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Many aphidophagous ladybird beetles lay clusters of eggs and sibling cannibalism occurs at hatching. Larvae that hatch early tend to cannibalize undeveloped eggs and those that hatch late. The cannibalized eggs, especially those that are sterile, represent a maternal investment in the cannibal and are regarded as "trophic" eggs, which increase their chances of surviving. The characteristics of cannibalized eggs, however, are poorly studied. In this study, I determined the sizes of eggs that developing and undeveloped eggs within egg clusters of Harmonia axyridis (Coleoptera: Coccinellidae). Developing eggs were significantly larger (on average 3.45% larger) than the undeveloped sterile ones, which were mainly small. Furthermore, the hatchability of eggs was also significantly associated with their size, inclusive of the maternal effects. These results indicate that the production of relatively small sterile eggs functions as circumstance-dependent maternal investment in improving the survival of the first instar larvae to hatch in each egg cluster.

Abstract In this study, we focused on the direct (i.e., predation) and indirect (i.e., potential threat from coexisting with a larger individual) effects of cannibalism and intraguild predation (IGP) during larval stages of two sibling ladybird beetles. These effects play an important role in the coexistence of the generalist–common Harmonia axyridis and specialist–rare H. yedoensis (Coleoptera: Coccinellidae). Direct predation effect of cannibalism and IGP was asymmetric in the two sibling ladybird beetles; the fourth instar larvae of H. axyridis were better intraguild predators than cannibals, while the reverse was true in the larvae of H. yedoensis. Neither cannibalism nor IGP significantly affected female body weight in either species. Larval H. axyridis surviving exposure to cannibalism or IGP had a reduced number of ovarioles as adults, whereas adult H. yedoensis ovarioles were not affected. For the indirect effects, longer developmental times in males and females and a lower total number of ovarioles in females were detected in H. axyridis. In H. yedoensis, shorter developmental time of males, lighter adult weight and fewer total ovarioles in females were observed. Olfactometer choice experiments clarified that the fourth instar larvae of H. axyridis avoided the first instar conspecific larvae, while those of H. yedoensis were attracted to the odors from H. axyridis and conspecifics. Thus, H. axyridis has an avoidance mechanism only for cannibalism but not for IGP, whereas H. yedoensis does not have any avoidance mechanism. These different behaviors in the direct and indirect effects of cannibalism and IGP observed in the laboratory may play important roles in the coexistence of generalist–common H. axyridis and specialist–rare H. yedoensis in natural conditions, compensating for the large handicap of H. yedoensis at reproductive interference from H. axyridis.

To develop nanosensors to probe neurotransmitters, we synthesized fluorescent-functionalized molecularly imprinted polymeric nanoparticles (fMIP-NPs) using monoamine neurotransmitters (serotonin and dopamine) immobilized on glass beads as templates. The size and fluorescence intensity of the fMIP-NPs synthesized with blended silane couplers increased with the presence of the target but were insensitive to the target analogs (L-tryptophan and L-dopa, respectively). However, when the template is anchored by a pure silane agent, both the fluorescence intensity and particle size of the fMIP-NPs were sensitive to the structural analog of the template. Another fMIP-NP was synthesized in the presence of poly([2-(methacryloyloxy)ethyl] trimethylammonium chloride (METMAC)-co-methacrylamide) grafted onto glass beads as a dummy template for acetylcholine. Acetylcholine increased the diameter and fluorescence intensity of the fMIP-NP, but choline had no effect. When the homopolymer of METMAC was used as a template, the fluorescence intensity and size of the resulting nanoparticles were not responsive to either acetylcholine or choline. The principle of increased fluorescence intensity due to specific interaction with the target substance is probably due to the increased distance between the fluorescent functional groups and decreased self-quenching due to the swelling caused by the specific interaction with the template. The results also indicate that MIP nanoparticles prepared by solid-phase synthesis can be used for targeting small molecules, such as the neurotransmitters addressed in this study, by adjusting the surface density of the template.

While insect predators need high-quality food to reach sexual maturity and reproduce, starvation following adult eclosion may occur when prey are missing in agroecosystems. This study explores whether the type of diet available at eclosion determines the future fecundity of newly emerged adult predators. In a laboratory experiment, three different diets (i.e., flowers of Perilla frutescens (L.) Britton, eggs of Ephestia kuehniella Zeller as prey, or no food) were offered to adult females of the ladybird beetle Harmonia axyridis Pallas during their first three days after adult eclosion. On the fourth day, each female was paired with a prey-fed male and the pair was subsequently fed with prey. Diet at eclosion did not affect pre-oviposition time, the number of eggs oviposited daily, or the viability of egg batches. High variability in pre-oviposition time among females was observed for all diets. Significant negative linear relationships were found between pre-oviposition time and both the number of eggs oviposited daily and the viability of egg batches. This study clarifies that the food readily available at adult eclosion does not affect the capacity of H. axyridis to reproduce, provided that adults find prey within a few days. More generally, it shows that the reproductive traits of H. axyridis allow this generalist predator to be highly adapted to heterogenous environments in its native range. It is concluded that the variety of habitats offered by diversified agroecosystems may highly benefit the ladybird beetle H. axyridis, and potentially enhance its ability to biologically regulate crop pests.

In predatory ladybirds male embryos in clutches of eggs infected with male-killing bacterial endosymbionts do not hatch and are consumed by female hatchlings. Moreover, it is predicted that infection with male-killing bacteria should be prevalent in a ladybird population if the fitness advantage to female ladybirds due to the resulting reallocation of resources is high. We compared male-killer bacterial prevalence in two sibling species of the ladybird genus Harmonia that use different host ranges. Harmonia yedoensis feeds mostly on the highly elusive pine aphid and its hatchlings can greatly enhance their ability to capture prey and thus their survival by consuming un-hatched eggs in the clutch. In contrast, Harmonia axyridis feeds on a wide range of prey and consumption of un-hatched eggs by the hatchlings does not necessarily increase their foraging success. In the study area where these two species of ladybird occur sympatrically 14 of the 22 females of H. yedoensis (64%) laid clutches of eggs of which approximately only a half hatched and did not produce male offspring, whereas only one of 12 of the females of H. axyridis (8%) exhibited symptoms indicating infection with male-killing bacteria. In the H. yedoensis that exhibited symptoms of being infected with male-killing bacteria, the total allocation of resources to each hatchling, which was provided by laying larger eggs and eating the un-hatched eggs in the clutch, was considerable. These results suggest that the difference in the prevalence of infection with male-killing bacteria in these two species of ladybird may reflect differences in the quality of their prey and their foraging behaviour.

The range and quality of prey species differ greatly among closely related species of predators. However, the factors responsible for this diversified niche utilization are unclear. This is because the predation and resource competition do not always prevent species coexistence. In this paper, we present evidence in support of reproductive interference as a driver of niche partitioning, focusing on aphidophagous insect. Firstly, we present closely related generalist and specialist species pairs in aphidophagous lacewings to compare the reproductive interference hypothesis with two other hypotheses that have been proposed to explain niche partitioning in lacewings and sympatric speciation through host race formation and sexual selection. Secondly, we present a case study that shows how reproductive interference can drive niche partitioning in sibling ladybird species. Thirdly, we show that many ladybird genera include species inhabiting the same region but having different food and habitat preferences, raising the possibility that reproductive interference might occur in these groups. Finally, we show that intraguild predation cannot always explain the niche partitioning in aphidophagous insects including hoverflies and parasitoids. On the basis of the evidence presented, we urge that future studies investigating predator communities should take account of the role of reproductive interference.

In Hymenoptera and Heteroptera, the absence of micropyles is one criterion for categorizing an egg as trophic. Undeveloped eggs are observed in more than 90% of the egg clusters of the ladybird beetle Harmonia axyridis Pallas. Traditionally, these undeveloped eggs are regarded as "trophic eggs." The surfaces of the eggs of H. axyridis were examined using scanning electron microscopy and the presence of micropyles in the shells of developing and undeveloped eggs determined. Micropyles are circularly distributed around the top of eggs and present in both developing and undeveloped eggs. The number of micropyles in the shells of developing and undeveloped eggs did not differ significantly. Our results indicate that the undeveloped eggs of H. axyridis have micropyles, suggesting that the mechanisms regulating the production of undeveloped eggs in H. axyridis differ from those resulting in the production of trophic eggs by Hymenoptera and Heteroptera.

The sibling species H. yedoensis Takizawa coexists sympatrically and simultaneously with H. axyridis only on pine trees in Japan. To elucidate the mechanisms enabling coexistence of these two sympatric sibling species, a laboratory experiment was performed that focused on differences in their maternal investment through eggs and the role of sibling cannibalism. The egg size (volume) of H. yedoensis was 24.91% larger than that of H. axyridis. Cluster size in H. axyridis was significantly larger than that in H. yedoensis; however, the total number of eggs and oviposition cost (by volume) per female in H. yedoensis were not significantly different from those in H. axyridis, although total number of clusters tended to be slightly higher in H. yedoensis than in H. axyridis. The percentage of undeveloped eggs per cluster in H. yedoensis was not significantly different from that in H. axyridis, whereas the percentage of developed eggs with delayed hatching per cluster was significantly larger in H. yedoensis than in H. axyridis. Moreover, the cost of sibling cannibalism per hatched larval cluster in H. yedoensis (worth 4.43 sibling eggs) was 3.36 times larger than that in H. axyridis.Therefore, maternal investment through egg and sibling cannibalism in developed eggs with delayed hatching are more intense in H. yedoensis than in H. axyridis, implying a higher larval survival rate through higher ability of prey capturing at the first instar. The results in this study suggest that the higher survival rate and accelerated development in H. yedoensis by the two maternal investments, i.e., a large egg and intense sibling cannibalism of developed eggs with delayed hatching, may play an important role in sympatric coexistence with the aggressive aphidophagous ladybird beetle H. axyridis.

The ladybird beetle Harmonia axyridis Pallas was investigated under laboratory conditions to clarify the relationship between food abundance or scarcity and ovarian development or oosorption. Four conditions were used: (1) fully fed for 24 h, (2) 24-h starvation, (3) 48-h starvation, and (4) 24-h starvation followed by 24-h re-feeding. Body length and initial body weight were not significantly related to the number of ovarioles per female. Both starvation conditions significantly increased the percentage of oosorptive individuals and ovarioles per female, and significantly decreased the percentage of mature ovarioles per female. Re-feeding for 24 h after a 24-h starvation resulted in a significantly higher percentage of mature ovarioles per female; however, the percentage of mature ovarioles remained lower than in the fully fed condition. Oosorption mainly occurred during the intermediate developmental stage of the ovarioles. The rates of ovarian development and oosorption in predatory H. axyridis were much faster compared with those in herbivorous ladybird beetles. Body length, initial body weight, and the number of ovarioles were significantly correlated with the number of eggs laid during the last 24 h of each experimental condition. From an analysis of the weight loss and the number of eggs laid during the last 24 h of each experimental condition, it appears that the realized weight of the eggs may be directly determined by the amount of food digested by the adult. The ovarian development and oosorption were asymmetric in the right and left ovaries. These may be important strategies for oviposition in H. axyridis, because selective provision of maturing ovarioles in the right or left ovary with digested nutrients would favor their development. In addition, the energy loss through oosorption during the intermediate developmental stage of oocytes would be less than the energy loss resulting from the resorption of mature oocytes. Therefore, one role of the ovary in H. axyridis, in addition to egg production, might be as a kind of energy storage system for increasing reproductive success. An immediate start of ovarian development under favorable feeding conditions and rapid oosorption during food scarcities may be an adaptive ovipositional and survival strategy for female adults of H. axyridis in response to heterogeneous and fluctuating resource conditions.

I hypothesized that sibling cannibalism is one of maternal investment in that a female controls sibling cannibalism. To test the hypothesis, I conducted a laboratory experiment and field observations to investigate sibling cannibalism in relation to cluster size and cluster site in the ladybird beetle, Harmonia axyridis Pallas. In the laboratory experiment, cluster size significantly affected the number of cannibalized eggs per cluster (R2 = 0.516), while cluster size was significantly affected by the oviposition interval. Furthermore, there was a marginally significant positive relationship between cluster size and the percentage of sibling cannibalism per cluster. In the field, cluster size and the direct distance from a cluster site to an aphid colony (an indicator of intensity of non-sibling cannibalism) significantly affected the number of cannibalized eggs per cluster (R2 = 0.472). Furthermore, there was a significant positive relationship between the direct distance from a cluster to the nearest aphid colony and cluster size. However, there was not a significant relationship between the distance and the percentage of sibling cannibalism. These results may be caused by the weakness of the female's power to control sibling cannibalism. Thus, a female H. axyridis controls cluster size through the intensity of non-sibling cannibalism, which may be one of oviposition strategies in this species.

Reproductive strategy affects population dynamics and genetic parameters that can, in turn, affect evolutionary processes during the course of biological invasion. Life-history traits associated with reproductive strategy are therefore potentially good candidates for rapid evolutionary shifts during invasions. In a series of mating trials, we examined mixed groups of four males from invasive and native populations of the harlequin ladybird Harmonia axyridis mating freely during 48 hours with one female of either type. We recorded the identity of the first male to copulate and after the 48 h-period, we examined female fecundity and share of paternity, using molecular markers. We found that invasive populations have a different profile of male and female reproductive output. Males from invasive populations are more likely to mate first and gain a higher proportion of offspring with both invasive and native females. Females from invasive populations reproduce sooner, lay more eggs, and have offspring sired by a larger number of fathers than females from native populations. We found no evidence of direct inbreeding avoidance behaviour in both invasive and native females. This study highlights the importance of investigating evolutionary changes in reproductive strategy and associated traits during biological invasions.

The simultaneous use of multiple biological control agents requires sufficient information of intraguild predation (IGP). In this work, the prey preferences of immature stages of the ladybird beetle Hippodamia variegata (Goeze) (Coleoptera Coccinellidae) for non-parasitized and parasitized adults of Aphis gossypii Glover (Hemiptera Aphididae) by the parasitoid wasp Lysiphlebus fabarum (Marshall) (Hymenoptera Braconidae) were investigated at the laboratory choice experiments. Moreover, the effects of these different feeding regimes of the immature stages were determined for the life history traits of emerging H. variegata and its progeny. Results revealed that the second and third instar larvae of H. variegata revealed no preference for the non-parasitized and parasitized aphids. The immature developmental time of H. variegata fed on the parasitized aphids was significantly shorter than those fed on the non-parasitized aphids; however, the weight of the emerged adults did not differ significantly. Contrary to this, in the next generation, the egg and hatched larval size of H. variegata fed on the parasitized aphids were significantly larger than those fed on the non-parasitized aphids, whereas the mean and total number of eggs laid by H. variegata fed on the parasitized aphids was significantly smaller than for those fed on the non-parasitized aphids. The feeding regimes at maternal rearing conditions revealed a significant effect on the progeny, as the weight of the emerging male and female adults increased. Combined with the simultaneous application of the two biological control agents, the negative effects resulting from feeding on the IG prey, L. fabarum, may be compensated by the fitness gained by the IG predator, H. variegata, and its progeny. Hence, the asymmetric IGP may not have a negative impact on the biological control of A. gossypii. [ABSTRACT FROM AUTHOR]

In central Japan, Harmonia yedoensis is a specialist ladybird that is confined to pine tree habitats, whereas its sibling species Harmonia axyridis is a generalist that feeds on a wide range of aphid species in nature. Interestingly, H. axyridis is not distributed in the Ryukyu Archipelago, southern Japan. We hypothesized that the ecological niche of H. yedoensis should be wider in the Ryukyu Archipelago, where its competitor species in central Japan, H. axyridis, is absent. We undertook fieldwork and a survey of published works to examine habitat utilization by H. yedoensis in the Ryukyu Archipelago. We found that H. yedoensis adults in the Ryukyu Archipelago visited several kinds of deciduous trees, including wild tamarind, Chinese hibiscus, Taiwan cherry and Malayan banyan, as well as pine trees. These observations suggest that habitat generalization has occurred in H. yedoensis in the Ryukyu Archipelago, where it does not compete with H. axyridis. [ABSTRACT FROM AUTHOR]

The wood-feeding cockroach Panesthia angustipennis spadica Shiraki (Blaberidae) is a gregarious species, and its groups containing both nymphs and adults have often been observed in the field. To clarify the effects of aggregation with a parent and siblings on nymphal development in P. angustipennis spadica, we raised nymphs of this species in the laboratory under the following three experimental categories: (a) aggregate with a female adult; (b) aggregate without a female adult; and (c) solitary. Survival rates did not significantly differ among the three categories. Unexpectedly, our results clearly showed that nymphs raised in aggregates with a female adult were smaller, lighter, and reached a lower instar than those raised without a female adult. These results indicate that the presence of a female adult does not positively affect nymphal development; i.e. there may be no direct parental care in this species as indicated by previous studies. Moreover, solitary nymphs grew faster and larger than nymphs in aggregates, showing that the aggregation with siblings also has no positive effect on nymphal development. Thus, it seems likely that external factors, e.g. predation and environmental conditions, may drive aggregating behavior in P. angustipennis spadica. [ABSTRACT FROM AUTHOR]

Abstract??The mushroom genusAmanitahas a spectrum of chemical compounds affecting survival and performance of animals. Ibotenic acid is one of such compounds found in someAmanitamushrooms. We studied the effects of ibotenic acid and its derivative, muscimol, on egg-to-pupa survival, pupation time, and pupal size in fiveDrosophilaspecies (Diptera: Drosophilidae),Drosophila bizonata,Drosophila angularis,Drosophila brachynephros,Drosophila immigrans, andDrosophila melanogaster. The first three species are mycophagous and use a wide range of mushrooms for breeding, whereasD. immigransandD. melanogasterare frugivorous. We reared fly larvae on artificial medium with 500, 250, 125, and 62.5??g/ml of ibotenic acid and/or musimol. The three mycophagous species were not susceptible to ibotenic acid, whereas the two frugivorous species were affected. In experiments withD. melanogaster, muscimol was less toxic than ibotenic acid. [ABSTRACT FROM AUTHOR]