Antennal sensory organs and glands of the harlequin ladybird,Harmonia axyridis

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is a sub‐cosmopolitan species. Native to Asia, it has been released during the 20th century for classical and augmentative biological control of several herbivorous insects, mostly aphids and coccids. Despite its recognized positive impact on biological control, H. axyridis is now considered among the most dangerous invasive species in Europe and in most places where it has established. This is mostly due to its ability to reduce the populations of native predatory species of the same trophic guild. When exploring a new area, H. axyridis adults use semiochemical cues to acquire information about the habitat. Presumably, these cues are perceived by the sensilla located on the antennae. Surprisingly, in spite of the huge literature existing on H. axyridis, the antennal sensory organs have been poorly characterized. Here, we used scanning and transmission electron microscopy (SEM, TEM) techniques to study H. axyridis antennae, with focus on the various types of sensilla and their distribution in male and female individuals. The presence of various classes of antennal sensilla belonging to the main types described in insects (chemoreceptors, mechanoreceptors, and thermo‐hygroreceptors) was highlighted, as well as the widespread presence of antennal glands. The investigations showed some peculiar characteristics not known in Coccinellidae, such as the concentration of sensory structures at the level of the distal part of the apical antennomere and the discovery of antennal glands associated with it. No sexual dimorphism was revealed, neither for the general structure of the antenna (similar number of antennomeres and presence of modifications), nor for the total length and width of the antenna, the relative size of the antennomeres, the types of antennal sensilla, of their distribution and abundance. The potential relevance of these sensory structures and antennal glands, reported for the first time in Coccinellidae, is discussed in the context of intra‐ and interspecific communication.     

Ecology and biological control application of multicoloured Asian ladybird, Harmonia axyridis: A review

The ecology of and biological control by multicoloured Asian ladybird beetle, Harmonia axyridis (Pallas) are reviewed. Our emphasis is on assembling and interpreting information on general characteristics, invasion and establishment, sexual activity, foraging and predation, development, survival and reproduction, predator-predator interactions, natural enemies, biocontrol, non-target effects and status of H. axyridis as a pest of fruits. Colonization of H. axyridis for aphid biocontrol in the USA have been successful in terms of its establishment, but its abundance is turning out to be a nuisance to humans. Its harmful non-target impact on beneficial organisms, humans and native species is becoming a debatable issue. The question on its present position, whether it is a biocontrol agent or pest, is a critical issue and discussed. Inferences from the empirical data are made and new avenues for future research are suggested.     

Factitious prey and artificial diets for predatory lady beetles: current situation,obstacles, and approaches for improvement: a review

Predatory lady beetles (Coleoptera: Coccinellidae) are important natural enemies of many pests in crop ecosystems throughout the world. Although several species are currently mass-reared and sold by biocontrol companies, there is an urgent need to reduce rearing costs. Cost-effective mass rearing of lady beetles involves the use of factitious prey/foods or artificial diets in lieu of natural prey. This review summarises the literature on the various kinds of factitious prey/foods and artificial diets as well as the feeding efficiency of predatory lady beetles. Factitious prey/foods generally satisfy the nutritional requirements for development of lady beetles, and lepidopteran immatures are the most widely used species. However, artificial diets, based on vertebrate protein and devoid of arthropod constituents, often yield disappointing results. We also highlight several successful examples and novel approaches for improving feeding efficiency. Moreover, available approaches and perspectives for speeding up the development and screening of cost-effective diets for mass rearing are stated in this review.     

Influence d'une alimentation préalable et du jeûne sur l'apparition de la recherche intensive des proies chez Semiadalia undecimnotata

Résumé Les prédateurs qui vivent aux dépens de ressources alimentaires groupées, modifient leur mode de déplacement après la capture et l'ingestion d'une proie. Ils passent de la recherche extensive (déplacements rapides et linéaires) à la recherche intensive (déplacements lents et sinueux). Chez les adultes et les larves de dernier stade de la coccinelle Semiadalia undecimnotata Schn. (Col., Coccinellidae), dans les premières heures qui suivent leur naissance ou leur mue, l'ingestion d'un premier puceron ne modifie pas les caractéristiques locomotrices individuelles: les déplacements restent de type extensif. Au début de chaque stade, cette coccinelle présente une période de sensibilisation à la proie rencontrée. Durant ces laps de temps, elle doit consommer plusieurs proies avant d'être capable d'adopter la recherche intensive. L'existence d'une période de jeûne avant cette première prise alimentaire favorise, au contraire, l'adoption de la recherche intensive.

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The ladybird, Semiadalia undecimnotata, like all other entomophagous insects feeding on relatively sedentary prey, exhibits two types of walking pattern during the search for prey: xtensive and intensive search. The latter is engaged following detection of visual or chemical cues. Changes in the ladybird search pattern: extensive search—a single prey capture—intensive search, were investigated in relation to experience of prior feeding-periods (experienced coccinellids) or to lack of such experience (naïve coccinellids) and with respect to changes in duration of fasting. The analysis of their pathways was performed initially by comparison of three locomotory paramètres: the number of stops (number/s), the walking speed (mm/s) and the turning-rate (degrees/s), and subsequently by statistical classification (principle components analysis). Unlike experienced coccinellids, observations of naïve coccinellids, indicate that a single feeding session during the three to five hours after emergence of an adult, or ecdysis of the larva, will change the walking pattern slightly. These animals will maintain extensive search or adopt a particular, intermediate locomotory pattern which is a development of extensive search but do not adopt intensive search. Fasting, for periods over twelve hours, favoured intensive search in populations studied. It is probable that the requirement to feed on more than one prey item before adopting intensive search involves concepts such as forms of learning (sensitization).

     

Effects ofBacillus thuringiensis var.San diego on predation effectiveness,development and mortality ofColeomegilla maculata lengi (Col.: Coccinellidae) larvae

Laboratory experiments were conducted to determine the effects of M-One^™ (Bacillus thuringiensis var.san diego) on larval instars ofColeomegilla maculata lengi Timberlake. Coccinellid larval development (from egg hatch to adult), completed on pollen treated with suspensions of M-One^™ at 20 ml/litre (5.6×10⁸ CPBIU/litre) and 200 ml/litre, took respectively 29.3 and 38.5 days compared with 21.9 days for the control (water). M-One^™ did not cause larval mortality.C. maculata third instars did not show any preference between eggs ofLeptinotarsa decemlineata (Say) treated with water or with M-One^™ at 20 ml/litre. However, at 200 ml M-One^™/litre, the number of eggs attacked was 34.7% lower than the eggs treated with water only, 48 h after the beginning of the test. These results indicate that the use of M-One^™, at the manufacturer's recommended field rate of 20 ml/litre, does not cause a major threat to larvalC. maculata populations.

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Résumé Des bioessais en laboratoire ont été effectués afin de déterminer si les larves de la coccinelle maculée,Coleomegilla maculata lengi Timberlake (Col.: Coccinellidae), sont affectées par M-One^™, un insecticide biologique préparé à partir de la bactérieBacillus thuringiensis var.san diego Berliner et utilisé dans la lutte contre le doryphore de la pomme de terre,Leptinotarsa decemlineata (Say) (Col.: Chrysomelidae). Le développement larvaire, effectué sur du pollen traité avec des concentrations de 20 ml M-One^™/litre (5,6×10⁸ unités internationales de doryphore/litre) et 200 ml M-One^™/litre, a nécessité 29,3 et 38,5 jours respectivement, comparativement à 21,9 jours pour le témoin (eau). M-One^™ n'a pas causé de mortalité chez les larves. Au cours d'un test de 48 h, les larves de stade III n'ont montré aucune préférence entre des œufs traités avec 20 ml M-One^™/litre et des œufs traités avec de l'eau. Par contre avec 200 ml M-One^™/litre, le nombre d'œufs attaqués a diminué significativement de 34,7% par rapport au témoin, 48 h après le début du test. Ces résultats indiquent que l'utilisation de M-One^™ à la concentration recommandée de 20 ml/litre ne représente pas une menace pour les populations larvaires de la coccinelle maculée.

     

Differential acceptance of and survivorship on a non‐host plant between two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii

Phytophagous specialists among insects occasionally accept plants they do not or only rarely utilize under natural conditions. This non‐host acceptance could represent an initial stage in host‐range evolution. Here, we examined adult acceptance of and larval survivorship on a non‐host plant – wild thistle, Cirsium kamtschaticum Ledeb. (Asteraceae) – for two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii Katakura (Coleoptera: Coccinellidae), which normally utilizes blue cohosh, Caulophyllum robustum Maxim. (Berberidaceae), in the wild. The two beetle populations showed lower acceptance of and survivorship on the non‐host plant (thistle) than on the normal host plant (blue cohosh). Furthermore, they differed significantly in their responses to thistle but not to blue cohosh. Even for the beetle population that more readily accepted thistle as adults, acceptance was moderate and the eclosion rate was low, indicating reduced performance on the non‐host plant. Although studies have infrequently focused on non‐host acceptance, this is likely common in phytophagous specialists and thus is a potentially important evolutionary factor, as it may determine the future direction of food‐range evolution.     

Why do neonates of aphidophagous ladybird beetles preferentially consume conspecific eggs in presence of aphids?

The neonates of aphidophagous ladybird beetles, Propylea dissecta, Coccinella transversalis and Coelophora saucia preferentially consumed conspecific eggs in presence of essential aphid prey, Aphis gossypii. This preference was strongest in P. dissecta and recorded in all density combinations of conspecific eggs and aphids. The neonates of all three species developed faster resulting in heavier second instars with lower mortality when fed on conspecific eggs as compared to mobile or defenceless aphids. In addition, they required less dry biomass of conspecific eggs than that of aphids. These results reveal that neonates are benefitted intrinsically when they feed on conspecific eggs. The neonates of P. dissecta exhibited discrimination for more nutritious food, as they contacted and ate conspecific eggs more frequently than aphids as their first meal. This discrimination was not found in Coccinella transversalis and Coelophora saucia possibly due to their bigger sizes and possible higher energy requirements. Although, the major effect of chemical cues was only noticed in P. dissecta, it could be concluded that chemicals (surface and within) of conspecific eggs possibly attract neonates for egg-cannibalism, as both eggs and aphids were consumed in greater numbers when coated with egg extracts. The reverse occurred when eggs and aphids were coated with aphid extracts.     

Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China

Abstract 1. We observed native populations of Harmonia axyridis (Pallas) around Beijing, China, over 2 years and performed choice and no‐choice mating tests between melanic and succinic (non‐melanic) beetles in the laboratory. 2. Succinic phenotypes outnumbered melanics by 5:1 in autumn, but melanics became equally abundant in spring, supporting previous inferences that melanism is advantageous in winter, but costly in summer. 3. Female H. axyridis expressed mate preference overtly, by rejecting less‐preferred phenotypes, and cryptically, by retaining their eggs for longer periods after matings with less‐preferred males, ostensibly to replace their sperm. 4. Succinic pairs formed more quickly in the spring generation, and melanic pairs in the autumn, and the time to copula was affected by both male and female phenotype. The strength of mate preference was contingent on female phenotype, suggesting melanic alleles had pleiotropic effects. 5. Whereas pair formation was under female control, the duration of copula was under male control and lasted longer in the autumn generation than in the spring. Copulations in the choice test tended to be shorter between similar phenotypes, suggesting that males invested more in dissimilar females when alternative mates were available. 6. Although spring and autumn generations were raised under identical conditions, significant contrasts were observed in their reproductive behaviour. 7. Two alternative hypotheses are advanced to explain why gender‐specific reproductive behaviours might vary between generations: maternally‐mediated epigenetic factors that influence the expression of genes in progeny as a function of maternal environment, and linkage disequilibria among alleles that cycle in frequency seasonally as a function of assortative mating.     

Landscape composition influences patterns of native and exotic lady beetle abundance

Aim Coccinellid beetles are important predators that contribute to pest suppression in agricultural landscapes. Since the introduction of the exotic coccinellids Coccinella septempunctata L. and Harmonia axyridis Pallas into the USA, several studies have reported a decline of native Coccinellidae in agroecosystems. We aimed to investigate the influence of landscape composition on native and exotic coccinellid abundance within soybean fields. Location Iowa, Michigan, Minnesota and Wisconsin. Methods As part of a 2‐year study (2005–06) on the biological control of the soybean aphid, Aphis glycines Matsumura, we examined coccinellid communities in 33 soybean fields using yellow sticky card traps. Landscape heterogeneity and composition were measured at multiple spatial scales ranging 1–3.5 km from focal soybean fields where coccinellid sampling took place. Results Exotic species made up 90% of the total coccinellid community in Michigan soybean fields followed by Wisconsin (84%), Minnesota (66%) and Iowa (57%). Harmonia axyridis was the dominant exotic coccinellid in all states comprising 45–62% of the total coccinellid community, followed by C. septempunctata (13–30%). Two additional exotic species, Hippodamia variegata (Goeze) and Propylea quatuordecimpunctata (L.) were also found in the region. Overall, the most abundant native coccinellid was Hippodamia convergens Guerin‐Meneville; however, its abundance varied across the region, comprising 0% (Michigan) to 28% (Iowa) of the total coccinellid community. Landscape structure significantly influenced the composition of coccinellid communities in soybean agroecosystems. We found that native coccinellids were most abundant in low‐diversity landscapes with an abundance of grassland habitat while exotic coccinellids were associated with the abundance of forested habitats. Main conclusion We propose that grassland dominated landscapes with low structural diversity and low amounts of forested habitat may be resistant to exotic coccinellid build‐up, particularly H. axyridis and therefore represent landscape‐scale refuges for native coccinellid biodiversity.     

Interactions between ants and aphidophagous and coccidophagous ladybirds

Aphidophagous and coccidophagous coccinellids come into conflict with homopteran-tending ants for access to food. Antagonistic interactions between coccinellids and ants may be competitive or non-competitive. Competitive interactions occur when coccinellids attack aphids or coccids that are being tended by ants for honeydew. Non-competitive interactions include all interactions away from ant-tended homopteran colonies. We here review observations and studies of such interactions. We note that most competitive interactions occur at times when untended aphids/coccids are scarce. We describe the chemical and physical defences that coccinellids use against ant aggression and consider whether these have evolved as general anti-predator deterrents or specifically in response to ants. Myrmecophilous coccinellids are then considered, with particular focus on the two most studied species, Coccinella magnifica and Platynaspis luteorubra. We note that the myrmecophily of the two species has the same adaptive rationale—to enable the ladybirds to prey on ant-tended aphids at times of aphid scarcity—but that it is based on different traits to facilitate life with ants. Finally, we consider the role of ants in the evolution of habitat specialisation in some coccinellids.