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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Egg parasitoids have evolved adaptations to exploit host-associated cues, especially oviposition-induced plant volatiles and odors of gravid females, when foraging for hosts. The entire host selection process is critical for successful parasitism and relevant in defining host specificity of parasitoids. We hypothesized that naïve egg parasitoid females reared on their coevolved host are able to exploit cues related to the coevolved host but not those from a novel host. We used the egg parasitoid Trissolcus japonicus , its coevolved host Halyomorpha halys , and the non-coevolved host Podisus maculiventris to evaluate this hypothesis. H. halys , a polyphagous pest native from Eastern Asia, has invaded North America and Europe, resulting in serious damage to crops. T. japonicus is the most effective egg parasitoid of H. halys in its native area and thus considered a major candidate for biological control. This parasitoid was detected in North America and Europe as a result of accidental introductions. Laboratory host range of T. japonicus includes P. maculiventris, an American predatory stink bug used as a biological control agent of several pests. Using T. japonicus reared on its natural host H. halys , we tested in a Y-tube olfactometer the responses of naïve parasitoid females to volatiles from tomato plants with a deposited egg mass and feeding punctures of either H. halys or P. maculiventris . Additionally, using two different olfactometer set-ups, we tested T. japonicus responses to volatiles emitted by eggs and mature males and females of H. halys or P. maculiventris . Tomato plants subjected to oviposition and feeding by H. halys were preferred by the wasp compared to clean plants, suggesting a possible activation of an indirect defense mechanism. Furthermore, T. japonicus females were attracted by cues from gravid females and mature males of H. halys but not from eggs. By contrast, naïve parasitoid females never responded to cues associated with P. maculiventris , although this non-target host is suitable for complete parasitoid development. Such lack of responses might reduce the probability of T. japonicus locating and parasitizing P. maculiventris under field conditions. Our experimental approach properly simulates the parasitoid host-location process and could be combined with the required host specificity tests for risk assessment in biological control programs.

Since the late 19th century, exotic ladybirds (Coleoptera: Coccinellidae) have been used extensively for suppressing herbivorous insects of economic importance. In recent decades, the introduction of non-native biological control (BC) agents has been greatly limited due to the awareness of the potential non-target effects of introductions. Nonetheless, recent episodes of biological invasions of economically important pests have raised the need to carefully consider whether the expected benefits of pest control go beyond the possible environmental risks of introduction. To better understand the factors that contributed to successful BC programs, here we review the literature behind classical and augmentative BC using exotic ladybirds. Additionally, by means of case studies, we discuss the BC efficacy of selected exotic species, e.g., Coccinella septempunctata L., Harmonia axyridis (Pallas), and Hippodamia variegata (Goeze), and their position within the communities of predators in the introduced areas of USA, Canada, and Chile. In Europe, much of the research on exotic ladybirds has been conducted on the undesired impact of H. axyridis. Therefore, we summarize the risk assessment data for this species and review the field research investigating the ecological impact on European aphidophagous predators. According to the BIO-CAT database of classical BC programs, 212 ladybird species belonging to 68 genera have been released in about 130 years of BC activity, with 14.6% of introductions having resulted in partial, substantial, or complete control of the target pest. However, because post-release evaluation of establishment and BC success has not always been conducted, this rate could underestimate the successful cases. Among other factors, ladybird establishment and pest suppression mostly depend on (1) intrinsic factors, i.e., high voracity, synchronized predator-prey life cycle, and high dispersal ability, and (2) extrinsic factors, i.e., adaptability to the new environment and landscape composition. This review contributes to improved understanding of ladybirds as exotic BC agents.