Behavioural switch of female Mediterranean fruit fly, Ceratitis capitata: mating and oviposition activity in outdoor field cages in Hawaii

1 Laboratory-reared normal, and wild female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), were assayed in outdoor field cages to assess the impact of a mating-induced behavioural switch on mating and subsequent oviposition activity. 2 Virgin females preferred interactions with males leading to mating over attraction to, and oviposition in, artificial yellow spheres containing guava odour or green apples hung in a guava tree. Laboratory-reared females previously mated with either laboratory-reared normal males or laboratory-reared irradiated (sterile) males showed little interest in remating with males and instead, were much more likely to be found arrested on artificial and real fruit and ovipositing. Oviposition on artificial fruit was five times greater by females that had mated with either normal or irradiated males than by virgin females. Wild females showed similar qualitative changes in the mating-induced behavioural switch; however, oviposition activity was significantly less than for laboratory-reared females. 3 These results confirm that mating has a profound effect on the behaviour of female Mediterranean fruit flies and that irradiated males are functionally equal with normal males (lab-reared or wild) in their ability to alter female behaviour. These results are discussed in the context of the sterile insect technique for control of Mediterranean fruit flies in the field.     

Host acceptance and sex allocation of Nasonia wasps in response to conspecifics and heterospecifics

Species recognition is an important aspect of an organism''s biology. Here, we consider how parasitoid wasps vary their reproductive decisions when their offspring face intra- and interspecific competition for resources and mates. We use host acceptance and sex ratio behaviour to test whether female Nasonia vitripennis and Nasonia longicornis discriminate between conspecifics and heterospecifics when ovipositing. We tested pairs of conspecific or heterospecific females ovipositing either simultaneously or sequentially on a single host, using strains varying in their recent history of sympatry. Both N. vitripennis and N. longicornis rejected parasitized hosts more often than unparasitized hosts, although females were more likely to superparasitize their own species in the sequential treatment. However, sex ratio behaviour did not vary, suggesting similar responses towards conspecifics and heterospecifics. This contrasts with theory predicting that heterospecifics should not influence sex ratios as their offspring do not influence local mate competition, where conspecifics would. These non-adaptive sex ratios reinforce the lack of adaptive kin discrimination in N. vitripennis and suggest a behavioural constraint. Discrimination between closely related species is therefore context dependent in Nasonia. We suggest that isolating mechanisms associated with the speciation process have influenced behaviour to a greater extent than selection on sex ratios.     

Oviposition behaviour in Paysandisia archon (Lepidoptera: Castniidae)

Paysandisia archon (Burmeister, 1880) (Lepidoptera: Castniidae) is a South American insect developing on palms and recently introduced in Europe where it damages most palm species. Understanding the oviposition behaviour would be decisive for risk assessment and pest management but key points on oviposition behaviour are missing. Using wind tunnel and field experiments, we investigated the oviposition timing, the attraction behaviour of mated females to palms and the different steps of oviposition behaviour. Results showed that oviposition behaviour occurred between 3 pm and 6 pm. In the field, gravid females were significantly more attracted by the palm crown than virgin females. The ovipositing females exhibited distinctive steps before ovipositing. Subsequent to alighting on the crown, pre-oviposition behaviour was characterized by two main behavioural steps: walking and probing the surface with antennae and ovipositor. After the choice of oviposition place, the gravid female remains motionless and the extendible ovipositor is deeply introduced into the upper fibrous part of the crown. About 10 eggs can be lays at the same place. Finally, the female starts to walk again and reinitiates the same behavioural sequences. This paper supports the hypothesis that odours from the crown may play a key role in gravid female attraction. The study assessed that P. archon lays on the palm crown, the part of the palm that should be treated for population monitoring.     

Behavior and Activity Pattern of Cephalonomia stephanoderis (Hymenoptera: Bethylidae) Attacking the Coffee Berry Borer,Hypothenemus hampei (Coleoptera: Scolytidae)

We describe behavioral sequences and daily activities of pre-ovipositing and ovipositing females of Cephalonomia stephanoderis (Hymenoptera: Bethylidae), an ectoparasitoid of the coffee berry borer, Hypothenemus hampei (Coleoptera: Scolytidae). Noticeable behavioral differences among preovipositing and ovipositing females include host examination, host stinging—probing, host feeding, and the oviposition per se. The female of C. stephanoderis feeds primarily on host eggs, but pupae are also exploited, mainly by pre-ovipositing females. After the onset of the oviposition period, C. stephanoderis examines the pupae repeatedly, stings them at frequent intervals, and spends more time feeding than during the pre-oviposition period. Host paralysis is linked both to host feeding and oviposition. It induces irreversible developmental arrest, which presumably allows preservation of the host until subsequent utilization, and contributes to successful offspring development, particularly by reducing host motility. Oviposition consists in a host selection process, a prolonged period of preparation of the potential host, and the egg-laying phase itself. Under our experimental conditions, pre-ovipositing and ovipositing females are active 17% and 36% of the day, respectively. Host handling time averages 6% and 23% in pre-ovipositing and ovipositing females, respectively. All coffee berry borer developmental stages are exploited by C. stephanoderis females, either for host feeding and/or oviposition activities. Such flexible behavior is advantageous given that host availability is limited inside the coffee berries.     

Characteristics of oviposition behaviour of the pollen beetle, Meligethes aeneus on four different host plants

Oviposition behaviour of Meligethes aeneus F. (Coleoptera: Nitidulidae) is characterised and quantified on four different plant species. Six behavioural components are identified: W—walking, WA—walking with abdomen on surface, R—resting, B—biting, AOH—placing abdomen over the bite hole and OVI—oviposition. Comparison of host acceptance behaviours on Brassica napus L., Brassica juncea (L.) Czern, Brassica nigra (L.) Koch and Sinapis alba L. showed that S. alba was accepted as a host only after a long exposure to the plant. Behaviour on the Brassica species was similar, however on B. nigra beetles spent a high proportion of time actually ovipositing. We conclude that important cues for oviposition are located both on the bud surface and inside the bud.     

Kin recognition by the predatory mite Iphiseius degenerans: discrimination among own, conspecific, and heterospecific eggs

1. Kin recognition is important in many social insects, but has also been found in several nonsocial insects such as parasitoids, where it plays an important role in oviposition behaviour. In nonparasitic arthropods, however, the fitness of ovipositing females also depends on the oviposition behaviour of related and unrelated females, especially when eggs are oviposited in clusters by several females. 2. In this paper, kin recognition in a predatory mite, Iphiseius degenerans, is studied. Mothers are capable of determining offspring sex ratio, and cannibalism on juvenile stages is a common phenomenon. Therefore, kin recognition is expected to occur in this predator. 3. Oviposition behaviour of this species is particularly interesting because it alternates foraging bouts in flowers with deposition of a single egg at a time on a leaf, where predation risk is lower. The eggs are not scattered but are deposited in clusters. After feeding in a flower, females therefore have to locate clusters of eggs. 4. Experiments on two‐choice arenas showed that females prefer to oviposit close to conspecific eggs rather than close to heterospecific eggs. Females also showed a preference for ovipositing near closely related conspecific eggs rather than more distantly related eggs. 5. Females tended to displace eggs of heterospecifics more frequently than eggs of conspecifics. 6. These behavioural observations show that females can discriminate not only between conspecific and heterospecific eggs but also between eggs that vary in degree of relatedness. This enables females to oviposit in clusters containing related eggs and thus avoid cannibalism by non‐kin and/or produce adaptive sex ratios despite the fact that the adults commute between flowers and leaves.     

A sex allocation cost to polyandry in a parasitoid wasp

The costs and benefits of polyandry are central to understanding the near-ubiquity of female multiple mating. Here, we present evidence of a novel cost of polyandry: disrupted sex allocation. In Nasonia vitripennis, a species that is monandrous in the wild but engages in polyandry under laboratory culture conditions, sexual harassment during oviposition results in increased production of sons under conditions that favour female-biased sex ratios. In addition, females more likely to re-mate under harassment produce the least female-biased sex ratios, and these females are unable to mitigate this cost by increasing offspring production. Our results therefore argue that polyandry does not serve to mitigate the costs of harassment (convenience polyandry) in Nasonia. Furthermore, because males benefit from female-biased offspring sex ratios, harassment of ovipositing females also creates a novel cost of that harassment for males.     

Oviposition of resistant and susceptible strains of Drosophila melanogaster in the presence of deltamethrin

Oviposition of three strains of Drosophila melanogaster in the presence of deltamethrin was observed. These strains had different levels of physiological susceptibility to deltamethrin. Two-choice tests were conducted with couples of flies in petri-dish arenas containing two oviposition dishes. On the first day of the experiment, females were given a choice between a treated oviposition dish and an untreated control dish. On the second day of the experiment, two control oviposition dishes were given to females. Although individual females showed a tendency to aggregate their eggs in one of the dishes, control experiments demonstrated an overall equal distribution of eggs between the dishes. When one of the two oviposition dishes in the arena was treated with deltamethrin, the percentage of females ovipositing and the mean number of eggs laid by females were reduced, compared with control arenas. Females avoided the treated oviposition dish and laid significantly more eggs on the control dish. Furthermore, when the deltamethrin concentration was increased on the first day, female flies postponed their oviposition and laid significantly more eggs on the second day. The resistant strain, SR, demonstrated the same capacity to select the untreated site for oviposition as the susceptible strain, but it showed a smaller oviposition reduction and egg retention. The relationship between physiological and behavioural susceptibility to deltamethrin is discussed.     

Seasonal decline in plant defence is associated with relaxed offensive oviposition behaviour in the viburnum leaf beetle Pyrrhalta viburni

1. Plant defence of Viburnum shrubs against oviposition by its specialist herbivore, the viburnum leaf beetle [VLB Pyrrhalta viburni(Paykull)], involves an egg‐crushing wound response in twigs. Although the response is variable among Viburnum species, it can have a strong impact on egg survivorship. Beetles typically aggregate egg masses with conspecifics along infested twigs, forming clusters that can overwhelm the twig response. It was investigated whether twig responses and beetle oviposition behaviour vary seasonally. 2. In a field experiment, twig defences decreased towards the end of the VLB oviposition period: wound response of the North American Viburnum dentatum L. and the European V. opulus L. was reduced by 100% and 54%, respectively, in September compared with the July to August period. 3. Oviposition trials demonstrated a corresponding behavioural change: VLB females displayed aggregative oviposition in August, but not in September. 4. Further tests revealed that late‐season VLB females reverted to aggregative oviposition after being kept on uninfested twigs, whereas females kept on heavily infested twigs did not. This behavioural change suggests that relaxation of aggregative oviposition originates from cues associated with high densities of egg masses. 5. Relaxation of aggregative oviposition may be adaptive (and beneficial for invasion) on shrubs with low levels of defences by reducing intra‐specific competition.     

Faster-X Evolution of Gene Expression in Drosophila

DNA sequences on X chromosomes often have a faster rate of evolution when compared to similar loci on the autosomes, and well articulated models provide reasons why the X-linked mode of inheritance may be responsible for the faster evolution of X-linked genes. We analyzed microarray and RNA–seq data collected from females and males of six Drosophila species and found that the expression levels of X-linked genes also diverge faster than autosomal gene expression, similar to the “faster-X” effect often observed in DNA sequence evolution. Faster-X evolution of gene expression was recently described in mammals, but it was limited to the evolutionary lineages shortly following the creation of the therian X chromosome. In contrast, we detect a faster-X effect along both deep lineages and those on the tips of the Drosophila phylogeny. In Drosophila males, the dosage compensation complex (DCC) binds the X chromosome, creating a unique chromatin environment that promotes the hyper-expression of X-linked genes. We find that DCC binding, chromatin environment, and breadth of expression are all predictive of the rate of gene expression evolution. In addition, estimates of the intraspecific genetic polymorphism underlying gene expression variation suggest that X-linked expression levels are not under relaxed selective constraints. We therefore hypothesize that the faster-X evolution of gene expression is the result of the adaptive fixation of beneficial mutations at X-linked loci that change expression level in cis. This adaptive faster-X evolution of gene expression is limited to genes that are narrowly expressed in a single tissue, suggesting that relaxed pleiotropic constraints permit a faster response to selection. Finally, we present a conceptional framework to explain faster-X expression evolution, and we use this framework to examine differences in the faster-X effect between Drosophila and mammals.     

A Quantified Ethogram for Oviposition in Triturus Newts: Description and Comparison of T. helveticus and T. vulgaris

Female newts of the genus Triturus deposit and wrap their eggs individually in the submerged leaves of aquatic macrophytes. Although this behaviour has previously been described, the different elements of the oviposition process have not been fully characterized nor any attempt made to quantify the behavioural elements. The study examined the oviposition behaviour of the two similarly sized species, Triturus helveticus and T. vulgaris on a standardized substrate macrophyte, Rorippa nasturtium–aquaticum. Continuous focal sampling was used to develop a baseline of discrete behavioural elements enabling quantification and comparison of oviposition behaviour between the two species. The results showed that the same pattern of elements was followed for each egg laid and the same key elements of the process were present in each newt species. Although these are broadly similar in size, there were striking differences in certain aspects of the oviposition sequence between the two species. Key findings were that leaf sniffing and leaf flexing and a measure of the duration of ovipositing were all significantly greater in females of T. helveticus and females of T. vulgaris laid significantly more eggs than those of T. helveticus in a standard observation period. The work presented here defines a baseline ethogram and shows how it can be used to reveal quantifiable differences in closely related species. This demonstrates its value in furthering our understanding of oviposition – a key aspect of female behaviour currently understudied in Triturus behavioural ecology, despite its intrinsic interest and value in understanding recruitment and maintenance of populations.     

Effect of adult females' predation risk on oviposition site selection in a water strider

In many egg‐laying species, females avoid ovipositing at sites where the predation risk is high. Previous studies have mainly focused on the risk for offspring. The effect of predation risk for the females has been considered in some taxa in which parents spend much time at an oviposition site for parental care or mating (e.g., birds, amphibians). In species in which females do not perform activities other than oviposition at sites, the effect of predation risk for females on oviposition site selection has been rarely investigated. We examined whether the predation risk for ovipositing females affects the decision on oviposition in the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae). Adult A. paludum are preyed upon by a backswimmer, Notonecta triguttata (Motschulsky) (Heteroptera: Notonectidae), but N. triguttata does not prey upon eggs or early instars of A. paludum. We allowed female A. paludum to oviposit under one of three conditions: in presence of N. triguttata, in presence of its chemicals, and in absence of the predator or its cues (control). Female A. paludum less frequently oviposited in presence of N. triguttata than in its absence. Oviposition frequency did not differ between females in presence of chemicals of N. triguttata vs. those in the control. Female A. paludum recognized the predation risk upon themselves from the presence of N. triguttata and avoided ovipositing. This study is the first to directly show that the predation risk upon ovipositing females changes oviposition site selection in species in which the time spent at an oviposition site is short.     

Oviposition decreased in response to enriched water: a field study of the pitcher-plant mosquito, Wyeomyia smithii

Abstract.  1. Environmental cues are known to influence oviposition behaviour in mosquitoes, with important consequences for larval survival and insect population dynamics. Enriched microhabitats have been shown to be preferred oviposition sites.
2. In a field experiment designed to determine whether ovipositing mosquitoes are sensitive to different levels of nutrient enrichment, new pitcher-plant ( Sarracenia purpurea ) leaves were opened and enriched with 0, 2, or 20 dead ants, and the number of pitcher-plant mosquito ( Wyeomyia smithii ) larvae resulting from subsequent oviposition were measured.
3. Oviposition rates were higher in leaves with low levels of enrichment (0 and 2 ants per leaf), although larval development was enhanced at the highest enrichment level.
4. Results suggest that, although these mosquito larvae are nutrient limited, ovipositing females preferentially avoid highly enriched leaves. This counterintuitive result may be due to low oxygen concentrations or a masked cue in enriched leaves, and contrasts with other oviposition studies.     

Comparing Behavioural Patterns of Thrips tabaci Lindeman on Leek and Cucumber

This study documents the behavioural repertoire of Thrips tabaci. Seventeen behavioural elements shown by adult females were videotaped, described and classified into seven behavioural categories. The comparison of thrips behaviour on leek and cucumber over 24 h (L:D = 16:8) revealed that on cucumber thrips spent significantly less time being inactive, but significantly more time with feeding resulting in significantly higher feeding damage compared to leek. Oviposition behaviour incidence and oviposition rate were somewhat, but not significantly lower on cucumber compared to leek. Sequences of leaf surface exploration lasted significantly longer on cucumber than on leek. Thrips females were as active during light as during dark periods. Our results show that T. tabaci females spend different amounts of time with resting, feeding and explorative behaviours on leek and cucumber, thus indicating differences in host use on these two crops.     

Juvenile prey induce antipredator behaviour in adult predators

It is generally assumed that the choice of oviposition sites in arthropods is affected by the presence of food for the offspring on the one hand and by predation risk on the other hand. But where should females oviposit when the food itself poses a predation risk for their offspring? Here, we address this question by studying the oviposition behaviour of the predatory mite Amblyseius swirskii in reaction to the presence of its counterattacking prey, the western flower thrips Frankliniella occidentalis. We offered the mites a choice between two potential oviposition sites, one with and one without food. We used two types of food: thrips larvae, which are predators of eggs of predatory mite but are consumed by older predator stages, and pollen, a food source that poses no risk to the predators. With pollen as food, the predators preferred ovipositing on the site with food. This might facilitate the foraging for food by the immature offspring that will emerge from the eggs. With thrips as food, female predators preferred ovipositing on the site without thrips. Predators that oviposited more on the site with thrips larvae killed more thrips larvae than females that oviposited on the site without food, but this did not result in higher oviposition. This suggests that the females killed thrips to protect their offspring. Our results show that predators display complex anti-predator behaviour in response to the presence of counter-attacking prey.     

Re-evaluation of the role of corpora cardiaca in calling and oviposition behaviour of giant silk moths

Re-investigation of the role of the corpora cardiaca in the reproductive behaviour of the giant silkmoths, Hyalophora cecropia and Antheraea polyphemus, showed that this pair of glands plays no essential role, either in “calling” behaviour by virgin females or in increased oviposition due to mating. Removal of corpora cardiaca-corpora allata complexes, either from diapausing pupae or from freshly eclosed adult females, had no effect on the calling behaviour or on its timing in either species. Moreover after mating, these operated females laid eggs in the typical mated oviposition pattern. Furthermore, females in which there was only a nervous connection between the brain and the abdomen but no haemolymph circulation called normally and oviposited after mating.Although the corpora cardiaca were not essential for calling behaviour, hormogenates of corpora cardiaca-corpora allata complexes and blood from calling or ovipositing females induced a typical “calling” response in 30–60% of the isolated virgin H. cecropia abdomens tested. This activity was not species-specific as it was also found in Manduca sexta, but the restriction of major activity to corpora cardiaca extracts and haemolymph suggested that a neurosecretory factor may modulate the normal neural control of calling behaviour.     

Experience-induced preference for oviposition repellents derived from a non-host plant by a specialist herbivore

Foraging adults of phytophagous insects are attracted by host‐plant volatiles and supposedly repelled by volatiles from non‐host plants. In behavioural control of pest insects, chemicals derived from non‐host plants applied to crops are expected to repel searching adults and thereby reduce egg laying. How experience by searching adults of non‐host volatiles affects their subsequent searching and oviposition behaviour has been rarely tested. In laboratory experiments, we examined the effect of experience of a non‐host‐plant extract on the oviposition behaviour of the diamondback moth (DBM), Plutella xylostella, a specialist herbivore of cruciferous plants. Naive ovipositing DBM females were repelled by an extract of dried leaves of Chrysanthemum morifolium, a non‐host plant of DBM, but experienced females were not repelled. Instead they were attracted by host plants treated with the non‐host‐plant extract and laid a higher proportion of eggs on treated than on untreated host plants. Such behavioural changes induced by experience could lead to host‐plant range expansion in phytophagous insects and play an important role in determining outcome for pest management of some behavioural manipulation methods.