybird, <Emphasis Type="Italic">Harmonia axyridis</Emphasis>: a global meta-analysis

The harlequin ladybird, Harmonia axyridis, is an important natural enemy of aphids throughout the world, but is now also considered an invasive alien species. We performed a meta-analysis of published life history data to address the question whether invading populations in Europe and North America have life history parameters that differ from native populations in Asia, explaining the beetle’s invasion success in new territories. In this meta-analysis, we accounted for important covariables that are often reported in published studies such as temperature, food source (aphids or eggs of Ephestia kuehniella), strain (laboratory or field populations) and photoperiod. Temperature was a key factor having consistent large effects on development rate, survival and reproductive characteristics of H. axyridis. Food source, strain, and photoperiod had effects on some, but not all characteristics, and their overall effect across characteristics was minor. Individuals of invasive populations had a shorter pre-oviposition period and higher fecundity at low temperatures than those of native populations, and a greater longevity across all temperatures. No differences in survival were found between native and invasive populations, while differences in development rate were not consistent, with opposing results obtained according to the way development rate was measured in trials reported in the literature. Results of this meta-analysis support the hypothesis that the life history of the beetle has changed during its invasion into North America and Europe. Invasive populations had a shorter pre-oviposition period and higher fecundity at low temperatures, as well as a greater longevity across all temperatures than native populations. These differences may partially explain the invasive success of H. axyridis.     

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.     

Functional traits,growth patterns,and litter dynamics of invasive alien and co-occurring native shrub species of chir pine forest in the central Himalaya,India

Across the continents, plant invasion is identified as one of the main threats to ecosystem functioning and stability. The main objective of this research was to evaluate the differences in the functional traits between invasive alien (Ageratina adenophora (Spreng.) and Lantana camara L.) and native (Berberis asiatica Roxb. Ex DC., Pyracantha crenulata (D. Don.) M. Roemer and Rubus ellipticus Sm.) shrub species of chir pine (Pinus roxburghii Sarg.) forest in the central Himalaya. Three 0.5 hectare chir pine forest stands were selected and individuals of similar diameter were tagged for comparative studies of leaf traits, growth pattern, and biomass accumulation in structural organs of each invasive alien and native species. Our one-way ANOVA and Tukey’s post hoc test results showed that both the invasive alien species have significantly (p?<?0.05) higher SLA, LWC, total chlorophyll content, foliar nutrient (N and P), RGR, LMR, SMR, nutrient uptake, and nutrient use efficiencies than native species. Leaf litter decomposition rate and nutrient release were also significantly (p?<?0.05) higher in both the invasive alien species. Native species, R. ellipticus, shared some of the traits, such as leaf area, chlorophyll content, RGR, LAR, LMR, and nutrient uptake efficiency with invasive alien species. The majority of traits differed among invasive alien and native species, implying that the success of invasive alien species is best described by being functionally distinct from native species. These findings indicate that invasive alien species had advanced functional traits which may be playing an important role in a rapid spread in the central Himalaya.

     

Ecological correlates of local extinction and colonisation in the British ladybird beetles (Coleoptera: Coccinellidae)

Five main drivers of population declines have been identified: climate change, habitat degradation, invasive alien species (IAS), overexploitation and pollution. Each of these drivers interacts with the others, and also with the intrinsic traits of individual species, to determine species’ distribution and range dynamics. We explored the relative importance of life-history and resource-use traits, climate, habitat, and the IAS Harmonia axyridis in driving local extinction and colonisation dynamics across 25 ladybird species (Coleoptera: Coccinellidae).Species were classified as continually present, continually absent, extinct, or colonising in each of 4,642 1-km² grid squares. The spatial distribution of local extinction and colonisation events (in the grid squares) across all species’ ranges were related to ecological traits, overlap with H. axyridis, climate, and habitat factors within generalised linear models (GLMs). GLMs were also used to relate species’ traits, range characteristics, and niche overlap with H. axyridis to extinction and colonisation rates summarised at the species level. Bayesian model averaging was used to account for model uncertainty, and produce reduced sets of models which were well-supported by data. Species with a high degree of niche overlap with H. axyridis suffered higher extinction rates in both analyses, while at the spatial scale extinctions were more likely and colonisations less likely in areas with a high proportion of urban land cover. In the spatial analysis, polymorphic species with large range sizes were more likely to colonise and less likely to go extinct, and sunny grid squares were more likely to be colonised. Large, multivoltine species and rainy grid squares were less likely to colonise or be colonised. In conclusion for ladybirds, extinction and colonisation dynamics are influenced by several factors. The only factor that both increased the local extinction likelihood and reduced colonisation likelihood was urban land cover, while ecological overlap with H. axyridis greatly increased extinction rates. Continued spread of H. axyridis is likely to adversely affect native species and urban areas may be particularly vulnerable.     

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.     

Harmonia axyridis invasions: Deducing evolutionary causes and consequences

I consider evolutionary approaches to deducing factors that have made the ladybird beetle Harmonia axyridis such a successful invader, and the contribution that studies of this species in its native range can make. Work aiming to demonstrate which (pre)adaptations have made the species so successful often fails to compare these putative characters with those of other ladybirds. This has led to a tendency for “argument by design”‐type claims on characters widely shared by non‐invasive coccinellids. There is good evidence from genetic studies that evolutionary change occurred in invasive populations, contributing to their success. There is some evidence for subsequent evolutionary change after the establishment of invasive H. axyridis, primarily in the native organisms with which the ladybird interacts. I show here that there appears to have been little adaptation in H. axyridis, over about 20 generations, to the alkaloids of one North American native intraguild prey, the ladybird Coleomegilla maculata. Studies of H. axyridis in its native range are important, as they provide a snapshot of the ancestral ladybird, unobscured by subsequent evolutionary change related to its invasiveness. They provide baseline data about phenomena such as interactions with natural enemies and intraguild predation, and they also can provide pointers as to how H. axyridis might further adapt in the regions it has colonized. Harmonia axyridis represents an ideal opportunity for greater international co‐operation between scientists studying this species in its native range in Asia and scientists studying it in Europe, America and Africa, where it is an invasive exotic.     

Effects of experimental climate warming and associated soil drought on the competition between three highly invasive West European alien plant species and native counterparts

It is widely suggested that climate warming will increase the impact of biological invasions, yet, to date studies on the combined effect of these two global changes are scarce. Here, we study how climate warming and associated soil drought affect the competition between native and invasive alien plant species. Three highly invasive alien plant species in West Europe, each with a native competitor, were grown either together or in isolation at ambient and at elevated air temperature (+3 °C) in climate-controlled chambers. Equal amounts of water were added to all communities. Soil drought observed in the heated chambers did not induce severe stress in the plants. In two species pairs, Fallopia japonica (Houtt.)–Cirsium arvense (L.) Scop. and Solidago gigantea Ait.–Epilobium hirsutum L. (alien invasive–native), the native species dominated in mixture, while the alien invasive species dominated in the third pair Senecio inaequidens DC.–Plantago lanceolata L. Warming did not modify the competitive balance in any of these pairs, in spite of enhancing the aboveground biomass of S. inaequidens and P. lanceolata and the greater photosynthetic rates in S. inaequidens. The results of this study cannot be extrapolated to all invasive or exotic species but may represent the possible future of three principle invaders and some of their key native counterparts. Future experiments are needed to identify response patterns of alien plants to climate warming more in general.     

Non‐native fishes and climate change: predicting species responses to warming temperatures in a temperate region

1. Temperate regions with fish communities dominated by cold‐water species (physiological optima <20 °C) are vulnerable to the effects of warming temperatures caused by climate change, including displacement by non‐native cool‐water (physiological optima 20–28 °C) and warm‐water fishes (physiological optima >28 °C) that are able to establish and invade as the thermal constraints on the expression of their life history traits diminish. 2. England and Wales is a temperate region into which at least 38 freshwater fishes have been introduced, although 14 of these are no longer present. Of the remaining 24 species, some have persisted but failed to establish, some have established populations without becoming invasive and some have become invasive. The aim of the study was to predict the responses of these 24 non‐native fishes to the warming temperatures of England and Wales predicted under climate change in 2050. 3. The predictive use of climate‐matching models and an air and water temperature regression model suggested that there are six non‐native fishes currently persistent but not established in England and Wales whose establishment and subsequent invasion would benefit substantially from the predicted warming temperatures. These included the common carp Cyprinus carpio and European catfish Silurus glanis, fishes that also exert a relatively high propagule pressure through stocking to support angling and whose spatial distribution is currently increasing significantly, including in open systems. 4. The potential ecological impacts of the combined effects of warming temperatures, current spatial distribution and propagule pressure on the establishment and invasion of C. carpio and S. glanis were assessed. The ecological consequences of C. carpio invasion were assessed as potentially severe in England and Wales, with impacts likely to relate to habitat destruction, macrophyte loss and increased water turbidity. However, evidence of ecological impacts of S. glanis elsewhere in their introduced range was less clear and so their potential impacts in England and Wales remain uncertain.     

Evolutionary ecology of microsporidia associated with the invasive ladybird Harmonia axyridis

Invasive species are characterized by the rapid growth and spread of their populations after establishing a foothold in new habitats, and there are now many examples of such species negatively affecting biodiversity and the economy. It is unclear why some species can become successful invaders, whereas most (even if closely related) remain noninvasive. We previously proposed a hypothesis that parasites associated with invading species can promote their invasive success if they are harmless toward the invaders but harmful to their competitors and/or predators in the newly colonized habitat. Here we discuss whether microsporidia that have recently been discovered in the invasive ladybird Harmonia axyridis contribute to its invasive success. We show that all H. axyridis beetles sourced from diverse collection sites all over the world carry abundant microsporidia. This suggests that both native and invasive H. axyridis populations are associated with these tolerated parasites, which were likely to have existed in native populations before expansion rather than being acquired in newly colonized areas. We describe the pathogenesis of the microsporidia during different developmental stages of H. axyridis and we address the possibility that the predation of its infected eggs and larvae by competing native ladybird species may lead to their infection and ultimately to their decline. Finally, we discuss our initial hypothesis: microsporidia that are tolerated by an invasive vector insect can be active against susceptible native competitors and/or predator species.