The influence of leaf age on the oviposition preference of Chrysophtharta bimaculata (Olivier) and the establishment of neonates

1 The degree of discrimination shown by a herbivore when selecting oviposition sites has been suggested as a key factor to understanding herbivore population dynamics. Chrysophtharta bimaculata (Coleoptera: Chrysomelidae) is a primary pest of Tasmanian eucalypt forests and can cause severe defoliation. Previous work suggests that females show discrimination when selecting oviposition sites. Our aim was to test the degree of oviposition discrimination exhibited by C. bimaculata with regards to leaf toughness, a character that is critical to neonate survival. 2 We conducted an experiment examining the leaf toughness critical for neonate survival and found that significant larval mortality occurs above a toughness of 46.9 g. We also determined that the maximum toughness of leaves upon which larvae established in the field was 48.2 g, supporting the laboratory result. 3 Field surveys showed that although the majority of eggs were laid on leaves suitable for larval establishment, many eggs were laid on unsuitable, tougher leaves. However, all eggs were normally placed within 20 cm of suitable leaves and glasshouse trials demonstrated the neonates could move this distance without mortality occurring. 4 We conclude that egg batch distribution and larval performance of C. bimaculata will influence the population dynamics of C. bimaculata in two ways. Firstly, the availability of expanding/newly expanding leaves of eucalypt hosts will determine larval carrying capacity. Secondly, at a more localized level, the deposition of large numbers of egg batches on both suitable and unsuitable leaves followed by successful neonate migration increases the risk of resource depletion and poor larval development.     

Oviposition preference of Anthocoris nemoralis and A. nemorum (Heteroptera: Anthocoridae) on pear leaves affected by leaf damage, honeydew and prey

Anthocorids are important predators of insect pests in pome fruit. Their choice of oviposition site determines the later distribution of nymphs. In two-choice experiments it was tested whether A. nemorum and A. nemoralis would show oviposition preference with regard to simulated insect damage, mimicked by piercing leaves with a fine pin, and whether the oviposition preference of A. nemoralis was affected by the presence of honeydew or eggs of their prey C. pyri. Leaves with simulated damage were preferred by A. nemoralis, but this was not the case for A. nemorum. Honeydew-treated leaves attracted more oviposition than honeydew-free leaves. On honeydew-treated leaves significantly more eggs were laid on the surface where honeydew had been applied, rather than the opposite. When C. pyri eggs were placed along the abaxial midvein, prey infestation attracted more oviposition. On the infested leaf more eggs were laid near the prey along the abaxial midvein than along the adaxial midvein. In contrast, when prey was placed along the adaxial midvein, no overall preference was found for prey-infested leaves, but on infested leaves, more eggs were laid near the prey along the adaxial midvein than along the abaxial midvein. Results showed that prey cues and presence of prey guide predator oviposition, even within the single leaf. The perspectives for biological control in orchards are discussed.     

Oviposition response of Simulium reptans (Diptera: Simuliidae) to the presence of conspecific eggs

Abstract.

• 1 The oviposition behaviour of Simulium reptans L. is described from two sites on the River Spey, Scotland. Female aggregations were observed immediately downstream of oviposition sites and were composed mainly of gravid flies (range 60–80%0) together with smaller numbers that were either freshly bloodfed (range 2–17%) or infected with mermithid nematodes (range 0–30%).
• 2 The time from landing on the oviposition sites to the onset of oviposition was recorded. The time in the presence of greater than 1-day-old eggs did not significantly differ from sites with no eggs present. However, the presence of freshly laid or I-day-old eggs significantly shortened the time to onset of oviposition. The cues that elicit oviposition are unknown but it is speculated that they may involve a pheromone.

     

Chemosterilization of the cabbage root fly under field conditions

Removing hedgerow feeding sites from within 40, 80 or 160 m of cauliflower plots, failed to reduce populations of the cabbage root fly (Erioischia brassicae (Bouché)). Oviposition on the plots was unchanged but the efficiency of yellow water-traps was improved. Traps along-side cleared hedgerows caught more gravid than non-gravid females. In cages, 94% of the eggs laid were sterile when both sterilant baits containing tepa [tris-(i-aziridinyl) phosphine oxide] and hedgerow flowers were present. In the field, maximum egg-sterility rarely exceeded 30% when sterilant was present when natural sterility averaged 8%. Chemosterilization was an inadequate control since sufficient eggs and larvae survived to produce root-damage indices of 49 % in contrast to 2 % on insecticide-treated plants. Most eggs were sterile when chemosterilant baits were placed 0.75 m apart and bait was most effective when placed around the crop during the pre-oviposition period and through the crop at the time of maximum oviposition. Increasing the concentration of tepa failed to increase the effectiveness of the bait but a higher proportion of the eggs laid were sterile on large than on small plots. When chemosterilant treatments were stopped, egg-fertility gradually reverted to the natural level. Protracted immigration, an innate tendency for females to disperse, reductions in the competitiveness of sterile males, and the failure of males to re-disperse once sterilized, appeared to be the main factors limiting the levels of sterility in root-fly populations exposed to tepa-baited lures 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.     

Ovipositional biology of Viburnum leaf beetle, Pyrrhalta viburni (Coleoptera: Chrysomelidae)

Pyrrhalta viburni Paykull, a new landscape pest in the United States, feeds in both the larval and adult stages on foliage of plants in the genus Viburnum. We measured lifetime oviposition capacity of mated and unmated females reared in the laboratory versus field-collected females, as well as ovipositional response to physical characteristics of the host plant. Both mated and unmated females produced eggs, but at different rates. Field-collected females and mated females reared in the laboratory laid similar numbers of egg masses containing similar numbers of eggs, but unmated females laid approximately one half as many eggs, the result primarily of smaller clutch size. Mated females reared in the laboratory had a preovipositional period of 11.4 +/- 1.7 versus 29 + 11.7 d for unmated females, and unmated females lived significantly longer than mated females. The angle and diameter of stems of V. trilobum, a very susceptible host, both greatly influenced oviposition; females laid most eggs on vertically oriented stems, and those of smallest diameter; when these factors were combined, stem diameter predominated. Females also had a very strong geotactic response, preferring to lay eggs on portions of stems toward gravity, even when stems were at fairly shallow angles.     

Laboratory and field evaluation of an oviposition trap for Culex quinquefasciatus (Diptera: Culicidae)

An ovitrap (BR-OVT) based on physical and chemical stimuli for attracting gravid Culex quinquefasciatus (Diptera: Culicidae) females was developed and evaluated under laboratory and field conditions. Attractants were assayed using alternative chamber bioassays prior to being used in the BR-OVT oviposition trap. A significant preference of gravid females for sites containing conspecific egg rafts was observed, as a response to the natural oviposition pheromone, as well as for sites treated with the synthetic pheromone erythro-6-acetoxy-5-hexadecanolide. Five- to 20-day old grass infusion was strongly attractive to gravid females for laying eggs. On the other hand, entomopathogenic Bacillus sphaericus (Bs) did not influence the choice of an oviposition site when used in combination with grass infusion and can therefore be used as a larvicide in ovitraps. Results from field trials showed that the BR-OVT with grass infusion and with or without Bs works as a preferred oviposition site for Cx. quinquefasciatus. The BR-OVT was more effective for egg collection when placed indoors and comparison with the number of egg rafts laid in cesspits over 40 days indicates that this very simple ovitrap may be a useful tool for monitoring populations of the most important of the vectors of bancroftian filariasis.     

Protection of potato crop against Phthorimaea operculella (Zeller) infestation using frass extract of two noctuid insect pests under laboratory and storage simulation conditions

The oviposition deterrent effect of water extract of Spodoptera littoralis and Agrotis ipsilon larval frass on Phthorimaea operculella adult females was studied using two types of larval food “Natural host and Semi-artificial diet” under laboratory and storage simulation (semi-field) conditions. Extracted frass of fed larvae on semi-artificial diet showed complete oviposition deterrent effect at treatments with 4th, 5th and 6th instars of S. littoralis, also at treatments with 1st–3rd and 6th instars of A. ipsilon, while the same effect was observed when the larvae fed on castor oil leaves as a natural host only at treatment with frass extract of A. ipsilon 6th instar larvae. Presence of low amounts of phenols and flavonoids in water extract of A. ipsilon larval frass resulted in relatively more effect as oviposition deterrent to fertile adult females on treated oviposition sites, while the opposite effect was obtained in S. littoralis larval frass experiments. At semi-field experiments, the percentage reduction of laid eggs reached 100% after two?days at treatments with frass extracts of 4th and 5th S. littoralis larval instars and A. ipsilon 6th instar larvae fed on semi-artificial diet and/or castor oil leaves. Percentage reduction of laid eggs for untreated sacks reached 93.24 and 48.95% after 2 and 30?days, respectively, when placed between treated sacks, in comparison with the mean number of laid eggs for isolated control.     

Distribution of second generation European corn borer,Ostrinia nubilalis, egg masses in field corn and relationship to subsequent tunneling damage

The relationship between second generation European corn borer (Ostrinia nubilalis Hübner) egg mass numbers and subsequent field corn damage, as measured by stalk cavity numbers, was studied in 79 fields in northeastern North Carolina over three years. A mean of 0.028 egg masses per plant (645 egg masses/23400 plants) was found over the course of the study. Significant differences in oviposition rate were detected between fields and years. Ca. 85% of egg masses were deposited in a five leaf zone surrounding the primary ear; of these, 89% were found on the lower four leaves in this zone. Egg masses appeared to be distributed randomly within fields but at low rates of incidence, and oviposition was relatively uniform between sampling areas within individual fields. Under moderate to high oviposition pressure (mean number of egg masses per plant over the duration of the oviposition period >ca. 0.02), eggs laid during the early phases of the oviposition period account for more subsequent stalk damage than eggs laid during the later phases of the oviposition period. Variations in second generation egg mass numbers accounted for ca. 70% of variation in stalk cavity numbers.     

Do Golden Egg Bugs (Phyllomorpha laciniata: Heteroptera, Coreidae) Require Conspecifics for Oviposition?

Golden egg bug (Phyllomorpha laciniata) females lay eggs on the bodies of conspecifics of both sexes. We investigated to what extent reproduction depended on the availability of conspecifics as oviposition substrate and the acceptability of the host plant as an alternative oviposition substrate in the absence of conspecifics. Mated females were placed in experimental enclosures each containing a sprig of fresh host plant. Each experimental female was subjected to one of three treatments: isolated from conspecifics (solitary), paired with another female, or paired with a male. Solitary females laid a few eggs on the host plant but then stopped laying eggs, and solitary females laid significantly fewer eggs than those enclosed with another female or a male. Females enclosed with a male laid no more eggs than those enclosed with a female. When two previously isolated females were later enclosed together, they soon renewed oviposition. Females in nature contained significantly more oviducal eggs than did females that were enclosed with other females for a short period. Thus the availability of suitable conspecifics as oviposition substrate stimulates the deposition of mature eggs, and reproduction depends on the presence of conspecifics of either sex as oviposition substrate.     

VARIATION IN OVIPOSITION PREFERENCE OF HELIOTHIS VIRESCENS IN RELATION TO MACROEVOLUTIONARY PATTERNS OF HELIOTHINE HOST RANGE

When cotton, geranium, petunia, tobacco, tomato, and velvetleaf were presented simultaneously to strains of the Heliothine moth, Heliothis virescens (F.), from Arizona (AZ), Mississippi (MS), and North Carolina (NC), oviposition choices were similar, while a strain from the Virgin Islands (VI) was significantly different from both the AZ and MS strains. In the MS-VI six-host test, the VI strain laid a higher proportion of its eggs on tomato and a lower proportion of its eggs on petunia than did the MS strain. Since petunia and tomato are in the same plant family (Solanaceae), differences between the strains in host preference do not appear to involve traits common to all members of a plant family. In the AZ-VI six-host test, the AZ strain laid a higher percentage of its eggs on tobacco and lower percentages on geranium and velvetleaf (Malvaceae) than did the VI strain. The percentages of eggs laid on cotton (Malvaceae) by the two strains were not significantly different. When moths were placed in cages with only tobacco and geranium, the AZ strain again laid a higher percentage of its eggs on tobacco than did the VI strain. However, when host choice was restricted to tobacco and cotton or velvetleaf and cotton, no significant difference between strains was found. Reciprocal hybrids of the AZ and VI strains were intermediate to the parental stocks in their preference for tobacco over geranium, indicating that a genetic mechanism was responsible for oviposition preference. The relevance of these findings to seemingly rapid fluxes in host range among members of the Heliothinae is discussed.     

Oviposition by Mamestra brassicae (L.) (Lep., Noctuidae) in relation to age, time of day and host plant

The age at which females begin to lay, the daily pattern of oviposition, the influence of host-plant material on egg-laying as well as the host preference of Mamestra brassicae L. (Lep., Noctuidae) were investigated under laboratory conditions. The females began to oviposit during the third, fourth and fifth scotophase after emergence. A great variability in daily female fecundity was observed. The highest level of oviposition occurred during the fifth day. Females oviposited during the whole scotophase, but maximal oviposition occurred during the second hour. The females without host-plant material laid fewer eggs than females with host-plant material (cabbage leaves Brassica oleracea var. capitata ). In two-choice tests, the females preferred to oviposit on cabbage rather than chrysanthemum, but there was no difference in the mean number of eggs laid on cabbage and tomato. There were significantly greater mean numbers of total eggs deposited on tomato than on chrysanthemum.     

In vitro rearing of the parasitoid Exorista larvarum (Diptera: Tachinidae) from eggs laid out of host

We evaluated the possibility of using the eggs laid out of host to rear Exorista larvarum (L.) (Diptera: Tachinidae), a larval parasitoid of Lepidoptera, on artificial media. In a first experiment, eggs oviposited on a plastic sheet (either by inexperienced or experienced females) showed the same in vitro hatching capability as those removed from the larvae of the factitious host Galleria mellonella L. Subsequently, eggs laid on the host integument or out of host, either by inexperienced or experienced females, were removed from the oviposition substrate and placed on a skimmed milkbased artificial medium. The percentages of hatched eggs, of puparia and adults, as well as the puparial weights did not differ significantly among the three treatments. These findings suggested that E. larvarum may be successfully reared in vitro with a total exclusion of the host insect. In a further test, no difference for the in vitro hatching time was found between the eggs oviposited either on a plastic sheet or on G. mellonella larvae in the same length of time (60 min). This result suggested that at oviposition the out-of-host eggs were unincubated, similarly to those that had been laid on the host larvae.     

Laboratory evaluation of Toxorhynchites splendens (Diptera: Culicidae) for predation of Aedes albopictus mosquito larvae

Biology of the mosquito Toxorhynchites splendens (Wiedemann) was studied in the laboratory to provide baseline data for using the predatory larvae of this species against those of Aedes albopictus (Skuse) in a biological control programme. The mean incubation time of Tx.splendens eggs was 43.8 h and the time required for newly-hatched larvae to initiate predation was 2.5 h. Mean numbers of prey larvae consumed and killed by each Tx.splendens larva totalled 389 and 345 respectively. The larval period of Tx.splendens was not significantly different for rearing individually or in groups of nine, with equal prey density, and duration of larval development was proportional to prey density. In mass rearing, larval cannibalism was usually observed during days 1-3 post-eclosion. The incidence of cannibalism decreased sharply on the fourth day after hatching when some larvae became fourth-instar. Adult female Tx.splendens usually commenced oviposition on day 4 after emergence. The number of eggs laid daily increased on day 7 and the peak oviposition of 6.3 eggs/female/day occurred on day 11. When oviposition containers were provided only intermittently, gravid females of Tx.splendens scattered most of their eggs on the dry floor of the cage. Viability of eggs laid by females aged 4-14 days was high (60-90%) but decreased to less than 40% as the females aged.     

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.     

The influence of tactile and other non-chemical factors on the ovipositional responses of the generalist herbivore Epiphyas postvittana

A bioassay to test the ovipositional responses of the generalist herbivore, Epiphyas postvittana, was developed, and various non-chemical factors that influenced the laying, and number of eggs laid, by females were tested. In choice experiments using various models, surface texture was found to be an important factor influencing ovipositional behaviour, with females preferring to lay on smoother rather than rougher surfaces. Another surface texture feature that influenced oviposition was a raised wax ridge on a smooth surface. The wax ridge stimulated more eggs to be laid within close proximity to the ridge. Observations of females showed that the increased numbers of eggs laid on models with a wax ridge resulted not from increased landing, time spent, or numbers of eggs laid per ovipositional bout, but rather through an increased probability of ovipositing after landing on models with a wax ridge compared to models without the ridge. Spatial features of the models were also found to influence egglaying by females. Females showed a tendency to lay on the upper rather than the lower surface of models, although this appeared to depend upon how stimulatory the upper surface of the model was. Females also laid more eggs on models with a dense array of leaves than on models with a more spread-out array.     

Induction of egg maturation and oviposition in the tick Ornithodoros parkeri (Acari: Argasidae)

Hemocoelic injection and vaginal insertion of selected male reproductive and non-reproductive tissue homogenates into fed-virgin female Ornithodoros parkeri stimulated varying degrees of ovum maturation and/or oviposition during 14 and 30 day observation periods, respectively. Mean times for oviposition and mean numbers of eggs laid per ovipositing female receiving hemocoelic injections of male reproductive tissue homogenates did not differ significantly from fed-mated controls. In addition, hemocoelic injection of male salivary glandular homogenate induced oviposition, yet synganglial homogenate did not. Although vaginal insertion induced both ovum maturation and oviposition, the effect was not as pronounced as when similar doses were administered by hemocoelic injection. These results indicate that a complex interrelated series of precopulatory and copulatory stimuli are necessary for oviposition to occur in fed O. parkeri.     

Egg-laying tactic in Phyllomorpha laciniata in the presence of parasitoids

Female insects are expected to choose oviposition sites that have the best conditions for offspring development and survival. Natural enemies, such as predators and parasitoids, may have a strong influence on the selection of oviposition substrates by phytophagous insects. The golden egg bug, Phyllomorpha laciniata (Villers) (Heteroptera: Coreidae) has an unusual reproductive strategy. Females mainly use conspecifics, both males and other females, as egg-laying substrates, but occasionally they oviposit on plants as well. Survival of the eggs is higher when eggs are carried by conspecifics than when they are laid on plants, due to predation and parasitism. We investigated egg-laying behavior in the forced presence of the egg parasitoid Gryon bolivari (Giard) (Hymenoptera: Scelionidae). Specifically, we studied whether females provide egg protection by avoiding oviposition under the risk of egg parasitization. We expected a lower oviposition rate under parasitoid presence, and the eggs, if any, to be placed preferably on conspecifics and not on plants, thus ensuring higher survival of the progeny. The results show that P. laciniata 's egg-laying rate was lower when they were enclosed with parasitoids than when parasitoids were absent, especially when plants were the only substrate to oviposit on. Moreover, females showed strong preference for laying eggs on conspecifics rather than on plants. Egg-laying in P. laciniata appears to be not only influenced by the availability of conspecifics, but also by the presence of egg parasitoids. This indicates that females may be able to detect G. bolivari and avoid oviposition when parasitoids are present. We discuss the possibility of conspecifics as enemy-free space.     

Oviposition behavior of Grapholita molesta Busck (Lepidoptera: Tortricidae) at different temperatures

Cultivation of temperate-climate fruits is economically important for Brazil. Grapholita molesta Busck is a pest that causes damage to apples, peaches, plums, and pears growing in different micro-regions of southern Brazil, and understanding its reproductive behavior is essential to develop control strategies. The objective of this study was to ascertain the influence of different temperatures (13, 16, 19, 22, and 25oC) on the oviposition behavior of G. molesta. Females of G. molesta were placed in individual plastic containers, and the pre-oviposition period and the number of eggs laid were assessed until adult death. Temperature influenced the pre-oviposition period, and females kept at 22o were the first to lay their eggs. Oviposition occurred over a longer period of time at 13oC than at the higher temperatures. The highest total number of eggs was obtained at 19oC, with the mean daily oviposition being directly proportional to the temperature. There was a negative interaction between the pre-oviposition period and the total number of eggs laid by females. The most suitable temperature for oviposition of G. molesta was 19oC.     

Do plant cues influence the oviposition behavior of Japanese beetles?

A bioassay arena was developed to test the response of the Japanese beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae), to various plant‐derived factors that may influence the selection of oviposition sites. In choice experiments with natural and artificial plant cues, females preferred to lay eggs in ryegrass rather than in artificial resources. Artificial grass stems received more eggs on average than bare ground, indicating the role of vertical objects on the soil surface as oviposition cues. Popillia japonica females were able to discriminate between the quality and quantity of the artificial stems, as more eggs were laid in soil blocks that had larger diameter stems or a higher number of stems per soil block. Beetles were capable of evaluating plant characteristics on the soil surface and adjusted their investment accordingly; more eggs were laid in resources that were chosen more frequently. Observations of females in choice arenas revealed how surface cues influence digging behavior and the time spent under the soil surface. The number of digging events was not significantly different among the three treatments with surface cues (ryegrass, grass roots, and artificial grass), but beetles spent significantly more time in the soil under ryegrass than any of the other treatments. These findings suggest that oviposition by P. japonica is selective and is influenced by plant‐derived cues evaluated before and after digging into the soil.