Oviposition behaviour and larval development of Anastrepha fraterculus from Argentina in citrus

Citrus peel physicochemical attributes are considered the main components conferring partial or even total resistance to fruit fly (Diptera: Tephritidae) infestation. Fruit fly females adapt their ovipositional strategies to overcome such resistance. Here, we explored the effects of citrus species (Rutaceae) on the ovipositional behaviour of the South American fruit fly, Anastrepha fraterculus (Wiedemann), and on its immature development. Particularly, we investigated the effects of (1) citrus species on oviposition behaviour and immature development, (2) citrus species on oviposition preference and on the location of the eggs at different depth in the citrus peel, and (3) harvest season and post‐harvest storage time on oviposition behaviour and immature development in lemon. Citrus species influenced ovipositional behaviour and affected survival of immature stages. Females laid eggs in lemon [Citrus limon (L.) Burm.], orange [Citrus sinensis (L.) Osbeck], and grapefruit (Citrus paradisi Macfadyen). In orange and lemon, larvae were found dead close to the oviposition areas, suggesting chemically mediated resistance mechanisms. Under choice conditions, females preferred grapefruit over lemon and bigger clutches were found in the layers where embryonic development is favoured. Unsuitability of lemon as a medium to complete development was neither affected by harvest season nor by storage time of the fruit after harvest. The physical and chemical characteristics of the peel were distinctive to each citrus species and may have affected the specific levels of resistance of these citrus species to infestation by A. fraterculus.     

An evolutionary interpretation of the “motivation to oviposit”

The ovipositional behavior of parasitoids and other insects is often described by phrases such as “motivation to oviposit” or “ovipositional drive”. This paper shows how an evolutionary (i.e. functional) interpretation can be given to such phrases. A detailed model for the parasitisation of Sycamore aphids by M. pseudoplatani is developed, using experiments by Collins and Dixon (1986). Two models are developed: i) one in which egg complement is the only state variable and ii) one in which egg complement and information concerning host densities are state variables. Comparisons of the behaviour of simulated parasitoids, using the decisions associated with the models, and the experiments suggest that both egg complement and information are important for the parasitoid's decision making. Accepting previously parasitized hosts may be optimal, and not simply an error in parasitoid perception. A number of other detailed predictions are made, such as the relative fitness of first and second eggs in superparasitized hosts and the nature of the memory of the parasitoid.     

Studies of Aedes aegypti (Diptera: Culicidae) ovipositional responses to newly identified semiochemicals from conspecific eggs

Abstract  The chemical factors influencing the selection of oviposition site by gravid females of various mosquito species have been the subject of numerous investigations. Recent studies have revealed this behaviour to be controlled by semiochemicals. Here we report studies on semiochemicals of egg origin and their effect on the ovipositional behaviour of Aedes aegypti. The compounds present in egg extracts of Ae. aegypti mosquitoes were isolated and identified by gas chromatography-mass spectrometry. They were then evaluated for their effect on ovipositional behaviour against gravid females of Ae. aegypti mosquitoes at different concentrations. Gravid female Ae. aegypti were found to be sensitive to all the identified compounds: 6-hexanolactone, methyl dodecanoate, dodecanoic acid, methyl tetradecanoate, tetradecanoic acid, methyl (Z)-9-hexadecenoate, methyl hexadecanoate (Z)-9-hexadecenoic acid, hexadecanoic acid, methyl (Z)-9-octadecenoate, methyl octadecanoate (Z)-9-octadecenoic acid and octadecanoic acid. Among them, dodecanoic and (Z)-9-hexadecenoic acids showed significant positive ovipositional response at different concentrations whereas all the esters showed deterrent/repellent ovipositional effect.     

Functions of the spermathecal muscle of the boll weevil,Anthonomus grandis

In female boll weevils, Anthonomus grandis, spermathecal filling was not affected by severing the spermathecal muscles. Females whose spermathecal muscles were severed 2 to 4 weeks after mating laid infertile eggs and resumed virginal ovipositional behaviour indicating the importance of this muscle in supplying sperm for egg fertilization. The presence of normal active sperm within the spermatheca in no way influenced ovipositional behaviour. In females whose spermathecal muscles had been severed, 22 per cent sperm displacement occurred after a second mating compared with 66 per cent for normal females. The physical displacement of sperm was thus largely dependent on a functional spermathecal muscle.     

Variance and covariance of ovipositional rates and developmental rates in the Phytoseidae (Acari: Mesostigmata): a phylogenetic consideration

Variance and covariance of ovipositional rates and developmental rates in the Phytoseiidae were analysed using comparative methods which consider phylogenetic effects. Nested analysis of variance showed that mean ovipositional rates and developmental rates of phytoseiid mites varied significantly between subfamilies, among genera within subfamilies, among subgenera within genera and among species within subgenera. For example, the mean ovipositional rate (eggs per day) was higher in the Amblyseiinae (2.04) than in the Phytoseiinae (1.39) and within the Amblyseiinae, it was higher in the genus Phytoseiulus (2.66) than in Amblyseius (1.80). Regressions using mean values of subgenera or higher taxa to account for phylogenetic effects showed significant correlation between mean ovipositional rates and developmental rates. The implications of this analysis for selecting species for future comparative analysis of phytoseiid life history variation are discussed.     

Contrary choices: possible exploitation of enemy-free space by herbivorous insects in cultivated vs. wild crucifers

Summary The pressure to escape natural enemies may shape how herbivorous insects use their plant resources. On wild crucifers, ovipositional preferences of the diamondback moth (Plutella xylostella; DBM) were similar to searching preferences of its main parasitoid, an ichneumonid wasp (Diadegma insulare). But on cultivated crucifers, these species had opposite preferences. In addition, DBM ovipositional preferences did not correlate with growth or reproduction on several foodplants. We interpret these patterns as evidence of evolution for use of enemy-free space in agricultural systems.     

Activity patterns and oviposition rates of Aphytis lingnanensis females,a parasitoid of California red scale Aonidiella aurantii: implications for successful biological control

1. Activity patterns of Aphytis lingnanensis Compere (Hymenoptera: Aphelinidae) females were determined. Females have a preovipositional period of 2–6.5 h and oviposit daily until they die. The average lifetime fecundity was 191 eggs (n = 10 females), with a maximum of 13 eggs per day, but usually far fewer. Females do not lay eggs at night as low light levels reduce the propensity to oviposit. 2. Aphytis lingnanensis females showed a repeatable pattern of activity during the course of the day, ovipositing actively during the first 1.5 h of their 7 h exposure to hosts. Thereafter, activity levels decreased sharply and females spent relatively long intervals resting. Activity occurred in fairly discrete bouts, with long periods of inactivity, ranging from 12 to 231 min between bouts. 3. Each day, females laid most eggs during their first bout of oviposition, with availability of mature eggs in the ovaries dictating the duration and intensity of ovipositional bouts. Once females completed a bout of oviposition in the laboratory, they moved away from the hosts even though many suitable hosts remained. They stood immobile, sheltering, until they matured more eggs. 4. These results suggest how general biological control models and dynamic state variable models of behaviour can be made more realistic. In particular, the behaviour of females does not remain constant over extended periods, particularly because of egg depletion, which suggests that the interaction between the ovipositional history of the individual and its physiological capabilities dictates the response of females to particular circumstances, and thus contributes to patterns of parasitism in the field. 5. It is predicted that A. lingnanensis females in the field are inactive at night and begin ovipositing when it becomes light enough in the morning. Egg depletion is likely if enough hosts are available, but should occur later in the day than was recorded in the laboratory. Egg-depleted females are likely to shelter while they mature more eggs, but may undertake interpatch movement.     

Intraspecific variability in host plant quality and ovipositionaI preferences in Eucheira socialis (Lepidoptera: Pieridae)

Abstract.

• 1 This study investigates interactions between Eucheira socialis (Pieridae: Lepidoptera), a strict monophagous herbivore, on Arbutus xalapensis (Ericaceae), a host plant with few herbivores. This tight association of insect on plant has many attributes conducive to reciprocal rather than diffuse evolution.
• 2 An indirect way of testing plant–insect coevolutionary theories is to test for the necessary conditions for reciprocal evolution in ecological time. Two conditions for coevolution were studied: (1) host plants vary in their suitability for larval growth and development, and (2) ovipositing insects discriminate among these plants based on their relative suitability.
• 3 Large differences in host plant suitability were found and relative differences were consistent from year to year.
• 4 There was no evidence that female insects based their ovipositional decisions on relative tree quality, which implies that factors other than host plant quality are involved in the maintenance and evolution of oviposition behaviour in Eucheira.
• 5 Of seven factors known to influence ovipositional preferences of insects among plants independent of potential larval success, the most likely causal factor in this system is the ability of females to balance a time/energy budget for finding potential oviposition sites, discriminating among them, and actually ovipositing.

     

Oviposition site selection on oats: the effect of plant architecture,plant and leaf age,tissue toughness,and hardness on cereal leaf beetle,Oulema melanopus

In sequentially planted oat stands, the cereal leaf beetle (CLB), Oulema melanopus (L.) (Coleoptera: Chrysomelidae: Lemini), is found in greater numbers, and lays more eggs, on later planted (younger) oats (Avena sativa L.) (Poaceae). Plant characteristics that could explain this ovipositional preference were examined in a series of experiments. Cage and open field whole plant preference tests confirmed the attraction of ovipositing females to younger oats. A cage effect illustrated the role of plant architecture (plant height) in CLB host selection. Two multiple‐choice and one no‐choice excised leaf experiments determined that characteristics of individual leaves associated with leaf insertion level (leaf number from base to apex) and age influence ovipositional site selection. Leaves of higher insertion level have higher nitrogen content, but fewer eggs are laid on those leaves. Two experiments examining the interaction between total leaf nitrogen and leaf insertion level showed that only leaf insertion level affected oviposition choice. Published literature suggests variation in secondary plant compounds cannot explain O. melanopus ovipositional preference among leaves. Grass leaves of higher insertion level have more extensively developed cells associated with tissue toughness and hardness. The data and supporting literature suggest tissue toughness and hardness are deterring oviposition on oat leaves of higher insertion level. However, newly eclosed larvae are able to feed on leaves usually avoided as oviposition sites. The explanation for this result may be a lack of correlation between host suitability and ovipositional preference.     

Adaptation for oviposition into concealed cycad ovules in the cycad weevils Antliarhinus zamiae and A. signatus (Goleoptera: Gurculionoidea)

Antliarhinus zamiae (Thunberg) and A. signatus Gyllenhal are the only insects known in which the larvae feed exclusively on the cycad gametophyte. In their host plants, species of the cycad genus Encephalartos (Zamiaceae), the gametophyte is protected by a three-layered integument and substantial sporophyll tissues. Adaptation for oviposition into the gametophyte may, therefore, be crucial to this feeding specialization. Ovipositional behaviour and ovipositor structure in four species of Antliarhinus suggest that ovipositional traits suited to ovule predation preceded the evolution of this habit and that ovipositional traits in ancestral species provided the right environment for the evolution of ovule predation. Current ovipositional traits in A. zamiae and A. signatus probably represent responses to changes in cone structure after the evolution of ovule predation.     

A STUDY ON THE OVIPOSITION SYNERGISTS FOR IN VITRO REARING TRICHOGRAMMA SPP.*

Abstract Solutions or suspensions of polyvinyl alcohol, gelatin, white latex, agar cassava powder, rice flour and starch were used as synergists to improve the oviposition of Trichogramma dendrolimi, T. confusum and T. evanescens with in vitro rearing method. The ovipositional response of Trichogramma spp. to these synergists was different. All the 7 preparations improved the trichogrammatid oviposition much on the artificial host “eggs”. Specific difference of trichogrammatid ovipositional response to these synergists was observed. So, the most effective and economical synergists can be used for increasing the efficiency of rearing different species of Trichogramma in vitro.     

Salinity and shade preferences result in ovipositional differences between sympatric tiger beetle species

1. Adult tiger beetles of the genus Cicindela often co‐occur within a habitat but larvae do not. Larvae are sedentary and form usually permanent burrows at the site of oviposition where they require 1–3 years for development. 2. To test niche partitioning based on ovipositional preference, the behaviour of two sympatric salt marsh tiger beetles, Cicindela circumpicta and C. togata (Coleoptera: Cicindelidae), were examined. 3. In laboratory studies, female C. circumpicta and C. togata distinguished between experimental salinities, with the former preferring 4 parts per thousand (ppt) and the latter preferring 12 ppt. In the field, C. circumpicta larvae were associated with lower salinities (1–3 ppt) and vegetation along the edges of salt flats while C. togata larvae were found on open salt flats often near halophytes (average salinity = 7.8 ppt). 4. In the field, females chose sites for oviposition in response to shade but not vertical landmarks. In a direct test, 53 of 56 new larval burrows occurred in shaded treatments, three in nonshaded controls. 5. Compared with nonshaded controls, shade increased survival of eggs that were collected in the laboratory and placed in the field. For C. circumpicta eggs, 78% placed in shaded treatments hatched, while significantly fewer (22%) hatched in exposed treatments. For C. togata, 43% of eggs placed in the shade hatched, while no eggs placed in exposed treatments hatched. 6. These results support the hypothesis of niche partitioning between C. circumpicta and C. togata based on ovipositional choice and resulting larval habitat.     

Symbiotic bacteria attenuate Drosophila oviposition repellence to alkaline through acidification

Metazoans harbor a wealth of symbionts that are ever-changing the environment by taking up resources and/or excreting metabolites.One such common environmental modification is a change in pH.Conventional wisdom holds that symbionts facilitate the survival and production of their hosts in the wild,but this notion lacks empirical evidence.Here,we report that symbiotic bacteria in the genus Enterococcus attenuate the oviposition avoidance of alkaline environments in Drosophila.We studied the effects of alkalinity on oviposition preference for the first time,and found that flies are robustly disinclined to oviposit on alkali-containing substrates.This innate repulsion to alkaline environments is explained,in part,by the fact that alkalinity compromises the health and lifespan of both offspring and parent Drosophila.Enterococcus dramatically diminished or even completely reversed the ovipositional avoidance of alkalinity in Drosophila.Mechanistically,Enterococcus generate abundant lactate during fermentation,which neutralizes the residual alkali in an egg-laying substrate.In conclusion,Enterococcus protects Drosophila from alkali stress by acidifying the ovipositional substrate,and ultimately improves the fitness of the Drosophila population.Our results demonstrate that symbionts are profound factors in the Drosophila ovipositional decision,and extend our understanding of the intimate interactions between Drosophila and their symbionts.     

Assessment of aphid colonies by hoverflies. III Pea aphids andEpisyrphus balteatus (de Geer) (Diptera: Syrphidae)

The syrphid flyEpisyrphus balteatus (de Geer) migrating from maple trees to leguminous plants in mid-April, assesses the qualitative and quantitative value of pea aphid colonies for securing successful development of her offspring. She selects young and “promising” pea aphid colonies of small size. This generally agrees with the “buy-futures” ovipositional tactic the syrphid fly adopts when it utilizes the maple aphids. The advantage of a “buy-futures” ovipositional tactic found in many syrphid species is discussed in comparison with other aphidophagous insects.     

Evaluation of Vigna angularis (Willd.) Ohwi and Ohashi accessions for resistance to Tetranychus urticae Koch (Acari:Tetranychidae)

Under laboratory conditions 88 azuki bean and three rice bean accessions were evaluated with a leaf-disc technique to select for those which expressed an antibiotic response to the TSSM ovipositional rate. Nine of the most promising accessions determined in the screening and Erimo, a commercial cultivar used as a control, were re-evaluated to determine the ovipositional rate and fecundity. Statistically significant differences between accessions were found: some of the accessions exhibited a reduced TSSM ovipositional rate and fecundity when compared with the susceptible control. The accessions ADZU279 and ADZU514 from Japan and ADZU577 from Ivory Coast demonstrated strong inhibition to TSSM fecundity. These three accessions may possess inherent potential in a breeding programme designed to enhance resistance to T. urticae.     

Status-dependent female mimicry in landlocked red-spotted masu salmon

We observed transient status-dependent female mimicry related to reproductive behaviour in landlocked red-spotted masu salmon (Oncorhynchus masou ishikawae). In females, the lower half of the lateral flank underwent solid blackening as an ovipositional character, and 70% of subordinate males showed a similar lateral pattern (M-type). In contrast, most of the paired males and 30% of subordinate males showed a different lateral pattern in which a melanic stripe ran along the lateral-line organ (N-type). It was possible to bring an M-type pseudo-fish model into closer proximity to a pair than an N-type model. In addition, the rate of successful sperm release in subordinate M-type males was higher than that in subordinate N-type males. Males could transform between N- and M-types according to fluctuations in competitive status among the males. A digital video image related to the article is available at .     

Ovipositional host choice by an invader accelerates displacement of its indigenous competitor

While competition for resources leading to invasion success is well recognized, avoidance of competition is much less so. Changes in behaviour that lead to avoidance are usually displayed by the weaker competitor. In our case though it was the stronger competitor that changed its behaviour by choosing a host for oviposition that was unacceptable to its competitor and avoiding one that was mutually acceptable; this accelerated the displacement of the competitor. We showed this by enclosing both invader (Middle East Asia Minor 1, B biotype) and indigenous (Australia, AN biotype) members of Bemisia tabaci species complex (sap sucking insects known as whiteflies) into field cages with either two plants that were a mutually acceptable oviposition host or one host that was acceptable to both and the other acceptable to the invader only. When only the mutually acceptable host was available, invader and indigenous females oviposited equally across the two plants. However, when given the choice, adult invaders still distributed themselves evenly across both hosts, but shifted their oviposition away from the mutually acceptable host and instead laid mostly on the host poorly utilized by the indigenous competitor. This indicates that the invader can change ovipositional choice to escape into competition free space.     

Ovipositional preference ofSiphoninus phillyreae and its fitness on seven host plant species

The relationship between ovipositional preference ofSiphoninus phillyreae (Haliday) (Homoptera: Aleyrodidae) and host plant suitability on seven host plant species (Citrus sinensis (L.) cv. ‘Washington’ [navel orange],Fraxinus uhdei (Wenz.) [shamel ash],Heteromeles arbutifolia Roemer [toyon],Malus domestica Mill. cv. ‘Granny Smith’, [apple],Pistacia vera L. cv. ‘Kerman’ [pistachio],Prunus persica (L.) cv. ‘O’Henry’ [peach], andPyrus communis L. cv. ‘Bartlett’ [pear]) was evaluated. Ovipositional preference ofS. phillyreae was determined by measuring egg density after adult female whitefies were given a simultaneous choice of all host plants for oviposition. Immature survival, developmental time, and adult size were examined to determine host plant suitability forS. phillyreae. All studies were performed under greenhouse conditions.S. phillyreae showed distinct ovipositional preference among host plant species. Host plant species had a significant effect on immature survival, but little or no effect on developmental time or forewing length. For four of the seven host plant species tested, there was an association between ovipositional preference and survival.     

Influence of larval experience on preference of a subterranean insect Delia antiqua on Allium hosts

Numerous studies have reported that larval experience can affect subsequent host plants selection and future oviposition decisions of many different species, but the investigation of pre‐imaginal experiences on host preference of adults has rarely been tested for soil‐dwelling insects. In this study, we present evidence that larval feeding experience can affect adult host preference in the onion maggot, Delia antiqua. By rearing D. antiqua on different host plants, we were able to examine the role of the natal host of different generations and the effect of larval density on host‐choice behaviour. We also performed bioassays by means of switched host treatment to evaluate the host‐selection principle. Choice bioassays among the three host species demonstrated that D. antiqua females preferred to oviposit on their natal host in each generation and host‐switching treatments. Additionally, increasing larval density could intensify this ovipositional preference on the natal host. The overall results showed that host preference of female D. antiqua is determined by larval experience and density. These findings also add support for the controversial Hopkins’ host‐selection principle.     

Host selection and sex ratio in a heteronomous hyperparasitoid

Abstract.

• 1 Encarsia tricolor Förster is a heteronomous hyperparasitoid: females develop as primary endoparasitoids of Homoptera, whereas males develop hyperparasitically in primary endoparasitoids, including conspecific females.
• 2 When offered pupae of Encarsia inaron (Walker) or conspecific pupae, E.tricolor showed a distinct preference to exploit E.inaron for male production.
• 3 Parasitoids were given the opportunity to gain different types of ovipositional experience on male and female hosts and were then offered patches containing mixtures of male and female hosts in various ratios. The mean sex ratio which they laid was not dependent on the relative availability of each type of host, whereas prior ovipositional experience had a significant effect.
• 4 Adaptive reasons for the observed oviposition strategies are suggested.