Tim Adriaens

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

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

The harlequin ladybird, Harmonia axyridis: global perspectives on invasion history and ecology

The harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), is native to Asia but has been intentionally introduced to many countries as a biological control agent of pest insects. In numerous countries, however, it has been introduced unintentionally. The dramatic spread of H. axyridis within many countries has been met with considerable trepidation. It is a generalist top predator, able to thrive in many habitats and across wide climatic conditions. It poses a threat to biodiversity, particularly aphidophagous insects, through competition and predation, and in many countries adverse effects have been reported on other species, particularly coccinellids. However, the patterns are not consistent around the world and seem to be affected by many factors including landscape and climate. Research on H. axyridis has provided detailed insights into invasion biology from broad patterns and processes to approaches in surveillance and monitoring. An impressive number of studies on this alien species have provided mechanistic evidence alongside models explaining large-scale patterns and processes. The involvement of citizens in monitoring this species in a number of countries around the world is inspiring and has provided data on scales that would be otherwise unachievable. Harmonia axyridis has successfully been used as a model invasive alien species and has been the inspiration for global collaborations at various scales. There is considerable scope to expand the research and associated collaborations, particularly to increase the breadth of parallel studies conducted in the native and invaded regions. Indeed a qualitative comparison of biological traits across the native and invaded range suggests that there are differences which ultimately could influence the population dynamics of this invader. Here we provide an overview of the invasion history and ecology of H. axyridis globally with consideration of future research perspectives. We reflect broadly on the contributions of such research to our understanding of invasion biology while also informing policy and people.

Harmonia + and Pandora + : risk screening tools for potentially invasive plants, animals and their pathogens

AbstractGiven the large number of alien species that may potentially develop into invasives, there is a clear need for robust schemes that allow to screen species for such risks. The Harmonia+ framework presented here brings together 30 questions that refer to distinct components of invasion. Together, they cover the stages of introduction, establishment, spread, and multiple kinds of impacts, viz. referring to the health of the environment (including wild species), cultivated plants, domesticated animals and man. In a complete assessment, input is provided by choosing among predefined ordinal answers and by supplementing these with textual clarification. Uncertainty is covered by indicating levels of confidence. By converting answers into scores, which are then condensed into summary statistics, Harmonia+ allows for quantitative output on stage-specific and general risks. Test assessments on five species emerging in Belgium showed the perceived environmental risks of Procambarus clarkii to be highest (0.72), and that of Threskiornis aethiopicus to be lowest (0.13). Given the considerable parallels that exist between invasive alien species and emerging infectious diseases, we additionally created Pandora, which is a risk analysis scheme for pathogens and parasites. It consists of 13 key questions and has the same structure as Harmonia+. Since diseases play a paramount role in biological invasions, results of Pandora assessments may feed into Harmonia+ through a slightly adapted, host-specific version named Pandora+. Harmonia+, Pandora and Pandora+ may be used both for prioritization purposes and for underpinning detailed risk analyses, and can be consulted online through http://ias.biodiversity.be.

Assessing the ecological risk posed by a recently established invasive alien predator: Harmonia axyridis 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.

Trans-national horizon scanning for invasive non-native species: a case study in western Europe

Horizon scanning for high-risk invasive non-native species (INNS) is crucial in preparing and implementing measures to prevent introductions, as well as to focus efforts in the control of species already present. We initiated a trans-national horizon-scanning exercise focused on four countries in western Europe: Great Britain, France, Belgium and The Netherlands, which share similar environmental and socio-economic characteristics. We followed a structured four-step approach combining existing knowledge about INNS, with a collaborative identification of priorities for research and management: (1) systematic review of potential INNS of concern, (2) discrimination of INNS into an Alert and Black List depending on their absence or presence in the study area respectively, (3) risk analysis of the Alert List, and (4) expert ranking of species in the Black List. Amongst species not yet present in the four countries (i.e. Alert List), assessors reliably pointed to the Emerald ash-borer (Agrilus planipennis) and Sosnowski’s hogweed (Heracleum sosnowskyi) as those INNS with the highest risk of invasion and impact. The Japanese knotweed (Fallopia japonica), Himalayan balsam (Impatiens glandulifera), zebra mussel (Dreissena polymorpha) and killer shrimp (Dikerogammarus villosus) were consistently highlighted as some of the most problematic INNS already present in the study area (i.e. Black List). The advantages of our combined approach include that it is inclusive of all-taxa, prioritizes both established and emerging biological threats across trans-national scales, and considers not only the ecological impact, but also potential direct economic consequences as well as the manageability of invasive species.

Testing the applicability of regional IUCN Red List criteria on ladybirds (Coleoptera, Coccinellidae) in Flanders (north Belgium): opportunities for conservation

Red Lists assess the extinction risk of species and are an important tool to prioritise species conservation and management measures. Worldwide, quantitative IUCN criteria are used to estimate the threat status of species at the regional level. In Flanders (north Belgium), about 70 000 distribution records of ladybirds were collected in 36% of all the grid cells (1 × 1 km2) since 1990 during a large‐scale citizen‐science project. Applying the IUCN criteria to the 36 resident ‘conspicuous’ ladybirds in Flanders resulted in two Regionally Extinct species, three Endangered species and six Vulnerable species. A further seven species were considered Near Threatened and the remaining 15 species (39%) were assessed as Least Concern. Three species were classified as Data deficient. Using the Red List status, we delineated ladybird hotspots that were mainly located in grid cells with large areas of Natura2000 sites. For calculating a distribution trend, we advocate the use of a high grid cell resolution (e.g. 1 × 1 km2 or 5 × 5 km2). The ladybird data set from Flanders provides evidence that IUCN Red List criteria can be applied to this charismatic, but relatively under‐surveyed insect group. For estimating the geographic range, the use of extent of occurrence instead of area of occupancy is advisable when the survey coverage is relatively low. We discuss the opportunities of the availability of a ladybird Red List for regional conservation measures.

Female morphs of a colour polymorphic damselfly differ in developmental instability and fecundity

Sex-limited colour polymorphism occurs in several animal taxa and is usually explained in the context of sexual selection. Specifically, for polymorphism restricted to the female sex, multiple phenotypes may have evolved in response to male harassment. Such male harassment is generally considered to entail differential costs to female morphs, which may ultimately result in fitness differences. However, contrary to this prediction, most previous studies do not support that female morphs (andromorphs and heteromorphs) differ in measures of quality and (or) fitness components. In this study, we evaluate quality and fitness differences between mated female morphs of the damselfly Enallagma cyathigerum . We suggest that many earlier studies may have failed to observe morph differences in quality or fitness because selection by male harassment was weak. Here, we selected a study population for which our expectation was that levels of per female capita male harassment were high. Nevertheless, also in this population mated female morphs did not differ in body size or condition (body mass/body length). However, mated female morphs did differ in levels of developmental instability: heteromorphs consistently showed a higher level of fluctuating asymmetry than andromorphs. Also, mated female morphs differed in fecundity: andromorphs had a lower clutch size than heteromorphs. In addition, larger females contained more eggs, but the slope of this relationship was steeper in heteromorphs. In conclusion, mated female morphs of the damselfly E. cyathigerum at our study site clearly differed in one quality estimate (developmental instability) and in our measure of fitness (fecundity).

Developing a framework of minimum standards for the risk assessment of alien species

1. Biological invasions are a threat to biodiversity, society and the economy. There is an urgent need to provide evidence-based assessments of the risks posed by inva-sive alien species (IAS) to prioritize action. Risk assessments underpin IAS policies in many ways: informing legislation; providing justification of restrictions in trade or consumer activities; prioritizing surveillance and rapid response. There are benefits to ensuring consistency in content of IAS risk assessments globally, and this can be achieved by providing a framework of minimum standards as a checklist for quality assurance. 2. From a review of existing risk assessment protocols, and with reference to the r equirements of the EU Regulation on IAS (1143/2014) and international agreements including the World Trade Organisation, Convention on Biological Diversity and International Plant Protection Convention, coupled with consensus methods, we identified and agreed upon 14 minimum standards (attributes) a risk-assessment scheme should include. 3. The agreed minimum standards were as follows: (1) basic species description; (2) likelihood of invasion; (3) distribution, spread and impacts; (4) assessment of intro-duction pathways; (5) assessment of impacts on biodiversity and ecosystems; (6) Assessment of impact on ecosystem services; (7) assessment of socio-economic impacts; (8) consideration of status (threatened or protected) of species or habitat under threat; (9) assessment of effects of future climate change; (10) completion possible even when there is a lack of information; (11) documents information sources; (12) provides a summary in a consistent and interpretable form; (13) includes uncertainty; (14) includes quality assurance. In deriving these minimum standards, gaps in knowledge required for completing risk assessments and the scope of exist-ing risk assessment protocols were revealed, most notably in relation to assessing benefits, socio-economic impacts and impacts on ecosystem services but also inclu-sion of consideration of climate change. 4. Policy implications. We provide a checklist of components that should be within in-vasive alien species risk assessments and recommendations to develop risk assess-ments to meet these proposed minimum standards. Although inspired by implementation of the European Union Regulation on invasive alien species, and as such developed specifically within a European context, the derived framework and minimum standards could be applied globally.

Stakeholder engagement in the study and management of invasive alien species

Invasive alien species are a major driver of global environmental change and a range of management interventions are needed to manage their effects on biodiversity, ecosystem services, human well-being and local livelihoods. Stakeholder engagement is widely advocated to integrate diverse knowledge and perspectives in the management of invasive species and to deal with potential conflicts of interest. We reviewed the literature in the ISI Web of Science on stakeholder engagement (the process of involving stakeholders (actors) in decision making, management actions and knowledge creation) in invasion science to assess and understand what has been done (looking at approaches and methodologies used, stakeholders involved, and outcomes from engagement) and to make recommendations for future work. Research on stakeholder engagement in invasion science has increased over the last decade, helping to improve scientific knowledge and contributing towards policy formulation and co-implementation of management. However, many challenges remain and engagement could be made more effective. For example, most studies engage only one stakeholder group passively using questionnaires, primarily for assessing local knowledge and perceptions. Although useful for management and policy planning, these stakeholders are not active participants and there is no two-way flow of knowledge. To make stakeholder involvement more useful, we encourage more integrative and collaborative engagement to (1) improve co-design, co-creation and co-implementation of research and management actions; (2) promote social learning and provide feedback to stakeholders; (3) enhance collaboration and partnerships beyond the natural sciences and academia (interdisciplinary and transdisciplinary collaboration); and (4) discuss some practical and policy suggestions for improving stakeholder engagement in invasion science research and management. This will help facilitate different stakeholders to work better together, allowing problems associated with biological invasions to be tackled more holistically and successfully.

Seasonal and diurnal variation in the proportions of female morphs of the damselfly Enallagma cyathigerum

In many damselfly species a female-limited colour polymorphism is encountered which is assumed to be the result of sexual conflict. Typically, one morph resembles the males body colouration (andromorph), while the other is dissimilar (heteromorph). Little is known about the extent of temporal variation in female morph proportions at the water where mating occurs. Knowledge about such variation should help to identify the factors that affect female morph proportion and the scales at which these factors operate. The objective of this study is to assess the occurrence of diurnal and seasonal variation in female morph proportions at the water for the damselfly Enallagma cyathigerum. Diurnal variation was evaluated at six nearby populations, while seasonal variation was examined at one of these populations. Furthermore, we considered temporal variation in female morph proportion in relation to proxies of male harassment (i.e., male density and operational sex ratio). Our findings indicate that female morph proportion varies throughout a day but is uniform on a seasonal scale. Variation in female morph proportions could not be explained by concomitant variation in male density or operational sex ratio. We suggest future study of male mate choice may consider temporal variation in female morph proportions at the water.