Innate preference hierarchies coupled with adult experience,rather than larval imprinting or transgenerational acclimation,determine host plant use in Pieris rapae

The evolution of host range drives diversification in phytophagous insects, and understanding the female oviposition choices is pivotal for understanding host specialization. One controversial mechanism for female host choice is Hopkins’ host selection principle, where females are predicted to increase their preference for the host species they were feeding upon as larvae. A recent hypothesis posits that such larval imprinting is especially adaptive in combination with anticipatory transgenerational acclimation, so that females both allocate and adapt their offspring to their future host. We study the butterfly Pieris rapae, for which previous evidence suggests that females prefer to oviposit on host individuals of similar nitrogen content as the plant they were feeding upon as larvae, and where the offspring show higher performance on the mother's host type. We test the hypothesis that larval experience and anticipatory transgenerational effects influence female host plant acceptance (no‐choice) and preference (choice) of two host plant species (Barbarea vulgaris and Berteroa incana) of varying nitrogen content. We then test the offspring performance on these hosts. We found no evidence of larval imprinting affecting female decision‐making during oviposition, but that an adult female experience of egg laying in no‐choice trials on the less‐preferred host Be. incana slightly increased the P. rapae propensity to oviposit on Be. incana in subsequent choice trials. We found no transgenerational effects on female host acceptance or preference, but negative transgenerational effects on larval performance, because the offspring of P. rapae females that had developed on Be. incana as larvae grew slower on both hosts, and especially on Be. incana. Our results suggest that among host species, preferences are guided by hard‐wired preference hierarchies linked to species‐specific host traits and less affected by larval experience or transgenerational effects, which may be more important for females evaluating different host individuals of the same species.     

Myrmecophilous butterflies utilise ant–treehopper associations as visual cues for oviposition

1. Selection of a safe oviposition site is important for herbivorous insects whose immature stages have limited mobility. Female herbivores rely on environmental cues for this choice, and presence of natural enemies or mutualistic partners may be important in this process. 2. Some butterflies have mutualistic interactions with ants (myrmecophily), in which caterpillars offer a nutritional liquid and gain protection against natural enemies. Participants in butterfly–ant mutualisms may utilise signals to initiate interactions, but the use of visual cues by ovipositing myrmecophilous butterflies remains uncertain. 3. Larvae of facultatively myrmecophilous Parrhasius polibetes (Lycaenidae) feed on Schefflera vinosa, and females prefer to oviposit near aggregations of the ant‐tended treehopper Guayaquila xiphias, where caterpillars survive better due to increased ant attendance. Given the conspicuousness of ant–treehopper associations, it was investigated whether butterflies use them as visual cues for oviposition and, if so, which participants of the association are used as cues: ants, treehoppers, or both. 4. Experiments using dried insects on paired branches revealed that females visually recognise ants and ant–treehopper associations, using them for egg‐laying decisions. However, presence of a treehopper aggregation alone had no effect on oviposition choices. 5. This is a first insight into the importance of visual discrimination for ovipositing myrmecophilous butterflies. The results show that facultative mutualisms can be important enough to promote a behavioural adaptation (visual detection of ants) reinforcing the interaction. Our research highlights the importance of the behavioural interface within complex multispecies systems.     

Host plant preference and offspring performance of a leaf‐mining moth,Caloptilia fraxinella,on two Fraxinus species

The preference‐performance or ‘mother‐knows‐best’ hypothesis states that female insects choose to oviposit on a host plant that increases the performance of their offspring. This positive link between host plant choice and larval performance is especially important for leaf miners with non‐motile larvae that are entirely dependent upon the oviposition choice of the female for host plant location. Preference and performance of the ash leaf coneroller, Caloptilia fraxinella (Ely) (Lepidoptera: Gracillariidae), a specialist on ash trees, Fraxinus spp. (Oleaceae), were tested in a series of laboratory and field experiments. Female C. fraxinella were exposed to two closely related hosts, black ash, Fraxinus nigra Marshall, and green ash, Fraxinus pennsylvanica Marshall var. subintegerrima (Vahl), in oviposition choice and wind tunnel flight experiments to determine which host is most attractive for oviposition. Caloptilia fraxinella females were inconsistent in host choice, yet performance of larvae was greater on green than black ash. In preference studies, C. fraxinella preferred to oviposit on black ash when leaflets were removed from the tree, but preferred intact green ash over black ash seedlings for oviposition and host location in a wind tunnel. In the field, however, more C. fraxinella visited black ash var. ‘Fallgold’ at leaf flush than green ash at the same sites. Age of the ash leaflet also influences oviposition in this leaf miner and females preferred new over old leaflets for oviposition. Performance of C. fraxinella larvae was evaluated in field and laboratory experiments and was greater on green ash than on black ash in both experiments based on larval survival and development time parameters. The stronger oviposition and host location preference in the field for black ash were not linked to enhanced performance of offspring, as green ash was the superior host, supporting higher larval survival and faster development. A stronger host location preference in the wind tunnel for green ash over black ash, however, suggests that under certain circumstances with this moth species, ‘mother (may) know best’.     

Oviposition preference but not adult feeding preference matches with offspring performance in the bronze bug Thaumastocoris peregrinus

Optimal foraging and optimal oviposition are two major forces leading to plant selection by insect females, but the contribution of these forces to the host‐selection process has been little studied for sucking herbivores. We studied feeding and oviposition behavior of a global pest, the bronze bug, Thaumastocoris peregrinus Carpintero & Dellapé (Heteroptera: Thaumastocoridae), using dual‐choice bioassays to evaluate the preference of females between host species, developmental leaf stage, or prior plant exposure to conspecifics. We assessed the link between these preferences and the performance of the offspring, by comparing survival and developmental time of nymphs reared on the various treatments. Finally, we compared the composition of the leaf wax of healthy and damaged leaves, and tested the effects of leaf wax on female preference behavior. Using healthy adult leaves of Eucalyptus tereticornis Sm. (Myrtaceae) as a reference, we found that females prefer to feed on Eucalyptus grandis W. Hill ex Maiden and E. tereticornis adult leaves that had been previously damaged by female conspecifics, whereas they reject juvenile leaves of E. tereticornis as food. Females also prefer to oviposit on leaves previously damaged by conspecifics but they rejected E. grandis as oviposition substrate. Nymphal performance varied among leaf treatments, suggesting a correlation with oviposition preference (but not feeding preference). Epicuticular wax extracts from damaged leaves contained higher concentrations of long‐chain, saturated linear alkanes, aldehydes, and alcohols than extracts from undamaged leaves. However, a choice assay failed to demonstrate an oviposition preference based on leaf surface wax chemistry. We discuss these findings in the context of the preference‐performance relationship.     

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.     

Choice of oviposition sites by Manduca sexta and its consequences for egg and larval performance

The preference‐performance hypothesis predicts that female insects should prefer to lay eggs in locations that enhance offspring performance. This study examines the choices of females regarding where to oviposit within plants, focusing on the hawkmoth Manduca sexta L. (Lepidoptera: Sphingidae) and its host Datura wrightii Regel (Solanaceae) in the southwestern USA. Smaller Datura leaves provide cooler microclimates for eggs (which may lead to faster embryonic development, shortening their exposure to egg predators) and more nitrogen for larvae. In contrast, large leaves reach temperatures that are stressfully high (which slows embryonic development) and provide less nitrogen for larvae. Thus, we would expect females to oviposit on small leaves. To examine whether leaf size influences female preference and offspring performance, we used laboratory and field studies to address the following questions. (1) On what size leaves do females typically oviposit? (2) Does the distribution of eggs in nature differ from that expected by chance? And (3) how does leaf size affect survival or growth of eggs and larvae? We find that oviposition choices of females do not lead to the highest probability of offspring survival. Females lay eggs on larger leaves, likely due to the greater accessibility of those leaves; however, eggs are more likely to hatch on small leaves. Larvae grow faster on large leaves, but larvae are also surprisingly mobile, suggesting that the consequences of oviposition site are minor once eggs have hatched. Larval mobility was seen only in the field, not in the laboratory, emphasizing the importance of field studies for predicting real‐world performance. Although females' leaf choices are potentially risky for eggs, the threats of high temperature and predation may vary sufficiently in space and time that there is no consistent selection for strong preferences. Furthermore, the fitness consequences for eggs and larvae largely offset each other and offspring are sufficiently mobile to cope with the conditions where they are laid.     

The Life and Death of Hopkins' Host-Selection Principle

Hopkins' host-selection principle (HHSP) refers to the observation that many adult insects demonstrate a preference for the host species on which they themselves developed as larvae. The meaning of HHSP has changed significantly since its first proposal in 1916. This review considers how the meaning of HHSP has changed over time and considers the various mechanisms that could contribute to a behavioral bias for the developmental host. The assumption that HHSP implies that the behavior of adult insects is conditioned by larval experience has resulted in widespread condemnation of HHSP. Despite a great deal of attention, there is still very little convincing evidence for preimaginal conditioning of host choice in insects. But growing evidence indicates that genetic variation in behavior and conditioning during the life span of an adult insect can contribute to a preference for the host on which an insect developed. Insects can acquire adult oviposition or feeding preferences through exposure to chemical residues from the environment of earlier life history stages. The concepts of host races and host fidelity have become familiar and acceptable, while the association of HHSP with preimaginal conditioning has led to a general rejection of the term.     

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.     

Effect of arbuscular mycorrhizal fungi (Glomus intraradices) on the oviposition of rice water weevil (Lissorhoptrus oryzophilus)

Root-feeding insects are important drivers in ecosystems, and links between aboveground oviposition preference and belowground larval performance have been suggested. The root-colonizing arbuscular mycorrhizal fungi (AMF) play a central role in plant nutrition and are known to change host quality for root-feeding insects. However, it is not known if and how AMF affect the aboveground oviposition of insects whose offspring feed on roots. According to the preference–performance hypothesis, insect herbivores oviposit on plants that will maximize offspring performance. In a greenhouse experiment with rice (Oryza sativa), we investigated the effects of AMF (Glomus intraradices) on aboveground oviposition of rice water weevil (Lissorhoptrus oryzophilus), the larvae of which feed belowground on the roots. Oviposition (i.e., the numbers of eggs laid by weevil females in leaf sheaths) was enhanced when the plants were colonized by AMF. However, the leaf area consumed by adult weevils was not affected. Although AMF reduced plant biomass, it increased nitrogen (N) and phosphorus concentrations in leaves and N in roots. The results suggest that rice water weevil females are able to discriminate plants for oviposition depending on their mycorrhizal status. The discrimination is probably related to AMF-mediated changes in plant quality, i.e., the females choose to oviposit more on plants with higher nutrient concentrations to potentially optimize offspring performance. AMF-mediated change in plant host choice for chewing insect oviposition is a novel aspect of below- and aboveground interactions.     

Egg laying preferences and larval performance in Dermestes maculatus

Dermestes maculatus DeGeer (Coleoptera: Dermestidae) is both a pest of dried stored products and, through its colonization of carrion, a forensically important species. However, little is known about the consequences of female oviposition site preferences on larval growth and development. To examine this, non‐virgin female beetles were offered a choice of food resources that had been aged to various extents to explore the adaptive nature of female oviposition preferences. Dermestes maculatus females consistently preferred to oviposit on muscle in contrast to either fat or bone marrow. Constraining larvae onto one of the three resource types confirmed that larvae grew faster and eclosed into larger adults when fed on muscle than when fed on either fat or bone marrow. In addition, the degree of sexual dimorphism was also related to food resource, with the greatest extent of size dimorphism (females larger than males) being evident on the preferred muscle resource. This conforms to the view that intraspecific variation in sexual size dimorphism is driven by intersexual differences in phenotypic plasticity, with females being able to reach greater size than males when conditions are good. The results indicate that D. maculatus female oviposition preferences are adaptive in that adult oviposition choice can enhance offspring fitness and so broadly conforms to the oviposition preference‐larval performance hypothesis as noted in a number of phytophagous insects.     

Selection of large host plants for oviposition by a monophagous leaf beetle: nutritional quality or enemy‐free space?

Abstract.  1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants.
2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival.
3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants.
4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field.
5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories.
6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.     

Continuous rearing of the predatory anthocorid Orius laevigatus without plant materials

The omnivorous anthocorid predator Orius laevigatus is an economically important biological control agent of several small arthropod pests including the western flower thrips Frankliniella occidentalis. Mass rearing systems for Orius bugs typically make use of plant materials such as bean pods as an oviposition substrate and moisture source. Omission of plant materials from the mass rearing system of these beneficial arthropods could drastically improve the cost‐effectiveness of their production and thus stimulate their use in augmentative biological control. This study investigated the effect of a plantless rearing system, using wax paper as a walking substrate, water encapsulated in Parafilm domes, and an artificial oviposition substrate made of Parafilm and moist cotton wool, on the developmental and reproductive fitness of O. laevigatus. Plantless rearing during four generations resulted in females with an 11% lower body weight and a pre‐oviposition period that was prolonged by 29%. However, other biological parameters were not negatively affected by the absence of plants. In addition, plantless‐reared females had similar predation rates on F. occidentalis larvae as their peers maintained on plants. Our study indicates that the omission of plant material from the production cycle of O. laevigatus is possible when Ephestia kuehniella eggs are provided as a nutritionally optimal food source.     

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.     

Host‐Plant Choice Behavior at Multiple Life‐Cycle Stages: The Roles of Mobility and Early Growth in Decision‐Making

The choice of food plants often assumes critical importance for a herbivore. Although many studies have investigated host‐plant choice behavior, few have examined preferences (vs. growth and survival) at multiple stages of the life cycle, notwithstanding the importance of identifying the critical stage(s) in an animal’s life history. Fern moths Herpetogramma theseusalis (Lepidoptera: Crambidae) provide an excellent opportunity to test host‐plant choice at several stages. Fern moth larvae feed on distantly related ferns, sensitive Onoclea sensibilis and marsh fern Thelypteris palustris, and adults oviposit on both species. We examined newly hatched larvae, overwintered larvae and ovipositing females to test hypotheses predicting when host‐plant choice takes place (overwintering and mobility hypotheses: overwintering stage determines choice of substrate vs. most mobile stage chooses) and the basis for choice (optimal oviposition and enemy‐free space hypotheses: resource producing highest fecundity vs. lowest losses to enemies). We also evaluated the hypothesis that host‐associated fitness trade‐offs explain host specialization. Only ovipositing females, the most mobile stage, exhibited a clear preference (for marsh fern), consistent with the mobility hypothesis. However, their preference for marsh fern fits neither the optimal oviposition hypothesis nor the enemy‐free space hypothesis; although some larvae initially grew faster on marsh fern, adults reared from the two ferns did not differ significantly in mass and experienced marginally lower parasitism on sensitive fern. Thus, we found no host‐associated fitness trade‐offs. Overwintering losses in marsh fern plots exceeded those in sensitive fern, and mixed plots supported the most overwintered larvae. Preference for marsh fern suggests that early success drives host‐plant choice, an advantage that later disappears. Temporal variability may prevent closer fits to the hypotheses, because both ferns provide the moths with acceptable resources throughout their life cycles.     

Cascading trait‐mediated interactions induced by ant pheromones

Trait‐mediated indirect interactions (TMII) can be as important as density‐mediated indirect interactions. Here, we provide evidence for a novel trait‐mediated cascade (where one TMII affects another TMII) and demonstrate that the mechanism consists of a predator eavesdropping on chemical signaling. Ants protect scale insects from predation by adult coccinellid beetles – the first TMII. However, parasitic phorid flies reduce ant foraging activity by 50% – the second TMII, providing a window of opportunity for female beetles to oviposit in high‐quality microsites. Beetle larvae are protected from ant predation and benefit from living in patches with high scale densities. We demonstrate that female beetles can detect pheromones released by the ant when attacked by phorids, and that only females, and especially gravid females, are attracted to the ant pheromone. As ants reduce their movement when under attack by phorids, we conclude that phorids facilitate beetle oviposition, thus producing the TMII cascade.     

Measuring the discrepancy between fecundity and lifetime reproductive success in a pollinating fig wasp

Lifetime reproductive success in female insects is often egg‐ or time‐limited. For instance in pro‐ovigenic species, when oviposition sites are abundant, females may quickly become devoid of eggs. Conversely, in the absence of suitable oviposition sites, females may die before laying all of their eggs. In pollinating fig wasps (Hymenoptera: Agaonidae), each species has an obligate mutualism with its host fig tree species [Ficus spp. (Moraceae)]. These pro‐ovigenic wasps oviposit in individual ovaries within the inflorescences of monoecious Ficus (syconia, or ‘figs’), which contain many flowers. Each female flower can thus become a seed or be converted into a wasp gall. The mystery is that the wasps never oviposit in all fig ovaries, even when a fig contains enough wasp females with enough eggs to do so. The failure of all wasps to translate all of their eggs into offspring clearly contributes to mutualism persistence, but the underlying causal mechanisms are unclear. We found in an undescribed Brazilian Pegoscapus wasp population that the lifetime reproductive success of lone foundresses was relatively unaffected by constraints on oviposition. The number of offspring produced by lone foundresses experimentally introduced into receptive figs was generally lower than the numbers of eggs carried, despite the fact that the wasps were able to lay all or most of their eggs. Because we excluded any effects of intraspecific competitors and parasitic non‐pollinating wasps, our data suggest that some pollinators produce few offspring because some of their eggs or larvae are unviable or are victims of plant defences.     

Trichobaris weevils distinguish amongst toxic host plants by sensing volatiles that do not affect larval performance

Herbivorous insects use plant metabolites to inform their host plant selection for oviposition. These host‐selection behaviours are often consistent with the preference–performance hypothesis; females oviposit on hosts that maximize the performance of their offspring. However, the metabolites used for these oviposition choices and those responsible for differences in offspring performance remain unknown for ecologically relevant interactions. Here, we examined the host‐selection behaviours of two sympatric weevils, the Datura (Trichobaris compacta) and tobacco (T. mucorea) weevils in field and glasshouse experiments with transgenic host plants specifically altered in different components of their secondary metabolism. Adult females of both species strongly preferred to feed on D. wrightii rather than on N. attenuata leaves, but T. mucorea preferred to oviposit on N. attenuata, while T. compacta oviposited only on D. wrightii. These oviposition behaviours increased offspring performance: T. compacta larvae only survived in D. wrightii stems and T. mucorea larvae survived better in N. attenuata than in D. wrightii stems. Choice assays with nicotine‐free, JA‐impaired, and sesquiterpene‐over‐produced isogenic N. attenuata plants revealed that although half of the T. compacta larvae survived in nicotine‐free N. attenuata lines, nicotine did not influence the oviposition behaviours of both the nicotine‐adapted and nicotine‐sensitive species. JA‐induced sesquiterpene volatiles are key compounds influencing T. mucorea females’ oviposition choices, but these sesquiterpenes had no effect on larval performance. We conclude that adult females are able to choose the best host plant for their offspring and use chemicals different from those that influence larval performance to inform their oviposition decisions.     

Predator related oviposition site selection of aquatic beetles (Hydroporus spp.) and effects on offspring life-history

1. Theory predicts that natural selection should favour females that are able to correctly assess the risk of predation and then use that information to avoid high‐risk oviposition sites to reduce the risk of offspring predation. Despite the potential significance of such behaviour on individual fitness, population dynamics and community structure, relatively few studies of oviposition behaviour connected to the risk of predation have been carried out. 2. However, some recent studies suggest that oviposition site selection in response to risk of predation may be a common phenomenon, at least among amphibians and mosquitoes. A vast majority of previous studies have, however, neglected to investigate how the offspring are affected, in terms of fitness related parameters, by the maternal oviposition site choice. 3. In an outdoor artificial pond experiment we tested the oviposition site selection of female aquatic beetles (Hydroporus spp.) in relation to the presence or absence of a predatory fish (Perca fluviatilis). In addition, we monitored how the oviposition site selection affected the behaviour, growth and food resource of the progeny. 4. We show that free‐flying females of the aquatic beetles Hydroporus incognitus and H. nigrita prefer to oviposit in waters without fish compared with waters with fish. Larval activity of Hydroporus spp. was unaffected by fish presence. Our results indicate that beetle larvae from females that do lay eggs in waters with fish show increased growth compared with larvae in waters without fish. We explain this difference in growth by a higher per‐capita food supply in the presence of a fish predator. This finding may have important implications for our understanding of how the variance of oviposition site selection in a population is sustained.     

The Immediate Source of the Oviposition-Deterring Pheromone Produced by Larvae of Adalia bipunctata (L.) (Coleoptera, Coccinellidae)

As is the case for other insects ovipositing on or in resources that are limited in time and/or space, the two-spot ladybird beetle, Adalia bipunctata (L.) produces an oviposition-deterring pheromone (ODP), which is produced by the larval stages. Foraging larvae touch the substrate with their tarsi and the anal disk on the tenth abdominal segment. The aim of this paper was to determine whether the ODP produced by larvae was deposited by the tarsi or the anal disk. Fourth instar larvae either had their anal disk and tarsi, or anal disk, or tarsi coated with a water-soluble mounting medium. Larvae so treated were allowed to walk on filter paper that was subsequently presented to gravid females. The tracks of larvae that had both their tarsi and anal disk masked did not inhibit oviposition. However, the tracks of larvae that had only their tarsi masked significantly inhibited oviposition but those of larvae that had only their anal disk masked did not. It is concluded that the ODP is deposited on the substrate by the anal disk on the tenth abdominal segment of larvae.     

Initial preference for plant species and state during oviposition site selection by an odonate

Endophytic egg‐laying odonates use an ovipositor to insert their eggs inside plant tissues. Before egg deposition, oviposition site selection consists of two crucial steps: (i) the initial choice, typically decided in species that oviposit in tandem within vertical substrates by the male when landing and then by the female by staying on the substrate or flying to another; and (ii) the insertion site choice, made by the female who uses her ovipositor to palpate the substrate. Some odonates prefer to deposit their eggs within specific plant species. Some are able to discriminate between living and dead substrates during the initial choice. However, the extent to which odonates discriminate among distinct plant species during the initial choice is unknown. We studied the initial site preference in Lestes macrostigma (Odonata: Lestidae) to determine whether the males and/or females show a distinct preference among five types of shoots when landing on or when palpating the substrate, respectively. Male L. macrostigma preferred to land on Bolboschoenus maritimus and dead Juncus spp. When focusing on J. maritimus, females preferentially palpated the substrate when the male landed on dead shoots. We suggest that the male preference for these substrates is consistent with that of the female during insertion site choice but also during egg deposition. Such behavior should reduce the duration of oviposition, with benefits of reducing the predation risk and increasing available time for foraging. The advantage in preferring these substrates should be linked to a selection pressure acting on egg development and/or survival.