Harmonia axyridis (Coleoptera: Coccinellidae) has been introduced widely for biological control of agricultural pests. Harmonia axyridis has established in four continents outside of its native range in Asia and it is considered an invasive alien species (IAS). Despite a large body of work on invasion ecology, establishment mechanisms of IAS and their interactions with natural enemies remain open questions. Parasites, defined as multicellular organisms that do not directly kill the host, could potentially play an important role in regulating host populations. This study presents a review of the parasites of H. axyridis, discussing their distributions and effects on host populations across the host's native and invasive range. These parasites are: Hesperomyces virescens Thaxt. fungi, Coccipolipus hippodamiae (McDaniel and Morrill) mites, and Parasitylenchus bifurcatus Poinar and Steenberg nematodes.

a b s t r a c tWe review aspects of the life histories of representative enemies of coccinellids (both entomophagous and phytophagous species) and expose both potential and real effects that they have on life parameters of their hosts. Lady beetles are attacked by a variety of natural enemies (bacteria, fungi, mites, nematodes, protozoa, wasps, flies). Few of these enemies have the ability to alter significantly the population dynamics of their hosts. This review should encourage further research to help define the role of natural enemies in the population dynamics of coccinellids. Ultimately, the conservation of beneficial lady beetles and the management of nuisance and pestiferous ones should be major emphases of research on coccinellid-natural enemy interactions.Published by Elsevier Inc.

Plants use trichomes as a morphological defense against attacks from herbivores. The literature was reviewed to test the hypothesis that trichome-bearing (pubescent) plants do not cause more harm than good to predators. Forty seven records on interactions between plant trichomes and predatory insects were found. Overall, the records reveal that trichomes have more harmful than beneficial effects on predators. Fortunately, most harmful effects are sublethal; they usually affect movement, development, oviposition and predation potential. In worst cases, sticky exudates from glandular trichomes entrap predators. The hooked tips on non-glandular trichomes impale predators. Entrapped and impaled predators often die from desiccation or starvation. Plant cultivars with high (rather than low) trichome density cause the most harm, and trichomes on tomato and some beans often cause more harm than good to predatory beetles, true bugs and lacewings. Whether these harmful effects have a net negative effect on plant fitness is poorly known and ripe for study. When developing and testing cultivars with increased trichome-based resistance to herbivory, the question as to whether these technologies are compatible with the functional role of those predators (single or combined species) capable of suppressing herbivore populations should be considered.

In the recent SI of BioControl and resultant book from this working group (Roy et al., 2012), we contributed a review paper on the chemical ecology of the invasive aphidophagous ladybird Harmonia axyridis (Sloggett et al., 2011). This paper focused on both the pure and applied aspects of this subject, including sections on: (1) chemical defence; (2) foods, feeding and reproduction; (3) H. axyridis chemistry, humans and human activity, and (4) future research perspectives.Harmonia axyridis' chemical defence is comprised of two alkaloids (harmonine and 3-hydroxypiperidin-2-one) and four methoxypyrazines. Based on assays of palatability and toxicity, it appears to be one of the most powerful chemical defence blends amongst ladybirds. There is some evidence that both harmonine and methoxypyrazine concentrations vary predictably with ladybird colour and pattern, although no such work has been carried out for 3-methoxypiperidin-2-one. Since our original review was published, the alkaloid harmonine has been shown to be a broad spectrum anti-microbial and powerful anti-malarial compound (Rörich et al., 2012).Our knowledge of food location in H. axyridis remains poor, although the ladybird appears to use multimodal cues, probably including chemical ones from both host plant and aphid prey. Like many other ladybirds, H. axyridis is deterred from oviposition by conspecific larval chemical cues. It is not deterred from oviposition by chemical cues from allospecifics, although in its native range other ladybird species avoid such cues from H. axyridis. This behaviour probably serves these species as a means of avoiding H. axyridis intraguild predation. Generally, H. axyridis ignores the deterrent hydrocarbon cues on the surface of ladybird egg intraguild prey and is little affected by intraguild prey defensive alkaloids, although a number of exceptions are known, especially in its invasive range.The habit of H. axyridis overwintering in large aggregations in buildings is not limited to its invasive range, although in its native range, such aggregations tend to be of very limited duration. Aggregations in buildings cause numerous problems, including the spotting of furnishings from reflex blood, allergic reactions in humans using the buildings and risks posed to pets or even children that may eat the ladybirds. Another problem caused by the abundance of H. axyridis is "wine taint", predominantly from methoxypyrazines, when the ladybirds are attracted to grape crops in autumn and are harvested and processed with the

The primary peer-reviewed literature pertaining to rearing of predatory beetles, true bugs, and lacewings was reviewed and synthesized. This study focused on the literature published from 1998 to 2007. Advances in rearing were revealed in relation to the influence of factitious prey and artificial diets on predator life parameters. Eggs of the Angoumois grain moth and Mediterranean flour moth were factitious prey that supported the development and reproduction of many species in lieu of natural prey. Artificial diets based on vertebrate protein were generally inferior to factitious prey for production of predators. A few exceptional cases demonstrated potential for continuous rearing of true bugs and lacewings on artificial diet, as a stand-alone food source. Factors that could ensure progressive rearing success might involve (1) exploiting the highly polyphagous nature of certain species, (2) exposing predators to artificial diet for multiple generations to incite adaptation, (3) formulating artificial diets to match the texture and chemical composition of preferred natural prey, and (4) encasing the diet within Parafilm Ò to resemble the shape of natural prey. Encasement may also reduce desiccation and retard spoilage of the diet. Identification of other factitious prey and development of artificial diets that ensure production of predators of high quality should be possible. This review was conducted, for the most part, in support of generalist insect predators used, or having the potential to be used, in augmentative biological control of pests on plants in semi-closed systems, such as greenhouses and interiorscapes.