Is aggregated oviposition by the blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) really pheromone‐mediated?

When female blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) oviposit in aggregations on carrion, even‐aged larval offspring reportedly develop faster, and fewer are parasitized or preyed upon. The benefits of aggregated oviposition equally affect con‐ and heterospecific larvae sharing a resource. The benefits imply that female blow flies engage in coordinated, pheromone‐mediated oviposition behavior. Yet, repeated attempts to identify oviposition pheromones have failed invoking doubt that they exist. Simply by regurgitating and feeding on carrion, flies may produce attractive semiochemicals. If flies were to aggregate in response to feeding flies rather than ovipositing flies, then the semiochemical cue(s) may be associated with the salivary gland. Working with L. sericata and P. regina and using liver as a surrogate oviposition medium, we test the hypotheses, and present data in their support, that (i) gravid or nongravid females ovipositing and/or feeding on liver enhance its attractiveness to gravid and nongravid females; (ii) females respond to semiochemicals from feeding heterospecific females; (iii) females respond equally well to semiochemicals from feeding con‐ and heterospecific females; (iv) macerated head tissues of females applied to liver enhance its attractiveness; and (v) females in direct contact with and feeding on liver, but not when next to yet physically separated from liver, enhance attraction of flies. We conclude that oviposition site‐seeking females do not respond to an oviposition pheromone. Instead, they appear to coopt semiochemicals associated with feeding flies as resource indicators, taking chances that resources are suitable for oviposition, and that ovipositing flies are present.     

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.     

Experience‐ and age‐mediated oviposition behaviour in the yellow fever mosquito Stegomyia aegypti (=Aedes aegypti)

In repeated behaviours such as those of feeding and reproduction, past experiences can inform future behaviour. By altering their behaviour in response to environmental stimuli, insects in highly variable landscapes can tailor their behaviour to their particular environment. In particular, female mosquitoes may benefit from plasticity in their choice of egg‐laying site as these sites are often temporally variable and clustered. The opportunity to adapt egg‐laying behaviour to past experience also exists for mosquito populations as females typically lay eggs multiple times throughout their lives. Whether experience and age affect egg‐laying (or oviposition) behaviour in the mosquito Stegomyia aegypti (=Aedes aegypti) (Diptera: Culicidae) was assessed using a wind tunnel. Initially, gravid mosquitoes were provided with a cup containing either repellent or well water. After ovipositing in these cups, the mosquitoes were blood‐fed and introduced into a wind tunnel. In this wind tunnel, an oviposition cup containing repellent was placed in the immediate vicinity of the gravid mosquitoes. A cup containing well water was placed at the opposite end of the tunnel so that if the females flew across the chamber, they encountered the well water cup, in which they readily laid eggs. Mosquitoes previously exposed to repellent cups became significantly more likely to later lay eggs in repellent cups, suggesting that previous experience with suboptimal oviposition sites informs mosquitoes of the characteristics of nearby oviposition sites. These results provide further evidence that mosquitoes modify behaviour in response to environmental information and are demonstrated in a vector species in which behavioural plasticity may be ecologically and epidemiologically meaningful.     

Oviposition in Yponomeuta cagnagellus: the importance of contact cues for host plant acceptance

Small ermine moths (Yponomeutidae: Lepidoptera) are specialist herbivores. Species within the genus Yponomeuta are each specialized on a limited number of plant species, mainly within genera belonging to the Celastraceae. European Yponomeuta species have developed new specialized host affiliations, mainly on rosaceous hosts. Since these host shifts are reputed to be of consequence for speciation, the role of the ovipositing female is of particular interest. Study of the pre-oviposition behaviour of gravid Y. cagnagellus (Hb.) moths on host (Euonymus europaeus), non-host (Crateagus monogyna) and artificial oviposition substrates, provided information on the nature of the cues used for host plant acceptance and the insect’s perception of these cues. Host selection by adult females occurs with contact chemoreceptors probably located on the antennae or tarsi. MeOH-soluble, non-volatile phytochemical compounds washed from the host plant’s surface and applied on an artificial twig are sufficient to stimulate a complete sequence of behavioural elements leading to oviposition. Volatiles do not have a large effect on the pre-oviposition behaviour.     

Importance of plant physical cues in host acceptance for oviposition by Busseola fusca

Plant and surrogate stems exhibiting specific combinations of physical cues were used to determine which plant‐related stimuli influence the oviposition of Busseola fusca (Fuller) (Lepidoptera: Noctuidae). The number of eggs and egg batches laid per female increased with an increase in diameter of both natural and artificial stems. Direct observations of the oviposition behaviour (walking, antennating, and sweeping with the ovipositor) indicated that the female moths preferred oviposition supports with a large diameter and non‐pubescent or smooth surfaces over pubescent or rough ones. Pubescence and rough surfaces significantly affected the behavioural steps leading to oviposition by interfering with the ovipositor sweep process necessary to find a suitable oviposition site. Furthermore, more eggs and egg batches were laid on soft than rigid supports. The rigidity of the support affected the proper insertion of the ovipositor for egg deposition. Our results underline the importance of physical stimuli in B. fusca's choice of an oviposition site, which may facilitate the identification of potential host plants or preferred oviposition sites on a plant for this species.     

Cover Caption

Females of the common green blow fly, Lucilia sericata, ovipositing on rat carrion. When females oviposit in aggregations, even‐aged larval offspring develop faster and fewer are preyed upon. These benefits imply that females engage in coordinated, pheromone‐mediated oviposition behavior. Yet, new data show that oviposition site‐seeking females do not respond to pheromones but to semiochemicals associated with gravid or non‐gravid feeding flies, taking chances that resources are suitable for oviposition and that ovipositing flies are present (see pages 651–660). Image: Sean McCann.     

Oviposition Behaviour of Lygus rugulipennis and its Preferences for Plant Wounds

Little information is available on the oviposition behaviour and strategies of Lygus bugs, and these topics are unknown for most Miridae. Lygus rugulipennis, a widespread polyphagous species in the Palaearctic Region, is a serious pest of both herbaceous crops and fruit trees. Here we report on the L. rugulipennis oviposition behaviour and preferences. Experiments were carried out utilizing fresh green beans, and data were recorded using a computerized system for behavioural acquisitions. The L. rugulipennis female examines the plant substrate by means of the labial tip and then probes into the tissue using her stylets. Once the substrate is suitable, the female inserts her ovipositor precisely in the point previously probed and lays an egg. To investigate females’ oviposition preferences, their behaviour was compared on healthy green beans vs. artificially wounded green beans, with different types of wounds and at different time intervals (1?h, 6?h, 12?h, 24?h and 48?h) after wounding. Through labial and stylet examination, females distinguished healthy and wounded tissues and significantly preferred to oviposit in the latter, with different egg densities depending on the type of wound. Such preference changed at different time intervals after wounding. The lower resistance of the wounded substrate to ovipositor insertion, compared with the healthy substrate, was confirmed through a penetrometer test. Lygus preference for damaged tissues is discussed in terms of oviposition strategies in the insect-plant interaction context and strategies to escape natural enemies.     

Long-term changes in the oviposition behaviour of the cabbage butterfly, Pieris rapae, induced by contact with plants

ABSTRACT. Contact between the tarsi and cabbage foliage was found to have important influences on the oviposition behaviour of Pieris rapae , whereas contact between the ovipositor tip and cabbage foliage showed no influence. Oviposition on a leaf disc was not influenced by the presence of previously laid eggs. Gravid female P.rapae showed an increased tendency to approach and land on host or non-host foliage, or on yellow or green cards, after tarsal contact with cabbage foliage. This effect persisted for at least 72h. It was accompanied by an increased tendency to oviposit on non-host plants, most obviously within seconds of contact with cabbage, but with some persistence after 72h. Contact with a non-host leaf (lettuce) reduced the tendency of females to land on plants, but this effect was less pronounced than the one induced by contact with cabbage. These behavioural changes were immediate as well as long-term and were not associated with changes in the ovaries, so they probably result from the direct effects of sensory input on the CNS. Such behaviour probably increased the likelihood that gravid females would locate host-plants for oviposition in the field. For tests of host-plant specificity, the implications are that the behaviour of gravid females towards an array of plants might differ according to the presence or absence of hosts, not only during the test, but also during the preceding several days.     

Quantitative genetics of oviposition behaviour and interactions among oviposition traits in the sand cricket

We performed a quantitative genetic study of oviposition behaviours and oviposition traits in the sand cricket Gryllus firmus. Egg survival in crickets depends on the depth at which they are inserted into the soil with the ovipositor. We examined whether egg depth depends on ovipositor length alone, or on both morphological and behavioural traits associated with oviposition. Heritability estimates were high (h² >0.5) for ovipositor length and small (h²=0.2) for oviposition behaviours. Negative genetic correlations between ovipositor length and some behavioural traits (digging depth and the behavioural component of egg depth) indicated compensation between oviposition traits on egg depth. Because of behavioural compensation, females with different ovipositor lengths subject to stabilizing selection on egg depth could have equal fitnesses. Females laid their eggs deeper, and their eggs were marginally more evenly distributed in dry than in wet sand. This suggests adaptive phenotypic plasticity in laying behaviour, but may also result from physical constraints of the substrate on the insertion of the ovipositor. The absence of significant between-family variation in oviposition traits in response to sand moisture indicates low evolutionary potential for phenotypic plasticity in oviposition traits according to soil moisture. In highly unpredictable environments, females could spread the risk of desiccation by laying eggs at different depths independently of environmental conditions (bet hedging). Our results show significant additive genetic influences on the ability of a female to spread risks as measured by genetic variation in egg distribution, suggesting that a bet-hedging strategy of egg laying has the potential to evolve in this population. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Ovipositing females of a short‐lived gall midge take time to assess suboptimal grass seed heads

Abstract The short‐lived adult wheat midge Sitodiplosis mosellana deposits eggs on the seed head of various grasses close to the developing seeds on which larvae feed. The time taken to make egg‐laying decisions is investigated using three types of wheat Triticum aestivum seed heads. Young Roblin, Old Roblin and Young Key differ in their effects on ovipositing females (72%, 22% and 6% of eggs in choice tests, respectively) and effects on feeding larvae (75%, 25% and 5% larval survival, respectively). Within seconds of arriving, the female is able to distinguish Young Roblin from the two lower‐ranked types. However, the lower‐ranked types are not rejected at this time. Instead, all head types are examined before the female eventually flies away. On Young Roblin, probing with the ovipositor is the first behaviour that occurs. Thereafter probing and insertion of the ovipositor occupy most of the female’s time and behavioural transitions tend to be ‘progressive’, signalling a shift from low to high intensity examining. Differences between females visiting Old Roblin and Young Key are significant but take longer to emerge. On both, sitting is the first behaviour but, over the next 5–10 min, the female on Young Key exhibits more sitting, walking and ‘regressive’ transitions than the female on Old Roblin. It is suggested that, when the ovipositing female is short‐lived and incapable of controlled flight in all but essentially windless conditions, her behaviour is designed to thoroughly, rather than rapidly, examine a suboptimal host before abandoning it for the uncertain future of finding a better host.     

The invasive moth Paysandisia archon in Europe: Biology and control options

The palm borer moth (Paysandisia archon, Burmeister) is a member of the Castniidae family originally from South America and is currently included in the A2 list of the OEPP/EPPO. This moth was introduced to Europe in 2000 through ornamental palms. Since its accidental introduction, it has become a major threat for natural stands of native palms, as well as for nurseries and gardens in the Mediterranean basin. To date, neither preventive nor control methods have been implemented for managing this pest under field conditions. In this review, we highlight the most relevant information on the biology of P. archon and summarize the available control strategies with a special focus on biocontrol-based treatments.     

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.     

Increasing the availability of food and the reproduction ofPoecilus cupreus in a cereal field by strip-management

The spruce bud moth,Zeiraphera canadensis Mutt & Free. (Lepidoptera: Tortricidae), displayed a bimodal activity pattern in two young white spruce,Picea glauca (Moench) Voss, plantations. During the day, when temperatures were highest and relative humidities lowest, most moths remained in the lower crown, presumably in response to the risk of hygrothermal stress and/or predation. Flight activity peaks occurred near dawn and dusk. Moths moved higher in the crowns shortly after sunset and many were observed flying above the tree canopy. This activity peak was associated with oviposition and mating. Oviposition started at sunset and continued for approximately 3 h. Females probed oviposition sites with their ovipositor before ovipositing, suggesting that they use sensillae on the ovipositor to evaluate potential oviposition sites. Most mating occurred between 22∶00 and 04∶00 h. Males hovered adjacent to branches containing females before alighting. After alighting, males usually approached to within 2–5 cm of females and then remained motionless for a short time before attempting mounting, indicating that some short-range communication usually occurs prior to mating. Reduced flight activity between midnight and dawn was not due to low temperatures, which were above the minimal temperatures for flight for males (10°C) and females (12°C) determined in the laboratory.     

Reproductive behaviour of mated and virgin females of a solitary sawflyDineura virididorsata

Sex ratios of the arrhenotokous sawflyDineura virididorsata Retz. (Hymenoptera: Tenthredinidae) are variable among populations in the field, but are biased in favour of females. We have concluded elsewhere that the sex ratios of this species cannot be satisfactorily explained by any general sex ratio model that is available. Here we present data on the different behavioural traits that are relevant to interpretation of the evolution of female-biased sex ratios under local mate competition. We report on oviposition behaviour, mating behaviour and the patterns of movement of ovipositing virgin and mated females. Our results show that adult emergence is synchronous and females will mate daily with different males (in the laboratory, at least). This would decrease the relatedness of offspring should it also take place in the field. In addition, ovipositing females (mated and virgin) leave their site of release in a relatively short time and deposit few eggs relative to their capabilities, so siblings are unlikely to be clumped. The time that virgins spend in deposition of an egg is not different from that spent by mated females, so there is no differential ‘investment’ in this regard. These behavioural observations agree with our previous conclusion that the preconditions of current sex ratio theories are not met byD. virididorsata.     

Use of Radio Telemetry for Studying Flight Movements of <Emphasis Type="Italic">Paysandisia archon</Emphasis> (Lepidoptera: Castniidae)

The moth Paysandisia archon is an invasive species that infests palm trees in the Mediterranean Basin. The immature stages occur exclusively inside the palm crown, whereas adults fly to mate and locate a new host plant. Here, we describe the use of small radio transmitters (0.27 g) for tracing the movements of P. archon adults. We report the first successful use of radio telemetry to track flight distances and space use of a flying moth. Our study was carried out using 11 tagged moths released on the Maguelone Peninsula in southern France. Although the males were successfully tracked within a restricted area, estimated at 4 ha, most of the tagged females immediately flew distances over 500 m, disappearing beyond the maximum detection range of the 30 ha study area. Flights for mating, resting and, likely, oviposition covered distances of 11.6 m to 224 m in males and 16.8 m to >500 m in females. We found evidence that both sexes are active during the warmer temperatures of the day. In contrast, the moths were inactive when the relative humidity was high. Moreover, the P. archon moths do not seem to reside on the host palm trees. Our data show a high mobility of females, which may be responsible for the spread of the first recorded Castniidae in France by laying eggs far from the palm trees where they emerged.     

Larval life-styles and oviposition behaviour of parasites and grazers

Summary We apply a theory for the computation of reproductive success for insects whose offspring can move from the host on which they hatch to compute the expected reproductive success associated with two kinds of oviposition behaviour. In the first type, accept or reject (AR), the ovipositing female either accepts a host for oviposition and lays her entire egg complement or rejects the host for oviposition. In the second type, clutch optimization (CO), the ovipositing female adjusts clutch according to host properties. The strongest prediction of our theory is that parasites which are both pro-ovigenic (emerge with essentially a full complement of eggs) and use the AR behaviour will be very rare.     

Pupation site and emergence time influence the mating success of bagworm females,Oiketicus kirbyi

In commercial oil palm plantations in Costa Rica, we tested the hypotheses that pupation site and emergence time affect the mating success of protogynous female bagworms,Oiketicus kirbyi (Guilding) (Lepidoptera: Psychidae). Greater proportions of female than male pupae on upper leaves of oil palms and greater proportions of mated females in the upper rather than lower crown strata support the hypothesis that selection of pupation site by female larvae influences the mating success of adults. Increasing captures of males with increasing trap height further suggest that enhanced mating success of females in tree tops may be attributed either to most effective dissemination of sex pheromone on higher sites, or to males foraging predominantly in the upper strata of oil palms. As the majority of females pupated in the middle rather than upper crown of oil palms, selection of pupation site by females may be affected by additional as yet unknown factors. Emergence of females significntly preceded emergence of males. Increasing proportions of mated females throughout the emergence seasons probably resulted from an increased ‘availability’ of males. In tropical rainforests with local variations inO. kirbyi developmental time and stage, protogyny may represent an evolutionary strategy that furthers outbreeding.     

Semiochemicals from ovaries of gravid females attract ovipositing female houseflies,Musca domestica

Chemical signals originating from the ovaries of gravid females of Musca domestica (Diptera: Cyclorrhapha: Muscidae) attract ovipositing females to common egg-laying sites. Behavioral experiments indicated that females preferred to oviposit in fermented wheat bran containing ovaries from reproductively mature houseflies. Females preferred to oviposit in fermented wheat bran than wet wheat bran. This effect was additive with the attraction to housefly ovaries. Solvent extracts from housefly ovaries were attractive to gravid females. Extracts obtained with hexane were most attractive to gravid females for egg laying, and extracts obtained with ethyl acetate attracted more egg laying than extracts obtained by dichloromethane. Gas chromatography (GC) and gas chromatography coupled with mass spectrometry (GC-MS) analysis showed that tricosane and (Z)-9-tricosene were the main components of the hexane extracts. Both tricosane and (Z)-9-tricosene were shown to elicit dose-dependent aggregation of gravid females in oviposition bioassays, but high doses of either chemical were not attractive.     

家蝇卵巢中的信息化学物质引诱雌蝇产卵的研究

产卵期家蝇MuscadomesticaL .卵巢中的信息化学物质诱引家蝇产卵。行为试验表明 ,雌蝇产卵时优先选择被成熟卵巢污染过的发酵麦麸 ,诱集的产卵百分率依次为 :发酵麦麸 + 1对卵巢 >湿麦麸+ 1对卵巢 >发酵麦麸 >湿麦麸 ;卵巢提取物同样对产卵雌蝇有诱集活力。产卵雌蝇卵巢正己烷、二氯甲烷以及乙酸乙酯的提取物均能诱集雌蝇产卵 ,诱集的产卵百分率依次为 :雌蝇卵巢正己烷提取物 >乙酸乙酯提取物 >二氯甲烷提取物。推测家蝇卵巢中的信息化学物质可能含有家蝇产卵聚集信息素的组分。     

Flight and oviposition behaviour of the African stem borer,Busseola fusca,on various host plant species

The African stem borer, Busseola fusca (Fuller) (Lepidoptera: Noctuidae), is an important pest of maize and sorghum in sub‐Saharan Africa. As in many other lepidopteran insects, the ability of B. fusca to recognize and colonize a variety of plants is based on the interaction between its sensory systems and the physical and chemical characteristics of its immediate environment. In this study, we tried to identify the behavioural steps of B. fusca leading to host selection and oviposition. Three Poaceae species commonly cultivated in Kenya for human consumption and animal forage were used in this study: the two most preferred hosts, maize (Zea mays L.) and sorghum [Sorghum bicolor (L.) Moench], and one non‐preferred host, Napier grass (Pennisetum purpureum Schumach). Wind tunnel observations revealed that volatiles produced by the different plant species did not appear to strongly influence the general orientation of B. fusca towards the plant, as similar behavioural steps were exhibited by the female moth regardless of the plant species involved. This indicated that the females were not able to recognize their preferred hosts from a distance. After landing, the female typically swept her ovipositor on the plant surface, simultaneously touching it with the tips of her antennae, and then oviposited. This behaviour was more frequently observed on maize and sorghum than on Napier grass, and indicated that both antennal and ovipositor receptors are used by the female moths to evaluate the plant surface before deciding to oviposit. As a result, the females laid more eggs on the two crops than on Napier grass. We conclude therefore that females recognized their preferred hosts only after landing. Tactile and contact‐chemoreception stimuli from the plants seemed to play a major role in oviposition decisions of B. fusca.