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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.

Native to Australasia, Eucalyptus (sensu lato) is one of the most planted genera of trees in the world. However, the sustainability of Eucalyptus species as plantation trees in non-native areas is increasingly threatened by the introduction and spread of Eucalyptus-feeding insects from Australia. We examine patterns and potential trends with respect to the global spread of Eucalyptus-feeding insects. Likely pathways of introduction and drivers of the rapid distribution of these insects, as well as management options are considered. The rate of introductions is shown to have increased nearly fivefold since the 1980s. As a result, the number of non-native pests of eucalypts outside of Australia has doubled in less than three decades. Furthermore, the rate of secondary spread among continents has also increased. Surprisingly, we found no association between area planted and the number of pests or new introductions. Only a small number of countries have been the points of first detection outside the native range; these countries have acted as bridgeheads to other regions. Quarantine regulations aimed at reducing the spread of invasive organisms appear to be ineffective at a global scale, and pathways allowing these invasions to occur are poorly understood or unknown. An expanded suite of management options are needed to provide resilience against the rapid accrual and homogenization of eucalypt pests, thereby ensuring the sustainability of eucalypt forestry worldwide.

Globalization is triggering an increase in the establishment of alien insects in Europe, with several species having substantial ecological and economic impacts. We investigated long-term changes in rates of species spread following establishment. We used the total area of countries invaded by 1171 insect species for which the date of first record in Europe is known, to estimate their current range radius (calculated as [invaded area](0.5)/pi). We estimated initial rates of radial spread and compared them among different groups of insects for all years (1800-2014) and for a subset of more recent decades (1950-2014). Accidentally introduced species spread faster than intentionally introduced species. Considering the whole period 1800-2014, spread patterns also differ between feeding guilds, with decreasing spread rates over residence time in herbivores but not in detritivores or parasitic species. These decreases for herbivorous species appeared mainly in those associated with herbaceous plants and crops rather than woody plants. Initial spread rate was significantly greater for species detected after 1990, roughly 3-4 times higher than for species that arrived earlier. We hypothesize that the political changes in Europe following the collapse of the Iron Curtain in 1989, and the further dismantling of customs checkpoints within an enlarged European Union (EU) have facilitated the faster spread of alien insect species. Also, the number of species first recorded in the Eastern Bloc of the politically-divided Europe before 1989 was lower than for the rest of Europe. A detailed analysis of six recent invaders indicated a dominant role of long-distance translocations related to human activities, especially with the plant trade, in determining rates of spread.

The advent of simple and affordable tools for molecular identification of novel insect invaders and assessment of population diversity has changed the face of invasion biology in recent years. The widespread application of these tools has brought with it an emerging understanding that patterns in biogeography, introduction history and subsequent movement and spread of many invasive alien insects are far more complex than previously thought. We reviewed the literature and found that for a number of invasive insects, there is strong and growing evidence that multiple introductions, complex global movement, and population admixture in the invaded range are commonplace. Additionally, historical paradigms related to species and strain identities and origins of common invaders are in many cases being challenged. This has major consequences for our understanding of basic biology and ecology of invasive insects and impacts quarantine, management and biocontrol programs. In addition, we found that founder effects rarely limit fitness in invasive insects and may benefit populations (by purging harmful alleles or increasing additive genetic variance). Also, while phenotypic plasticity appears important post-establishment, genetic diversity in invasive insects is often higher than expected and increases over time via multiple introductions. Further, connectivity among disjunct regions of global invasive ranges is generally far higher than expected and is often asymmetric, with some populations contributing disproportionately to global spread. We argue that the role of connectivity in driving the ecology and evolution of introduced species with multiple invasive ranges has been historically underestimated and that such species are often best understood in a global context.

During the last two centuries, thousands of insect species have been transported (largely inadvertently) and established outside of their native ranges worldwide, some with catastrophic ecological and economic impacts. Global variation in numbers of invading species depends on geographic variation in propagule pressure and heterogeneity of environmental resistance to invasions. Elton’s diversity-invasibility hypothesis, proposed over sixty years ago, has been widely explored for plants but little is known on how biodiversity affects insect invasions. Here we use species inventories from 44 land areas, ranging from small oceanic islands to entire continents in various world regions, to show that numbers of established insect species are primarily driven by diversity of plants, with both native and non-native plant species richness being the strongest predictor of insect invasions. We find that at large spatial scales, plant diversity directly explains variation in non-native insect species richness among world regions, while geographic factors such as land area, climate and insularity largely affect insect invasions indirectly via their effects on local plant richness.

The success of a biological invasion is context dependent, and yet two key concepts—the invasiveness of species and the invasibility of recipient ecosystems—are often defined and considered separately. We propose a framework that can elucidate the complex relationship between invasibility and invasiveness. It is based on trait-mediated interactions between species and depicts the response of an ecological network to the intrusion of an alien species, drawing on the concept of community saturation. Here, invasiveness of an introduced species with a particular trait is measured by its per capita population growth rate when the initial propagule pressure of the introduced species is very low. The invasibility of the recipient habitat or ecosystem is dependent on the structure of the resident ecological network and is defined as the total width of an opportunity niche in the trait space susceptible to invasion. Invasibility is thus a measure of network instability. We also correlate invasibility with the asymptotic stability of resident ecological network, measured by the leading eigenvalue of the interaction matrix that depicts trait-based interaction intensity multiplied by encounter rate (a pairwise product of propagule pressure of all members in a community). We further examine the relationship between invasibility and network architecture, including network connectance, nestedness and modularity. We exemplify this framework with a trait-based assembly model under perturbations in ways to emulate fluctuating resources and random trait composition in ecological networks. The maximum invasiveness of a potential invader (greatest intrinsic population growth rate) was found to be positively correlated with invasibility of the recipient ecological network. Additionally, ecosystems with high network modularity and high ecological stability tend to exhibit high invasibility. Where quantitative data are lacking we propose using a qualitative interaction matrix of the ecological network perceived by a potential invader so that the structural network stability and invasibility can be estimated from the literature or from expert opinion. This approach links network structure, invasiveness and invasibility in the context of trait-mediated interactions, such as the invasion of insects into mutualistic and antagonistic networks.

A literature survey identified 403 primary research publications that investigated the ecological effects of invasive alien insects and/or the mechanisms underlying these effects. The majority of these studies were published in the last 8 years and nearly two-thirds were carried out in North America.These publications concerned 72 invasive insect species, of which two ant species, Solenopsis invicta and Linepithema humile, accounted for 18% and 14% of the studies, respectively.Most publications investigated effects on native biodiversity at population or community level. Genetic effects and, to a lesser extent, effects on ecosystem services and processes were rarely explored. We review the effects caused by different insect invaders according to: their ecosystem roles, i.e. herbivores, predators, parasites,parasitoids and pollinators; the level of biological organisation at which they occur; and the direct and indirect mechanisms underlying these effects. The best documented effects occur in invasive ants,Eurasian forest herbivores invasive in North America,and honeybees. Impacts may occur through simple trophic interactions such as herbivory, predation or parasitism. Alien species may also affect native species and communities through more complex mechanisms such as competition for resources,disease transmission, apparent competition, or pollination disruption, among others. Finally, some invasive insects, particularly forest herbivores and ants, are known to affect ecosystem processes through cascading effects. We identify biases and gaps in our knowledge of ecological effects of invasive insects and suggest further opportunities for research.

Insect species associated with human goods continue to be accidentally introduced into new locations. A small proportion of these introduced species become invasive, causing a range of impacts in the receiving community. It is therefore important to evaluate the patterns of which species become invasive and which strategies are most successful in managing them. This review assesses the distribution, abundance, impact and management of the invasive Vespidae worldwide. We identified 34 vespid species known to be introduced around the world, but the seven most invasive species are all eusocial. Most introduced Vespidae only occur in one or two countries, but some areas have become geographic hotspots of invasion: Hawaii (15 species), North America (eight species), New Zealand (five species), Australia (four species) and South America (four species). Two invasive species, Vespula vulgaris and V. germanica have become particularly widespread and abundant with a range of impacts on biodiversity and ecosystem function. Other successful invasive species include several Polistes spp., which affect local biodiversity through direct predation or competition for food or space. Toxic baiting has been the most successful control strategy against invasive vespids to date, although this has mostly been small scale experimental management as it has proved difficult to develop commercial control products. Development of shelf-stable lures or baits combined with suitable toxins or pathogens could overcome some of the commercial impediments. Several attempts at biological control using parasitoids have not successfully reduced invasive wasp populations, although the biocontrol agent has only established in one case. The social structure of colonies and their high reproductive efficiency have facilitated invasion by these species, but it also means management at the population level will be difficult. This emphasises the need to prevent such invasions from occurring in the first place.

Biological control is a valuable and effective strategy for controlling arthropod pests and has been used extensively against invasive arthropods. As one approach for control of invasives, exotic natural enemies from the native range of the pest are introduced to areas where control is needed. Classical biological control began to be used in the late 1800s and its use increased until, beginning in 1983, scientists began raising significant concerns and questions about nontarget and indirect effects that can be caused by these introductions. In recent years, similar issues have been raised about augmentative use of exotic natural enemies. Subsequently, international guidelines, national regulations and scientific methods being used for exotic natural enemies in biological control have changed to require appropriate specificity testing, risk assessment and regulatory oversight before exotic natural enemies can be released. National and international standards aimed at minimizing risk have increased awareness and promoted more careful consideration of the costs and benefits associated with biological control. The barriers to the implementation of classical and augmentative biological control with exotic natural enemies now are sometimes difficult and, as a consequence, the numbers of classical biological control programs and releases have decreased significantly. Based in part on this new, more careful approach, classical biological control programs more recently undertaken are increasingly aimed at controlling especially damaging invasive arthropod pests that otherwise cannot be controlled. We examine evidence for these revised procedures and regulations aimed at increasing success and minimizing risk. We also discuss limitations linked to the apparent paucity of postintroduction monitoring and inherent unpredictability of indirect effects.