Spotlight on the positive effects of the ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis> on agriculture

In the midst of considerable negativity surrounding the ladybird Harmonia axyridis (Pallas), this paper sheds some light on the positive effects that this predator has had on agriculture. Using resources available at the USDA, National Agricultural Library (DigiTop literature database, Navigator platform), I searched the abstracts of published literature on H. axyridis using the search term “Harmonia axyridis” or “Leis axyridis.” After deleting duplicate records, there were 1193 total records on this ladybird in the database dating from 1961–2015. Of this total, 90 records (7.54 %) highlighted positive effects of H. axyridis on agriculture in terms of its spectacular voracity, predation capacity, and effectiveness in suppressing plant pests, primarily aphids and scales, in forests, orchards, crop fields, gardens, greenhouses, and in laboratory bioassays. More concerted research is necessary to develop methodologies to exploit the positive behaviors of H. axyridis, while managing its negative ones.     

Reaping benefits from an invasive species: role of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in natural biological control of <Emphasis Type="Italic">Aphis glycines</Emphasis> in North America

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is an invasive species present in numerous agroecosystems in North America. Despite adverse impacts as a threat to native biodiversity, a nuisance household invader and a pest in fruit production, H. axyridis also plays a beneficial role as a major component of assemblages of generalist predators in several agricultural crops. Here, we review the role of H. axyridis as a natural enemy of Aphis glycines Matsumura (Hemiptera: Aphididae), an invasive pest of soybean, Glycine max (L.) Merrill (Fabales: Fabaceae), in North America. Harmonia axyridis is often the most abundant predator species attacking A. glycines in soybean agroecosystems. This predator has the potential to both prevent and suppress A. glycines outbreaks. Further studies are needed to fully understand and utilize the potential of H. axyridis as a natural enemy in the management of A. glycines and other agricultural pests in agroecosystems worldwide.     

Heritable bacterial endosymbionts in native and invasive populations of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis Pallas (1773) (Coleoptera: Coccinellidae) is the well-studied system of invasive insect species. Native and invasive parts of the area of H. axyridis are isolated geographically. We studied the species composition and the distribution of bacterial symbionts Spiroplasma and Rickettsia in seven localities of the native area and six localities of the invasive area of H. axyridis. Rickettsia was detected in H. axyridis populations for the first time. We found that the proportion of beetles infected with Rickettsia in native and invasive populations of H. axyridis is about 0.03. Spiroplasma was found only in native populations of H. axyridis. The proportion of infected individuals with Spiroplasma in native populations of H. axyridis is about 0.08. All studied native populations of H. axyridis are infected with Spiroplasma, while all invasive populations are not. We discuss the possible influence of Spiroplasma and Rickettsia in the formation of invasive populations of H. axyridis.     

Chemical defences of native European coccinellid eggs against intraguild predation by the invasive Asian coccinellid, <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas)

Harmonia axyridis (Pallas) is a coccinellid of Asian origin that has recently invaded substantial parts of Europe and is suspected to affect native coccinellid populations through intraguild predation and competition for food. Previous work has shown that two species from the Calvia genus appeared to be well protected against H. axyridis predation. To deepen our understanding on chemical protection of Calvia spp. and the predation risk by H. axyridis, we tested for susceptibility and palatability of Calvia spp. and H. axyridis eggs against predation by H. axyridis neonate larvae. Results show that eggs of C. quatuordecimguttata were mostly not eaten by H. axyridis, while eggs of the congeneric C. decemguttata were found to be largely unprotected against predation by the invasive coccinellid. We also observed that H. axyridis first instars successfully cannibalized on conspecific eggs. Removing the surface chemicals from C. quatuordecimguttata eggs resulted in significantly reduced protection from being preyed upon by H. axyridis, while applying these extracts onto C. decemguttata and H. axyridis eggs resulted in increased protection against H. axyridis larvae. The importance of surface chemicals in the interactions between H. axyridis and native coccinellids was confirmed by GC–MS analysis, showing a high diversity of hydrocarbons located on the surface of C. quatuordecimguttata eggs, i.e. more than twice as many when compared to C. decemguttata. Survival of H. axyridis larvae feeding on eggs of C. quatuordecimguttata, C. decemguttata or conspecific eggs, from which surface chemicals were removed by washing them with hexane, was not different from survival on unwashed eggs.     

Long term changes in the communities of native ladybirds in Northern Italy: impact of the invasive species <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas)

Since the mid-2000s, the exotic coccinellid species Harmonia axyridis (Pallas) has established itself in Northern Italy, raising serious concerns about possible repercussions on native ladybirds. In this study we compared the ladybird assemblages in 2015–2016 with those sampled in 1995–1996, before the arrival of H. axyridis. Surveys were carried out in the same sites and with the same methods for both periods. Aphidophagous ladybirds were sampled in hedgerows and herbaceous habitats at field margins by mechanical knockdown and sweeping net, respectively. The changes in the structure of ladybird communities after the arrival of H. axyridis were significantly different between arboreal and herbaceous habitats. Harmonia axyridis is currently the dominant species in shrubs and trees, and all the native ladybird species taken together account for only approximately one third of the total individuals sampled in 2015–2016. On the other hand, the relative abundance of the exotic species in herbaceous habitats was low, and it has had a negligible relevance on the structure of ladybird communities on grasslands. Among native species, Adalia bipunctata (L.), whose ecological niche largely overlaps with that of H. axyridis, suffered the largest decline between 1995–1996 and 2015–2016. The co-occurrence of H. axyridis invasion and the regression of A. bipunctata suggest a direct impact of the exotic species, because no other major modifications occurred in the studied areas between sampling periods.     

Determining the effects of life stage,shared prey density and host plant on intraguild predation of a native lacewing (<Emphasis Type="Italic">Chrysoperla carnea</Emphasis>) by an invasive coccinellid (<Emphasis Type="Italic">Harmonia axyridis</Emphasis>)

Negative impacts of non-native Harmonia axyridis (Pallas) on members of the native aphid enemy guild have been widely hypothesised but mainly only assessed with other coccinellid species, and mostly in small experimental arenas. Here we investigated the interactions between H. axyridis and Chrysoperla carnea Stephens larvae. In small-scale (Petri dish) arenas 2nd-instar C. carnea were at risk of predation from larval (2nd and 4th-instar) and adult (male and female) H. axyridis while 3rd-instar C. carnea were only at minimal risk from 4th-instar and adult female H. axyridis. Plant species, aphid species and aphid density did not affect intraguild predation of 2nd-instar C. carnea by 4th-instar and adult H. axyridis in mesocosm experiments. Chrysoperla carnea consumed similar numbers of Megoura viciae Buckton, Aphis fabae Scop. and Acyrthosiphon pisum Harris aphids while H. axyridis consumed fewer M. viciae than the other two species. The greatest suppression of A. pisum was achieved in treatments with both C. carnea and H. axyridis. Life stage and the sex of H. axyridis as well as the life stage of C. carnea are important variables affecting intraguild predation and these attributes should be considered when assessing the potential threat of other potentially invasive alien predators.     

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undaria pinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.     

Predators and parasitoids of the harlequin ladybird, <Emphasis Type="Italic">Harmonia axyridis</Emphasis>, in its native range and invaded areas

The harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae) has rapidly spread in several continents over the past 30 years and is considered an invasive alien species. The success of H. axyridis as an invader is often attributed to weak control by natural enemies. In this paper, we provide an overview of current knowledge on predators and parasitoids of H. axyridis. The common feature of predators and parasitoids is that they directly kill exploited organisms. Currently available data show that H. axyridis, displaying a variety of chemical, mechanical, and microbiological anti-predator defenses, is usually avoided by predators. However, some birds and invertebrates can eat this ladybird without harmful consequences. The primary defenses of H. axyridis against parasitoids include immune response and physiological and nutritional unsuitability for parasitoid development. These defenses are probably relatively efficient against most ladybird parasitoids, but not against flies of the genus Phalacrotophora. The latter are idiobiont parasitoids and hence can evade the host’s immune response. Indeed, rates of parasitism of H. axyridis by Phalacrotophora in the Palaearctic region (both in the native range in Asia and in Europe) are relatively high. While strong evidence for enemy release on the invasive populations of H. axyridis is lacking, several cases of parasitoid acquisition have been recorded in Europe, North America, and South America. We conclude that enemy release cannot be excluded as a possible mechanism contributing to the spread and increase of H. axyridis in the early stages of invasion, but adaptation of parasitoids may lead to novel associations which might offset previous effects of enemy release. However, further work is required to elucidate the population-level effects of such interactions.     

The contribution of a beneficial insectary plant <Emphasis Type="Italic">Scaevola aemula</Emphasis> to survival and long-term establishment of flightless <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in greenhouses

The fairy fan flower, Scaevola aemula R. Br., is a primary candidate insectary plant for maintaining populations of generalist predators. We conducted release experiments in greenhouses of cultivated eggplants to evaluate the effects of intercropping S. aemula on the establishment of flightless Harmonia axyridis Pallas. Compared with a monoculture of eggplant, all release experiments showed that flightless H. axyridis remained in greater numbers in plots with S. aemula planted alongside eggplant. In the release experiment of flightless H. axyridis larvae, the incidence of aphids in the plot with transplanted S. aemula was suppressed compared with that in the release plot without transplanted S. aemula. In a laboratory experiment, the longevity of flightless H. axyridis adults on blossom stems of S. aemula was greater than when open flowers and buds were removed, suggesting that the insects fed on floral resources such as the pollen of S. aemula. Our findings showed that the floral resources of S. aemula can enhance aphid suppression by improving the establishment of flightless H. axyridis.     

Evaluation of the role of the national parks for Persian leopard (<Emphasis Type="Italic">Panther pardus saxicolor</Emphasis>, Pocock 1927) habitat conservation (case study: Tandooreh National Park,Iran)

Having knowledge of the habitat requirements of top predators is important for their conservation and also for the stability of wildlife communities since they are keystone species of ecosystems. This study aims to predict suitable habitats for the Persian leopard (Panther pardus saxicolor, Pocock 1927) in Tandooreh National Park, Iran. This species is the largest leopard subspecies in the Middle East and is at high risk of extinction. Presence points for the species have been obtained from field data, literature review, atlas data, and IUCN records. Variables related to human disturbance, terrain, land use/land cover, and the leopards’ prey were used to predict the habitat selection of the Persian leopard by employing binary logistic regression. Our model predicted that the presence of Persian leopards is restricted to the southern, north-western, and western areas of the park which are under severe fragmentation threat. Moreover, leopards avoided areas near human development such as roads. We have also found that the availability of prey such as Capra aegagrus, Ovis vignei, and Ovis orientalis is one of the important parameters to predict habitat suitability for leopards. Our model provides a suitable tool for national park managers to identify biodiversity hotspots since leopards are often associated with high biodiversity.     

Potential for exploitative competition,not intraguild predation,between invasive harlequin ladybirds and flowerbugs in urban parks

In aphidophagous insect communities invaded by the harlequin ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae), intraguild predation (IGP) is widely implicated in the displacement of native predators, however, indirect trophic interactions are rarely assessed. Using molecular gut-content analysis, we investigated the relative frequencies of IGP by H. axyridis on the predatory flowerbug Anthocoris nemoralis Fabricius (Heteroptera: Anthocoridae) and prey overlap for a shared prey, the lime aphid Eucallipterus tiliae L. (Hemiptera: Aphididae), in Tilia × europaea crowns in urban parks. The frequency of IGP by H. axyridis was low: 2.7 % of larvae and 3.4 % of adults tested positive for A. nemoralis DNA. The presence of lime aphid DNA in predators was higher: 56.5 and 47.9 % of H. axyridis larvae and adults, respectively, contained E. tiliae DNA, whereas 60.8 % of adult A. nemoralis tested positive for aphid DNA. Incorporating insect densities revealed that the density of H. axyridis larvae had a strong negative effect on the likelihood of detecting aphid DNA in A. nemoralis. Prey overlap for E. tiliae was widespread in space (2–13 m height in tree crowns) and time (May–October 2011) which suggests that interspecific exploitative competition, mediated by predator life-stage, more so than IGP, is an important indirect trophic interaction between co-occurring H. axyridis and A. nemoralis. Whether exploitative competition equates to displacement of A. nemoralis populations requires further investigation. Our results emphasize the need to incorporate indirect interactions in studies of insect communities following invasion, not least because they potentially affect more species than direct interactions alone.     

Conspecific tolerance and heterospecific competition as mechanisms for overcoming resistance to invasion by an intertidal crab

The success of the invasive Asian shore crab, Hemigrapsus sanguineus, stems partly from its ability to exclude established crab species from preferred rocky and cobble intertidal habitat. Here, we assessed preference and competition for habitat types (cobble vs. sand) for H. sanguineus and two competitor species; the previous invasive green crab, Carcinus maenas, and the native rock crab, Cancer irroratus, in New England. In simple laboratory experiments, we paired similarly sized heterospecifics and conspecifics from each species, and also grouped combinations of C. maenas and H. sanguineus in a series of four-individual sets in order to dissect the outcome of intra- and inter-specific competition at different densities. Individually, all three species preferred cobble substrate. With paired conspecifics, H. sanguineus individuals would cohabitate in cobble, whereas C. maenas and C. irroratus individuals each excluded conspecifics from cobble. In heterospecific pairs, H. sanguineus excluded both C. maenas and C. irroratus from cobble. C. maenas and C. irroratus, were equally likely to exclude the other species, but rarely excluded H. sanguineus. In larger assemblages, H. sanguineus preferentially grouped under cobble, whereas C. maenas were more evenly distributed among habitat types. These observations demonstrate that conspecific tolerance and heterospecific competition can be effective, complementary mechanisms for overcoming invasion resistance. Such mechanisms help explain the well-studied success of H. sanguineus following its introduction into New England coastal habitats, and the resulting exclusion of preexisting crab species.     

An approach to bioassess pelagic ciliate biodiversity at different taxonomic resolutions in response to various habitats in the Amundsen Sea (Antarctica)

The rapid melting of glaciers and loss of sea ice will result in changes in habitat conditions that may drive substantial changes in biodiversity. In order to bioassess the changing polar ecosystem and evaluate biological conservation, pelagic ciliate communities at different taxonomic resolutions were studied at five habitats in the Amundsen Sea during the austral summer from December 2010 to January 2011. Distinctive spatial patterns were observed in the communities among the five habitats (oceanic areas, transitional areas, polynyas, edges of glaciers, and edges of sea ice) in response to environmental variability (e.g., temperature, salinity, chlorophyll a, and nutrients). The distributions in the numbers of different taxonomic levels and of three biodiversity indices (Shannon-Wiener H′, Pielou’s J′, and Margalef D) also revealed clear spatial variability with the maximum mean species number and indices in the polynya and maximum genus and family numbers in the transitional area. The presence/absence of data at taxonomic resolutions up to the family level provided sufficient information to evaluate the ecological patterns of pelagic ciliate communities and could accurately reflect habitat variations. The k-dominance curves illustrated clearly that maximum diversity was presented in the polynya at the species level and in the transitional area at the genus and family level. We suggest that the diversity at higher taxonomic resolutions should be considered more in future monitoring. Our findings provide basic data and an approach toward answering important questions about biological conservation, especially the biodiversity at various taxonomic resolutions in response to the increasing climate changes in polar ecosystems.     

Abiotic constraints and biotic resistance control the establishment success and abundance of invasive <Emphasis Type="Italic">Humulus japonicus</Emphasis> in riparian habitats

Dispersal, abiotic and biotic constraints are all involved in explaining the success of invasive plants but how these factors influence the different life stages of an invader remains poorly known. Focusing on highly invaded riparian habitats we asked: (1) how do propagule pressure, resource availability and resident vegetation influence the success of the invasive Asian vine Humulus japonicus at different stages of its life cycle (i.e. seedling, vegetative and flowering) (2) what is the influence of increasing resource availability on the performance and trait plasticity of H. japonicus compared to a functionally similar co-occurring native species? To answer the first question we performed a repeated field survey along the Gardon River (S France) with detailed measurements of distance to the riverbed, soil characteristics, light availability, and resident vegetation cover. To answer the second question we used a greenhouse experiment to compare the biomass and three functional traits of H. japonicus and Galium aparine along a gradient of increasing water and nitrogen availability. Initial germination success was only determined by abiotic constraints, while the role of biotic resistance increased for later stages with establishment success favoured by the interaction of low resident vegetation cover and high soil fertility, and final integrated success favoured by high light availability. H. japonicus performed better and showed higher plasticity in plant height than G. aparine under increased resource availability while their biomass did not differ in the lower part of the resource gradient. Our study demonstrates that by combining field and experimental studies and analysing responses at different life stages we can gain a more complete understanding of how ecological filters shape successful invasions in the course of the life cycle.     

Effect of habitat structure on reproduction and prey capture of a rare carnivorous plant, <Emphasis Type="Italic">Pinguicula lutea</Emphasis>

Habitat degradation is one of the greatest threats to biodiversity worldwide and the main contributor to the decline of many carnivorous plant species. For carnivorous plants in the southeastern United States, including many Pinguicula species (butterwort, Lentibulariaceae), degradation via altered fire regime has been implicated in their decline. Despite this decline, limited empirical research has been conducted examining the influence of habitat structural changes (through natural succession or human management) on reproduction and prey capture by carnivorous plants. The objectives of our study were to compare reproduction and prey capture for Pinguicula lutea (yellow butterwort) in habitats with different vegetation structures in the Florida Panhandle, where differences were largely due to management history. Pinguicula lutea is a self-compatible carnivorous plant that inhabits fire-dependent longleaf pine savannas of the southeastern United States and is threatened in the state of Florida. In 2014 and 2015, 13 sites were identified occupying three different habitat structures: maintained (intermittently mowed), grassy (dominated by Aristida stricta var. beyrichiana), and woody (encroachment by Hypericum and Ilex). Reproductive output was determined by assessing fruit set and ovule fertilization rate at each site. Additionally, prey availability and prey capture were assessed at each habitat site. In general, there were no differences in either measure of reproduction across habitat structure types. There were differences in prey abundance of Collembola, Diptera, and total arthropods both in terms of availability and capture. Total arthropod availability and prey capture were lowest in grassy sites compared to maintained habitat sites and woody habitat sites. Microclimatic conditions associated with each habitat structure and leaf morphology or physiology could explain the observed arthropod abundance and prey capture patterns. This study is the first ecological assessment of plant–insect interactions for Pinguicula species of the southeastern US and highlights the importance of habitat quality and management for this understudied group of carnivorous plants.     

Contrasting impacts of highly invasive plant species on flower-visiting insect communities

Invasive alien plants threaten biodiversity, ecosystems and service provision worldwide. They can have positive and negative direct and indirect effects on herbivorous insects, including those that provide pollination services. Here, we quantify how three highly invasive plant species (Heracleum mantegazzianum, Impatiens glandulifera and Fallopia japonica) influence the availability of floral resources and flower-visiting insect communities. We compared invaded with comparable uninvaded areas to assess floral resources and used pan-trapping to quantify insect communities. Only F. japonica influenced floral resource availability: sites invaded by this species had a higher flowering plant species richness and abundance of open floral units than uninvaded sites, probably due to its late flowering and the paucity of other flowering species at this time of year. Fallopia japonica was also associated with higher abundances of bumblebees, higher overall insect diversity and higher hoverfly diversity than uninvaded areas. Differences in pollinator communities were also associated with I. glandulifera and H. mantegazzianum, despite there being no detectable differences in floral resources at these sites. Specifically, there were more bumblebees and solitary bees in I. glandulifera sites, and a higher overall diversity of insects, particularly hoverflies. By contrast, H. mantegazzianum sites had a lower abundance of solitary bees and hoverflies. These findings confirm that invasive plant species have a range of species-specific effects on ecological communities. This supports the emerging view that control of invasive species, as required under international obligations, is not simple and that potential losses and gains for biodiversity must be carefully evaluated on a case-by-case basis.     

<Emphasis Type="Italic">Macrocystis integrifolia</Emphasis> and <Emphasis Type="Italic">Lessonia trabeculata</Emphasis> (Laminariales; Phaeophyceae) kelp habitat structures and associated macrobenthic community off northern Chile

Macrocystis integrifolia and Lessonia trabeculata form vast kelp beds providing a three-dimensional habitat for a diverse invertebrate and fish fauna off northern Chile. Habitat modifications caused by the El Niño Southern Oscillation (ENSO) are likely to alter the inhabiting communities. The aim of this study was to reveal relationships between distinct habitat structures of a M. integrifolia kelp bed, a dense L. trabeculata kelp bed and L. trabeculata patches colonizing a barren ground, and the associated dominant macrobenthic key species. Seasonally 15 sampling units (10 m² each) of any of the three habitats were monitored by SCUBA divers, which counted sporophytes and macroinvertebrates living between the latter. Furthermore, samples of plants were analysed in the laboratory to measure the morphological variables: total plant length, maximal holdfast diameter, stipe number, number of dichotomies per stipe, frond width and total drained wet mass. Multivariate analysis showed that the L. trabeculata kelp bed is denser, with a higher number of dichotomies per stipe, whereas sporophytes of M. integrifolia are longer with more stipes and wider fronds. Sporophytes of L. trabeculata patchily present on barren ground are shorter and have more stipes compared with those in the dense L. trabeculata kelp bed. Thus, the habitats provide different three-dimensional structures. The associated macrobenthic communities show a variable degree of overlapping; however, key faunal assemblages were distinguished for each habitat. Our study provides evidence that habitat diversity drives species diversity, the more homogeneous, monospecifically composed kelp bed habitats show comparatively low diversity, mainly caused by the dominance of the ascidian P. chilensis and T. tridentata in the M. integrifolia bed, and the mussel A. ater only present in the L. trabeculata bed. Species richness and diversity is highest in the heterogeneous habitat where L. trabeculata patches interrupt the barren ground. Our study revealed morphological differences between M. integrifolia and L. trabeculata kelp beds reflected in stipe number, plant length, dichotomies per stipe, and wet mass, which influence the composition of the associated characteristic fauna and its functional relations i.e. T. niger and T. tridentata.     

Parasites of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>: current research and perspectives

Harmonia axyridis (Coleoptera: Coccinellidae) has been introduced widely for biological control of agricultural pests. Harmonia axyridis has established in four continents outside of its native range in Asia and it is considered an invasive alien species (IAS). Despite a large body of work on invasion ecology, establishment mechanisms of IAS and their interactions with natural enemies remain open questions. Parasites, defined as multicellular organisms that do not directly kill the host, could potentially play an important role in regulating host populations. This study presents a review of the parasites of H. axyridis, discussing their distributions and effects on host populations across the host’s native and invasive range. These parasites are: Hesperomyces virescens Thaxt. fungi, Coccipolipus hippodamiae (McDaniel and Morrill) mites, and Parasitylenchus bifurcatus Poinar and Steenberg nematodes.     

Assessing the ecological risk posed by a recently established invasive alien predator: <Emphasis Type="Italic">Harmonia axyridis</Emphasis> as a case study

Invasive alien predators are a serious threat to biodiversity worldwide. However, there is no generic method for assessing which local species are most at risk following the invasion of a new predator. The harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), is an alien in Europe and many other parts of the world where it affects other species of ladybirds through competition for food and intra-guild predation (IGP). Here, we describe a method developed to assess which European ladybird species are most at risk following the invasion of H. axyridis. The three components of the risk assessment are: the likelihood that the assessed native species encounters H. axyridis in the field, the hazard of competition for food, and the IGP hazard. Thirty native European ladybird species were assessed through data obtained from field observations, laboratory experiments and literature reviews. The species that are considered most at risk are found on deciduous trees, have immature stages which are highly vulnerable to IGP by H. axyridis, and are primarily aphidophagous. These species should be the focus of specific studies and possibly conservation actions. The risk assessment method proposed here could be applied to other alien predators which are considered a threat to native species through competition and predation.     

Alpine bistort (<Emphasis Type="Italic">Bistorta vivipara</Emphasis>) in edge habitat associates with fewer but distinct ectomycorrhizal fungal species: a comparative study of three contrasting soil environments in Svalbard

Bistorta vivipara is a widespread arctic-alpine ectomycorrhizal (ECM) plant species. Recent findings suggest that fungal communities associated with B. vivipara roots appear random over short distances, but at larger scales, environmental filtering structure fungal communities. Habitats in highly stressful environments where specialist species with narrower niches may have an advantage represent unique opportunity to test the effect of environmental filtering. We utilised high-throughput amplicon sequencing to identify ECM communities associated with B. vivipara in Svalbard. We compared ECM communities in a core habitat where B. vivipara is frequent (Dryas-heath) with edge habitats representing extremes in terms of nutrient availability where B. vivipara is less frequent (bird-manured meadow and a nutrient-depleted mine tilling). Our analysis revealed that soil conditions in edge habitats favour less diverse but more distinct ECM fungal communities with functional traits adapted to local conditions. ECM richness was overall lower in both edge habitats, and the taxonomic compositions of ECM fungi were in line with our functional expectations. Stress-tolerant genera such as Laccaria and Hebeloma were abundant in nutrient-poor mine site whereas functional competitors genera such as Lactarius and Russula were dominant in the nutrient-rich bird-cliff site. Our results suggest that ECM communities in rare edge habitats are most likely not subsets of the larger pool of ECM fungi found in natural tundra, and they may represent a significant contribution to the overall diversity of ECM fungi in the Arctic.