Genetics and selective breeding of variation in wing truncation in a flightless aphid control agent

Augmentative biological control by predaceous ladybird beetles can be improved by using flightless morphs, which have longer residence times on the host plants. The two‐spot ladybird beetle, Adalia bipunctata (L.) (Coleoptera: Coccinellidae), is used for the biological control of aphids in greenhouses and on urban trees. Flightlessness due to truncated wings occurs at very low frequency in some natural populations of A. bipunctata. Pure‐breeding strains of this 'wingless' genotype of A. bipunctata can easily be obtained in the laboratory. Such strains have not been commercialized yet due to concerns about their reduced fitness compared to wild‐type strains, which renders mass production more expensive. Wingless strains exhibit, however, wide intra‐population phenotypic variation in the extent of wing truncation which is related to fitness traits. We here use classical quantitative genetic techniques to study the heritability and genetic architecture of variation in wing truncation in a wingless strain of A. bipunctata. Split‐families reared at one of two temperatures revealed strong family‐by‐temperature interaction: heritability was estimated as 0.64 ± 0.09 at 19 °C and 0.29 ± 0.06 at 29 °C. Artificial selection in opposite directions at 21 °C demonstrated that the degree of wing truncation can be altered within a few generations resulting in wingless phenotypes without any wing tissue (realized h² = 0.72), as well as those with minimal truncations (realized h² = 0.61) in two replicates. The latter lines produced more than twice as many individuals. This indicates that selective breeding of wing truncation may be exploited to improve mass rearing of flightless strains of A. bipunctata for commercial biological control. Our work illustrates that cryptic variation can also be a source for the selective breeding of natural enemies.     

Intraguild predation by Harmonia axyridis (Coleoptera: Coccinellidae) on native insects in Europe: molecular detection from field samples

Intraguild predation (IGP) is a potential mechanism of negative effects on native species populations by invasive non‐native species such as Harmonia axyridis. Molecular techniques (polymerase chain reaction) were used to probe for the presence of various insect (coccinellid, syrphid and chrysopid) prey DNA in the guts of 177 H. axyridis larvae field‐collected in England, France, Germany, Slovakia and the Czech Republic in 2010. Three of the four target prey species were detected in the guts of H. axyridis at the following rates: Adalia decempunctata 9.6%, Adalia bipunctata 2.8% and Episyrphus balteatus 2.8%. IGP on Chrysoperla carnea was not detected. IGP detection of at least one target species was made in England, France, Slovakia and the Czech Republic, but not in Germany. These results strengthen the evidence that H. axyridis is a very generalist predator. Results are discussed in the context of their ecological significance.     

Mating stimulates female feeding: testing the implications for the evolution of nuptial gifts

Nutritional benefits from nuptial gifts have been difficult to detect in some species, raising the question: what maintains nuptial feeding when gifts do not benefit females? The sensory trap hypothesis proposes that nuptial feeding may be explained by pre‐existing sensory responses that predispose females to ingest gifts. Recent studies have shown that male seminal proteins can induce a nonspecific increase in female feeding after mating, which may represent a sensory trap for nuptial feeding if it results in increased intake of post‐mating gifts. I tested these ideas using female beetles that ingest a spermatophore after mating. I show that males stimulate strongly increased female feeding post‐mating. However, there was little evidence for dose dependence in the feeding response that could allow males to stimulate feeding beyond the female optimum. Moreover, the post‐mating feeding response could not explain nuptial feeding: despite feeding more in general, newly mated females were less likely than nonmated females to ingest spermatophore gifts.     

Condition-dependent ejaculate size and composition in a ladybird beetle

Sexually selected male ejaculate traits are expected to depend on the resource state of males. Theory predicts that males in good condition will produce larger ejaculates, but that ejaculate composition will depend on the relative production costs of ejaculate components and the risk of sperm competition experienced by low- and high-condition males. Under some conditions, when low condition leads to poorer performance in sperm competition, males in low condition may produce ejaculates with higher sperm content relative to their total ejaculate and may even transfer more sperm than high-condition males in an absolute sense. Previous studies in insects have shown that males in good condition transfer larger ejaculates or more sperm, but it has not been clear whether increased sperm content represents a shift in allocation or simply a larger ejaculate, and thus the condition dependence of ejaculate composition has been largely untested. We examined condition dependence in ejaculate by manipulating adult male condition in a ladybird beetle (Adalia bipunctata) in which males transfer three distinct ejaculate components during mating: sperm, non-sperm ejaculate retained within the female reproductive tract, and a spermatophore capsule that females eject and ingest following mating. We found that high condition males indeed transferred larger ejaculates, potentially achieved by an increased rate of ejaculate transfer, and allocated less to sperm compared with low-condition males. Low-condition males transferred ejaculates with absolutely and proportionally more sperm. This study provides the first experimental evidence for a condition-dependent shift in ejaculate composition.     

Overwintering, phenology and fecundity of Harmonia axyridis in comparison with native coccinellid species in Italy

Classical biological control is generallyunderstood as an environmentally safe practiceof insect pest management. However, questionshave been raised about possible negativeeffects for native species. As part of a riskassessment study a semi-field trial was carriedout in northern Italy in order to compare theoverwintering of the native species Propylea quatuordecimpunctata (L.) and Adalia bipunctata (L.) (Coleoptera:Coccinellidae) with the exotic Harmoniaaxyridis (Pallas) (Coleoptera: Coccinellidae).All the experiments were carried out from April1998 to April 2001. Harmonia axyridis wasable to overwinter successfully in northernItaly. Fecundity, oviposition rate, longevityand rate of increase of overwintering femaleswere calculated and the phenology of the threespecies was studied. Harmonia axyridisoverwintering mortality (31.9%) was lower incomparison with mortality of native species(68.9% for P. quatuordecimpunctata and61.3% for A. bipunctata). As aconsequence, post-overwintering rate ofincrease of the exotic species was higher.Longevity of overwintered females was similaramong the species. Mean fecundity of H. axyridis (783.8 eggs per female) was slightlyhigher than A. bipunctata (720.2 eggs perfemale), and much higher than P. quatuordecimpunctata (193.7 eggs per female). H. axyridis and A. bipunctatacompleted four generations and P. quatuordecimpunctata three generations in a year. The results show that H. axyridisappears to posses a high potential for establishment in Italy.     

Does the satiation hypothesis account for the differences in efficacy of coccidophagous and aphidophagous ladybird beetles in biological control? A test with Adalia bipunctata and Cryptolaemus montrouzieri

Mills (1982) formulated the satiationhypothesis in order to explain why ladybirdsare generally more effective in the biologicalcontrol of coccids than of aphids:aphidophagous ladybirds are supposedly lessefficient because they become more rapidlysatiated than coccidophagous ladybirds. If thisis true, the former should spend less timeeating than the latter. This hypothesis istested in this study using the coccidophagousCryptolaemus montrouzieri Mulsant(Coleoptera: Coccinellidae) and theaphidophagous Adalia bipunctata (L.) (Coleoptera: Coccinellidae). Contrary to Mills'(1982) prediction the coccidophagous ladybirddid not feed continuously and even spent lesstime feeding than the aphidophagous ladybird.Furthermore, the gut capacity of C. montrouzieri is smaller than that of A. bipunctata.     

Isolation and characterization of highly polymorphic microsatellite loci in the 2‐Spot Ladybird,Adalia bipunctata

Contrary to theoretical predictions, female 2‐spot ladybirds (Adalia bipunctata) mate many more times than necessary to maintain high fertilisation success and may gain through the acquisition of material or genetic benefits. In order to investigate this mating system in detail, microsatellite markers have been isolated using a modified enrichment technique. Thirty‐nine loci were successfully amplified by polymerase chain reaction (PCR), of which only two were monomorphic. Detailed characterization of ten loci revealed very high levels of polymorphism. These markers are likely to be invaluable tools with which to study population genetics and patterns of paternity in this species.     

The inheritance of a wingless character in the 2spot ladybird (Adalia bipunctata)

A viable wingless 2spot ladybirdAdalia bipunctata (L.) was found in the wild. Breeding through four generations revealed that the wingless trait was controlled by a recessive allele which displays variable levels of expression. The wingless ladybird is discussed in relation to its potential as a biocontrol agent. One ladybird also occurred in this stock which is suggestive of a supergene controlling the colour polymorphism in this species.     

Effect of intraguild predation on the survival and development of three species of aphidophagous ladybirds: consequences for invasive species

Abstract 1 Survival and development of hatchling larvae of three aphidophagous ladybirds (Coleoptera: Coccinellidae), Harmonia axyridis Pallas, Coccinella septempunctata brucki Mulsant and Adalia bipunctata Linnaeus, when fed their own and the other species eggs were recorded. 2 In all three species, the larvae survived when fed conspecific eggs. 3 The percentage of larvae of H. axyridis that survived decreased to 35% and 85% when fed eggs of A. bipunctata and C. s. brucki, respectively. All the larvae of A. bipunctata and C. s. brucki died after eating eggs of H. axyridis. None of the larvae of C. s. brucki died after eating eggs of A. bipunctata, whereas 46% of those of A. bipunctata died after eating eggs of C. s. brucki. 4 In general, larvae were reluctant to eat the eggs of other species. However, larvae of C. s. brucki showed less reluctance than H. axyridis to eat the eggs of A. bipunctata. 5 The consequence of this for invasive species of ladybird is discussed.     

A study of nonrandom mating in a British population of the two-spot ladybird with a high frequency of the melanic morph

In some studies of the two-spot ladybird (Adalia bipunctata), melanic males have been found in excess over the typical morph in matings. Data suggest that a genetic female mating preference is responsible. The mating advantage of melanic males may be important in maintaining a polymorphism between melanic and typical ladybirds in many populations in the United Kingdom (U.K.). It has been reported that preference frequency varies linearly with melanic frequency throughout most of the U.K. One particular population ofAdalia bipunctata near Aberdare, South Wales, is noted for its high frequency of melanic individuals. It has been suggested that local environmental factors account for the high melanic frequency in this population. It is also possible, however, that a female mating preference may be at least partly responsible for the high frequency of melanics (as has been proposed for the rest of the U.K.). In this study, experiments have been performed to determine the level of female mating preference in the Aberdare population. No evidence was found for any mating advantage to melanic males. There was inconsistent and unexpected evidence that melanic females were overrepresented in matings, but the cause for this was unclear. Female mating preference does not appear, therefore, to be responsible for the high melanic frequency in the population ofAdalia bipunctata near Aberdare. There is not a simple association between mating preference and melanic frequency in U. K. populations of the two-spot ladybird.