Artículo de publicación ISIThe 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

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Inferences about introduction histories of invasive species remain challenging because of the stochastic demographic processes involved. Approximate Bayesian computation (ABC) can help to overcome these problems, but such method requires a prior understanding of population structure over the study area, necessitating the use of alternative methods and an intense sampling design. In this study, we made inferences about the worldwide invasion history of the ladybird Harmonia axyridis by various population genetics statistical methods, using a large set of sampling sites distributed over most of the species' native and invaded areas. We evaluated the complementarity of the statistical methods and the consequences of using different sets of site samples for ABC inferences. We found that the H. axyridis invasion has involved two bridgehead invasive populations in North America, which have served as the source populations for at least six independent introductions into other continents. We also identified several situations of genetic admixture between differentiated sources. Our results highlight the importance of coupling ABC methods with more traditional statistical approaches. We found that the choice of site samples could affect the conclusions of ABC analyses comparing possible scenarios. Approaches involving independent ABC analyses on several sample sets constitute a sensible solution, complementary to standard quality controls based on the analysis of pseudo-observed data sets, to minimize erroneous conclusions. This study provides biologists without expertise in this area with detailed methodological and conceptual guidelines for making inferences about invasion routes when dealing with a large number of sampling sites and complex population genetic structures.

Brown, Peter M.J.; Bizama, Gustavo. 2016. Rapid spread of Harmonia axyridis in Chile and its effects on local coccinellid biodiversity. Diversity and Distributions, 22 (9). 982-994. 10.1111/ddi.12455 Contact CEH NORA team at noraceh@ceh.ac.ukThe NERC and CEH trademarks and logos ('the Trademarks') are registered trademarks of NERC in the UK and other countries, and may not be used without the prior written consent of the Trademark owner. The spread and impact of this species has been documented in Europe and North America, but no 28 information exists for South America. The aims of this study were to: 1) document the process of 29 invasion of H. axyridis in Chile, 2) compare the abundance of H. axyridis in different habitats with 30 varying degrees of disturbance, and 3) assess change in the coccinellid assemblages in alfalfa fields 31 over the six years following invasion. 32 Location Chile 33Methods The spread of H. axyridis was estimated using information from citizen scientists alongside 34 records from the National Pest Surveillance System. The abundance of H. axyridis in different habitat 35 types and of all coccinellids in alfalfa fields was assessed using yellow sticky traps. In alfalfa, the 36 variations in species richness, Shannon and Simpson diversity and equitability indices through time 37 were compared. 38

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We conducted an experimental landscape study to test the hypotheses that: (1) habitat removal results in short-term increases in population density in the remaining habitat patches (the crowding effect); (2) following habitat removal, density is higher in landscapes with more, smaller patches and more habitat edge (i.e., a higher level of habitat fragmentation per se) than in less fragmented landscapes, for the same total amount of habitat on the landscapes; (3) this positive effect of fragmentation per se on density is larger in landscapes with smaller inter-patch distances; and (4) these last two effects should be reduced or disappear over time following habitat removal. Our results did not support the first hypothesis, but they provided some support for the other three hypotheses, for two of the four Coccinellid species studied. As in other empirical studies of fragmentation per se on population density, the effects of fragmentation per se were weak and positive (when they did occur). This is the first study to document a transient effect of fragmentation per se on population density, and to show that this effect depends on inter-patch distances. We suggest that fragmentation per se increased the rate of immigration to patches, resulting in higher population densities in more fragmented landscapes.

The resource concentration hypothesis (Root 1973) predicts that specialist herbivorous insects should be more abundant in large patches of host plants, because the insects are more likely to find and stay longer in those patches. Between August 1989 and January 1990 we experimentally tested Root's hypothesis by analyzing the numerical response of four species of herbivorous insects associated with patches of 4, 16, 64 and 225 cabbage plants, Brassica oleracea var. capitata. In addition, we studied the colonization of patches by adults of Plutella xylostella (L.) (Lepidoptera: Plutellidae), and the migration of their larvae in patches of different sizes. No herbivorous insect densities differed significantly with patch size. Adults of P. xylostella colonized all kind of patches equally. Larvae did not migrate between patches, and their disappearance rate did not differ between patches. The resource concentration hypothesis is organism-dependent, being a function of the adult and juvenile herbivore dispersal behavior in relation to the spatial scale of patchiness.

Aim Both anthropogenic habitat disturbance and the breadth of habitat use by alien species have been found to facilitate invasion into novel environments, and these factors have been hypothesized to be important within coccinellid communities specifically. In this study, we address two questions: (1) Do alien species benefit more than native species from human-disturbed habitats? (2) Are alien species more generalized in their habitat use than natives within the invaded range or can their abundance patterns be explained by specialization on the most common habitats? Location Chile.Methods We quantified the use of different habitat types by native and alien coccinellid beetles by sampling individuals in nine habitat types that spanned a gradient in disturbance intensity, and represented the dominant natural, seminatural and agricultural habitats in the landscape. ResultsOur results provide strong support for the hypotheses that moredisturbed habitats are differentially invaded. Both the proportion of alien individuals and the proportion of alien species increased significantly with increasing disturbance intensity. In contrast, we found no evidence that alien species were more generalized in their habitat use than native species; in fact, the trend was in the opposite direction. The abundance of specialized alien coccinellid species was not correlated with the area of the habitat types in the landscape.Main conclusion The results suggest that successfully established alien coccinellid species may be 'disturbance specialists' that thrive within human-modified habitats. Therefore, less-disturbed agroecosystems are desirable to promote the regional conservation of native species within increasingly human-dominated landscapes.