Richard F. Comont

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.

Escape from parasitism by the invasive alien ladybird, Harmonia axyridis

Alien species are often reported to perform better than functionally similar species native to the invaded range, resulting in high population densities, and a tendency to become invasive. The enemy release hypothesis (ERH) explains the success of invasive alien species (IAS) as a consequence of reduced mortality from natural enemies (predators, parasites and pathogens) compared with native species. The harlequin ladybird, Harmonia axyridis, a species alien to Britain, provides a model system for testing the ERH. Pupae of H. axyridis and the native ladybird Coccinella septempunctata were monitored for parasitism between 2008 and 2011, from populations across southern England in areas first invaded by H. axyridis between 2004 and 2009. In addition, a semi‐field experiment was established to investigate the incidence of parasitism of adult H. axyridis and C. septempunctata by Dinocampus coccinellae. Harmonia axyridis pupae were parasitised at a much lower rate than conspecifics in the native range, and both pupae and adults were parasitised at a considerably lower rate than C. septempunctata populations from the same place and time (H. axyridis: 1.67%; C. septempunctata: 18.02%) or in previous studies on Asian H. axyridis (2–7%). We found no evidence that the presence of H. axyridis affected the parasitism rate of C. septempunctata by D. coccinellae. Our results are consistent with the general prediction that the prevalence of natural enemies is lower for introduced species than for native species at early stages of invasion. This may partly explain why H. axyridis is such a successful IAS.

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

AbstractFive 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.

Alien arthropod predators and parasitoids: interactions with the environment

Many species of entomophagous arthropods have been introduced either intentionally (through the practice of biological control) or unintentionally to new regions. We examine interactions of these aliens with their new environments in the context of rapid global change linked to human activity. We consider effects of such interactions on establishment and spread of the alien species and effects on indigenous biota and ecosystems. Major elements of global change that affect alien-environment interactions include landscape modifications by humans (e.g., cultivation, habitat loss and fragmentation) and increases in atmospheric CO2 and other gases resulting in climate change and other effects (e.g., changes in food quality for herbivores that affect higher trophic levels as well). Alien arthropod predators can alter landscapes for their benefit, to the detriment of indigenous species. A brief review also of blood-feeding alien arthropods makes clear that interactions with the environment critically influence invasions of zoophagous arthropods in general.

Evaluating promotional approaches for citizen science biological recording: bumblebees as a group versus Harmonia axyridis as a flagship for ladybirds

Over the past decade, the number of biological records submitted by members of the public have increased dramatically. However, this may result in reduced record quality, depending on how species are promoted in the media. Here we examined the two main promotional approaches for citizen science recording schemes: flagship-species, using one charismatic species as an umbrella for the entire group (here, Harmonia axyridis (Pallas) for Coleoptera: Coccinellidae), and general-group, where the group is promoted as a whole and no particular prominence is given to any one species (here, bumblebees, genus Bombus (Hymenoptera: Apidae)). Of the two approaches, the general-group approach produced data that was not biased towards any one species, but far fewer records per year overall. In contrast, the flagship-species approach generated a much larger annual dataset, but heavily biased towards the flagship itself. Therefore, we recommend that the approach for species promotion is fitted to the result desired.