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Concept: Biological pest control agents


Harmonia axyridis has been introduced as a biological control agent in Europe and the USA. Since its introduction, it has established and spread, and it is now regarded as an invasive alien species. It has been suggested that intraguild predation is especially important for the invasion success of H. axyridis. The aim of this study was to compare the intraguild predation behaviour of three ladybird species (Coccinella septempunctata, Adalia bipunctata, and H. axyridis). Predation behaviour was investigated in semi-field experiments on small lime trees (Tilia platyphyllos). Two fourth-instar larvae placed on a tree rarely made contact during 3-hour observations. When placed together on a single leaf in 23%-43% of the observations at least one contact was made. Of those contacts 0%-27% resulted in an attack. Harmonia axyridis attacked mostly heterospecifics, while A. bipunctata and C. septempunctata attacked heterospecifics as often as conspecifics. In comparison with A. bipunctata and C. septempunctata, H. axyridis was the most successful intraguild predator as it won 86% and 44% of heterospecific battles against A. bipunctata and C. septempunctata respectively, whilst A. bipunctata won none of the heterospecific battles and C. septempunctata won only the heterospecific battles against A. bipunctata. Coccinella septempunctata dropped from a leaf earlier and more often than the other two species but was in some cases able to return to the tree, especially under cloudy conditions. The frequency with which a species dropped did not depend on the species the larva was paired with. The results of these semi-field experiments confirm that H. axyridis is a strong intraguild predator as a consequence of its aggressiveness and good defence against predation from heterospecific species. The fact that H. axyridis is such a strong intraguild predator helps to explain its successful establishment as invasive alien species in Europe and the USA.

Concepts: Biological pest control, Invasive species, Introduced species, Coccinellidae, Harmonia axyridis, Tilia, Coccinella septempunctata, Biological pest control agents


The harlequin ladybird, Harmonia axyridis, has emerged as a model species for invasion biology, reflecting its remarkable capacity to outcompete native ladybird species when introduced into new habitats. This ability may be associated with its prominent resistance to pathogens and intraguild predation. We recently showed that the constitutive antibacterial activity present in the haemolymph of H. axyridis beetles can be attributed to the chemical defence compound harmonine. Here, we demonstrate that H. axyridis differs from other insects, including the native ladybird Coccinella septempunctata, by reducing rather than increasing the antimicrobial activity of its haemolymph following the injection of bacteria. However, both species produce new or more abundant proteins in the haemolymph, indicating that bacterial challenge induces innate immune responses associated with the synthesis of immunity-related proteins. Our results suggest that H. axyridis beetles can switch from constitutive chemical defence to inducible innate immune responses, supporting hypothesis that inducible antimicrobial peptides protect host beetles against pathogens that survive constitutive defences. These alternative antimicrobial defence mechanisms may reflect a trade-off resulting from fitness-related costs associated with the simultaneous synthesis of harmonine and antimicrobial peptides/proteins.

Concepts: Immune system, Bacteria, Microbiology, Innate immune system, Coccinellidae, Harmonia axyridis, Coccinella septempunctata, Biological pest control agents


Invasive species that proliferate after colonizing new habitats have a negative environmental and economic impact. The reason why some species become successful invaders, whereas others, even closely related species, remain noninvasive is often unclear. The harlequin ladybird Harmonia axyridis, introduced for biological pest control, has become an invader that is outcompeting indigenous ladybird species in many countries. Here, we show that Harmonia carries abundant spores of obligate parasitic microsporidia closely related to Nosema thompsoni. These microsporidia, while not harming the carrier Harmonia, are lethal pathogens for the native ladybird Coccinella septempunctata. We propose that intraguild predation, representing a major selective force among competing ladybird species, causes the infection and ultimate death of native ladybirds when they feed on microsporidia-contaminated Harmonia eggs or larvae.

Concepts: Ecology, Invasive species, Introduced species, Coccinellidae, Harmonia axyridis, Coccinella septempunctata, Biological pest control agents, Coccinella


Ladybug taint (LBT) is a wine fault caused by the inadvertent incorporation of ladybeetles to the wine during the winemaking process. Harmonia axyridis, also known as the multicolored Asian lady-beetle (MALB), was the only species considered responsible for causing the taint. A second species, Coccinella septempunctata (7 Spot), has recently also been implicated. The main objectives of this study were to identify and quantify the compounds associated with ladybug taint of white and red wine from these two Coccinellidae species (Harmonia axyridis and Coccinella septempunctata), and determine the most odor-active compounds in LBT-affected wines.

Concepts: Wine, Oenology, Acids in wine, Coccinellidae, Harmonia axyridis, Coccinella septempunctata, Biological pest control agents, Coccinella


Numerous species of gall midges (Diptera: Cecidomyiidae) have been recorded from saltbush (Chenopodiaceae: Atriplex) around the world but only 11 of them belong to the large cecidomyiid genus Asphondylia. Of these, two species were described in the late 19th century from complex bud galls on Atriplex halimus in the Mediterranean Basin. In the present study Asphondylia punica is redescribed, A. conglomerata is synonymized with it, and Asphondylia scopuli is described from Atriplex lanfrancoi, an endemic plant to the Maltese Islands. Descriptions are accompanied by information about the galls and life history of the gall midges, and a review of the parasitic Hymenoptera associated with A. scopuli is provided. Four species of parasitoids were found and attributed to the families Eurytomidae, Pteromalidae, Eupelmidae and Eulophidae, of which the pteromalid Mesopolobus melitensis is described as new.

Concepts: Nematocera, Species, Flies, Parasitoid, Apocrita, Biological pest control agents, Cecidomyiidae, Atriplex


Understanding the characteristics and drivers of dispersal is crucial for predicting population dynamics, particularly in range-shifting species. Studying long-distance dispersal in insects is challenging, but recent advances in entomological radar offer unique insights. We analysed 10 years of radar data collected at Rothamsted Research, U.K., to investigate characteristics (altitude, speed, seasonal and annual trends) and drivers (aphid abundance, air temperature, wind speed and rainfall) of high-altitude flight of the two most abundant U.K. ladybird species (native Coccinella septempunctata and invasive Harmonia axyridis). These species cannot be distinguished in the radar data since their reflectivity signals overlap, and they were therefore analysed together. However, their signals do not overlap with other, abundant insects so we are confident they constitute the overwhelming majority of the analysed data. The target species were detected up to ∼1100 m above ground level, where displacement speeds of up to ∼60 km/h were recorded, however most ladybirds were found between ∼150 and 500 m, and had a mean displacement of 30 km/h. Average flight time was estimated, using tethered flight experiments, to be 36.5 minutes, but flights of up to two hours were observed. Ladybirds are therefore potentially able to travel 18 km in a “typical” high-altitude flight, but up to 120 km if flying at higher altitudes, indicating a high capacity for long-distance dispersal. There were strong seasonal trends in ladybird abundance, with peaks corresponding to the highest temperatures of mid-summer, and warm air temperature was the key driver of ladybird flight. Climatic warming may therefore increase the potential for long-distance dispersal in these species. Low aphid abundance was a second significant factor, highlighting the important role of aphid population dynamics in ladybird dispersal. This research illustrates the utility of radar for studying high-altitude insect flight and has important implications for predicting long-distance dispersal.

Concepts: Insect, Temperature, Aphid, Coccinellidae, Harmonia axyridis, Coccinella septempunctata, Biological pest control agents, Coccinella


Haller’s rule states that brains scale allometrically with body size in all animals, meaning that relative brain size increases with decreasing body size. This rule applies both on inter- and intraspecific comparisons. Only 1 species, the extremely small parasitic wasp Trichogramma evanescens, is known as an exception and shows an isometric brain-body size relation in an intraspecific comparison between differently sized individuals. Here, we investigated if such an isometric brain-body size relationship also occurs in an intraspecific comparison with a slightly larger parasitic wasp, Nasonia vitripennis, a species that may vary 10-fold in body weight upon differences in levels of scramble competition during larval development. We show that Nasonia exhibits diphasic brain-body size scaling: larger wasps scale allometrically, following Haller’s rule, whereas the smallest wasps show isometric scaling. Brains of smaller wasps are, therefore, smaller than expected and we hypothesized that this may lead to adaptations in brain architecture. Volumetric analysis of neuropil composition revealed that wasps of different sizes differed in relative volume of multiple neuropils. The optic lobes and mushroom bodies in particular were smaller in the smallest wasps. Furthermore, smaller brains had a relatively smaller total neuropil volume and larger cellular rind than large brains. These changes in relative brain size and brain architecture suggest that the energetic constraints on brain tissue outweigh specific cognitive requirements in small Nasonia wasps.

Concepts: Neuroanatomy, Brain, Density, Larva, Human brain, Wasp, Brain size, Biological pest control agents


Integrated Pest Management (IPM) is a decision making process used to manage pests that relies on many tactics, including cultural and biological control, which are practices that conserve beneficial insects and mites, and when needed, the use of conventional insecticides. However, systemic, soil-applied neonicotinoid insecticides are translocated to pollen and nectar of flowers, often for months, and may reduce survival of flower-feeding beneficial insects. Imidacloprid seed-treated crops (0.05 mg AI (active ingredient) /canola seed and 1.2 mg AI/corn seed) translocate less than 10 ppb to pollen and nectar. However, higher rates of soil-applied imidacloprid are used in nurseries and urban landscapes, such as 300 mg AI/10 L (3 gallon) pot and 69 g AI applied to the soil under a 61 (24 in) cm diam. tree. Translocation of imidacloprid from soil (300 mg AI) to flowers of Asclepias curassavica resulted in 6,030 ppb in 1X and 10,400 ppb in 2X treatments, which are similar to imidacloprid residues found in another plant species we studied. A second imidacloprid soil application 7 months later resulted in 21,000 ppb in 1X and 45,000 ppb in 2X treatments. Consequently, greenhouse/nursery use of imidacloprid applied to flowering plants can result in 793 to 1,368 times higher concentration compared to an imidacloprid seed treatment (7.6 ppb pollen in seed- treated canola), where most research has focused. These higher imidacloprid levels caused significant mortality in both 1X and 2X treatments in 3 lady beetle species, Coleomegilla maculata, Harmonia axyridis, and Hippodamia convergens, but not a fourth species, Coccinella septempunctata. Adult survival were not reduced for monarch, Danaus plexippus and painted lady, Vanessa cardui, butterflies, but larval survival was significantly reduced. The use of the neonicotinoid imidacloprid at greenhouse/nursery rates reduced survival of beneficial insects feeding on pollen and nectar and is incompatible with the principles of IPM.

Concepts: Insect, Beetle, Flower, Lepidoptera, Coccinellidae, Coccinella septempunctata, Biological pest control agents, Hippodamia convergens


The use of biological control agents to control pests is an alternative to pesticides and a tool to manage invasive alien species. However, biocontrol agents can themselves become invasive species under certain conditions. The harlequin ladybird (Harmonia axyridis) is a native Asian biocontrol agent that has become a successful invader. We reviewed articles containing “Harmonia axyridis” to gather information on its presence and surveyed entomologists researching Coccinellidae around the world to investigate further insights about the current distribution, vectors of introduction, habitat use and threats this species pose. The harlequin ladybird has established populations in at least 59 countries outside its native range. Twenty six percent of the surveyed scientists considered it a potential threat to native Coccinellidae. Published studies and scientists suggest Adalia bipunctata, native to Europe, is under the highest risk of population declines. Strict policies should be incorporated to prevent its arrival to non-invaded areas and to prevent further expansion range. Managing invasive species is a key priority to prevent biodiversity loss and promote ecosystem services.

Concepts: Biodiversity, Biology, Ecology, Biological pest control, Invasive species, Coccinellidae, Harmonia axyridis, Biological pest control agents


For an insect to be able to efficiently attach to surfaces, the adhesive pads on the distal parts of its legs must establish large contact areas. In case of hairy adhesive pads this requires flexibility of the contact-forming bristles, called adhesive tarsal setae. However, too flexible setae would have a low mechanical stability resulting in a decreased attachment ability of the pads. Here we show that the adhesive tarsal setae of the ladybird beetle Coccinella septempunctata feature pronounced gradients in the material composition and properties along their length. The Young’s modulus ranges from 1.2 MPa at the tips, where we found the incorporation of high proportions of the elastic protein resilin, to 6.8 GPa at the bases of the setae. These gradients likely represent an evolutionary optimization, which increases the performance of the adhesive system by enabling effective adaptation to rough surfaces while simultaneously preventing lateral collapse of the setae.

Concepts: Beetle, Pascal, Young's modulus, Hair, Coccinellidae, Coccinella septempunctata, Biological pest control agents, Coccinella