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.     

Distribution ofChilo partellus (Lepidoptera: Pyralidae) egg batches on maize

Distribution ofChilo partellus egg masses was studied in field, greenhouse, and laboratory experiments. The eggs were laid in batches mainly on the lower side and the lower leaves of the plant. The egg batch size ranged from 1 to 169 eggs, with a median of 33.5 eggs per batch (average, 40.5). Oviposition ofC. partellus is described at two levels. The first level, choice of oviposition plants, followed a random distribution. The second level, number of egg batches per plant, followed an aggregated distribution in the field, where more than one egg batch was deposited on the same plant by the same female, which was found on 25% of the oviposition plants. A mechanism for egg-layingC. partellus females to perceive preceding oviposition or injured plants could not be detected. Oviposition site choice seemed to be mediated by tactile stimuli.     

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.     

Oviposition Behavior of the Rice Brown Planthopper, Nilaparvata lugens (Stål), and Its Electronic Monitoring

Oviposition behavior of Nilaparvata lugens (Stål) (Homoptera: Delphacidae) on rice, Oryzae sativa L., was monitored using a videocamera and an AC (alternating-current) electronic measuring system. Oviposition by N. lugens always took place during stylet penetration into the plant. After penetration of the ovipositor, there were three specific ovipositional sequences observed; sawing insertion of the ovipositor, release of an egg in the plant, and partial or complete withdrawal of the ovipositor from the plant. Distinctive waveforms were recorded in correlating with these events. From a sequence of waveform patterns, it was possible to determine the number of eggs and egg masses laid inside the plant tissue and the duration of oviposition behavior. Waveform analysis revealed that N. lugens laid an egg mass consisting of on average 5.7 eggs. This occurred 4.3 min after a period of about 6.4 min of stylet penetration, and this sequence was repeated several times in 24 h. Electrical and/or tethering effects on oviposition appear to be negligible within 24 h of recording.     

Oviposition Deterrence Is Likely an Effect,Not a Mechanism,in the Leaf Beetle <Emphasis Type="Italic">Chrysophtharta bimaculata</Emphasis> (Olivier) (Coleoptera: Chrysomelidae)

Oviposition deterrence is common in many insects as an evolutionary mechanism to reduce subsequent larval competition. We investigated a suspected case of oviposition deterrence by the paropsine chrysomelid, Chrysophtharta bimaculata. In paired choice tests, gravid females were found to prefer ovipositing on host leaves without conspecific eggs, confirming the presence of an apparent oviposition deterrence mechanism. Washing egg batches in water, hexane, or ethanol did not change this preference, suggesting that a soluble marking pheromone was not involved. Furthermore, it is unlikely that a plant-derived oviposition deterring substance is produced as beetles showed no significant oviposition preference between leaves which had been oviposited upon, but then had the eggs removed, and those that had never been oviposited upon. In trials using artificial leaves and mimic egg batches, “leaves” with “egg batches” placed near the tip of the leaf (the preferred site of oviposition in this species) were significantly less likely to be laid upon than artificial leaves where mimic eggs were placed away from the tip. In combination, the results strongly infer that oviposition deterrence in C. bimaculata is due to the mechanical blocking of the oviposition site by the first laid egg-batch, rather than a specific oviposition deterring cue. The apparent oviposition deterrence in this insect may well be an outcome or evolutionary effect of oviposition-site selection, rather than a clear adaptive mechanism to decrease larval competition.     

Preference of Bt‐resistant and susceptible Busseola fusca moths and larvae for Bt and non‐Bt maize

The sustainability of genetically engineered insecticidal Bacillus thuringiensis Berliner (Bt) maize, Zea mays L. (Poaceae), is threatened by the evolution of resistance by target pest species. Several Lepidoptera species have evolved resistance to Cry proteins expressed by Bt maize over the last decade, including the African maize stem borer, Busseola fusca (Fuller) (Lepidoptera: Noctuidae). The insect resistance management (IRM) strategy (i.e., the high‐dose/refuge strategy) deployed to delay resistance evolution is grounded on certain assumptions about the biology and ecology of a pest species, for example, the interactions between the insect pest and crop plants. Should these assumptions be violated, the evolution of resistance within pest populations will be rapid. This study evaluated the assumption that B. fusca adults and larvae select and colonize maize plants at random, and do not show any preference for either Bt or non‐Bt maize. Gravid female B. fusca moths of a resistant and susceptible population were subjected to two‐choice oviposition preference tests using stems of Bt and non‐Bt maize plants. Both the number of egg batches as well as the total number of eggs laid on each stem were recorded. The feeding preference of Bt‐resistant and susceptible neonate B. fusca larvae were evaluated in choice test bioassays with whorl leaf samples of specific maize cultivars. Although no differential oviposition preference was observed for either resistant or susceptible female moths, leaf damage ratings indicated that neonate larvae were able to detect Bt toxins and that they displayed feeding avoidance behaviour on Bt maize leaf samples.     

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.     

Some host-plant factors affecting oviposition by aphidophagous Syrphidae (Diptera)

Although most aphidophagous syrphid species lay their eggs close to aphids, Platycheirus peltatus (Meig.), Melanostoma scalare (F.) and M. mellinum (L.) laid freely on uninfested brussels sprout plants. The existence of species that will lay in the absence of aphids may be valuable in biological control and also provides a useful tool for the investigation of non-aphid oviposition stimuli. Plant species and plant appearance were both important factors in P. peltatus oviposition. Particular sites on the plant were chosen for oviposition by Syrphus balteatus (Deg.). The nature of the substrate at the oviposition site affected the size of the egg batches laid there by M. scalare, M. mellinum, P. peltatus and P. clypeatus (Meig.). The possibility is discussed that plant-location is the older-established system of host-finding and that an aphid-location system has gradually superseded it during the evolution of the obligatorily aphidophagous Syrphinae.     

绿盲蝽成虫的产卵行为与习性

本文观察记录了绿盲蝽Apolygus lucorum(Meyer-Dür)雌性成虫的产卵行为,并研究了其产卵习性。结果发现,绿盲蝽卵主要产在植物组织中,单次产卵平均历时31.4 s;产卵主要在夜间进行,白天的产卵量仅占全天的6.6%。成虫交配后主要产可育卵,后期偶产不育卵;而未交配个体大部分能产不育卵。在25℃下,成虫从7日龄起开始产卵,16日龄前产卵量占总产卵量的48.9%;17～40日龄间产卵量约占40%。在棉株上,约65%卵分布在中部(第4到第7果枝),同时94.3%卵集中在叶柄、叶脉、蕾柄和铃柄上。     

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.     

Dynamics of oviposition in Danaus plexippus (Insecta: Lepidoptera) on milkweed, Asclepias spp

Where a butterfly chooses to lay an egg will influence the subsequent survival of her offspring. In this paper we report on the effect of a number of variables which influence the choice of oviposition site in Danaus plexippus L. Experiments were conducted under both flight cage and field conditions. The field observations consisted of recording the within and between plant egg dispersions across different patch sizes. Laboratory experiments looked at egg laying preferences as affected by plant species, age and condition. Butterflies were selective in their choice of oviposition site. Eggs were laid singly on the underside of medium sized leaves towards the top of a plant. More eggs were laid per plant on single isolated plants than on plants within a patch. The number of eggs per plant increases with plant height but decreases with plant age. Females preferred young plants or plants with fresh regrowth of leaves. These characteristics could override species preferences which were, in decreasing order, Asclepias curassavica. A. fruticosa and A. physocarpa. Species preferences varied between butterflies and with female age. Neither the presence of eggs nor larvae on a plant deterred oviposition. These results are compared with previous observations of egg laying in this species.     

An algal cyclostat with computer-controlled dynamic light regime

Platycnemis pennipes deposits eggs into plant tissue at or just below the water surface. On a number of different plant species egg deposition rate was found to be similar, while oviposition duration varied. Eggs were not always to be found in the substrate after a female had probed the substrate with her ovipositor. Actual oviposition took place only during longer stays.     

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.     

Reduced realised fecundity in the pine looper Bupalus piniarius caused by host plant defoliation

1. Whether the potential fecundity of herbivorous insects is realised or not may depend on female behaviour, which in turn may be influenced by host plant acceptability. Female Bupalus piniarius were observed to discriminate against needles growing out the year following defoliation (current‐year needles) of its host plant Pinus sylvestris. 2. It was hypothesised that the discriminatory behaviour was due to current‐year needles being less secure as a substrate. Field and laboratory experiments were designed to test this hypothesis and to estimate the discrepancy between potential and realised fecundity when females were offered defoliated branches. 3. In a laboratory oviposition experiment, B. piniarius females were exposed to branches bearing either current‐year needles only or both mature and current‐year needles. Daily oviposition rate, egg batch size, longevity, and mature eggs and fat retained at death were recorded for each female. In field experiments, the rate at which eggs dropped from expanding needles and the capacity of neonate larvae hatching from the dropped eggs to colonise a tree were assessed. 4. Significantly fewer eggs were laid when females were exposed to defoliated branches. 5. Twenty‐six and 16% of the eggs laid on current‐year needles dropped from the needles in 1998 and 1999 respectively, whereas no eggs dropped from mature needles in 1998 and only one egg (< 1%) dropped in 1999. 6. A very small proportion of larvae hatching on the forest floor (simulated egg drop) was able to recolonise host trees. 7. These results emphasise the importance of oviposition behaviour on realised fecundity when analysing insect population dynamics. In the case of B. piniarius, egg placement, although a minor detail during the normal course of events, became of key importance when defoliation deprived females of their preferred egg attachment site.     

Stylus of the odonate endophytic ovipositor: a mechanosensory organ controlling egg positioning

Using light and scanning electron microscopy, a sensory field consisting of 15-20 campaniform sensillae is described on the base of the stylus of the endophytic ovipositor of Odonata. It is hypothesised that two symmetric styli equipped with this number of sensillae can function as a mechanosensory organ responsible for control of precise egg positioning in plant stems during oviposition. In laboratory experiments with females of damselflies Lestes sponsa and Lestes barbarus (Lestidae), it was demonstrated that the distance between laid eggs is not dependent on the presence of styli. Removal of styli from both sides did not influence a shift of oviposition to one side. Females with one removed stylus shifted the clutch line in the opposite direction toward the removed stylus. Additionally, removal of styli influenced positions of single eggs in egg sets, and disturbed the capacity for complex oviposition. Thus, both morphological and experimental data support the hypothesis that styli participate in the control of egg line and egg patterning in the clutch.     

Physical and chemical factors influencing landing and oviposition by the cabbage root fly on host-plant models

Various plant models were used in both choice and no-choice tests in the laboratory to assess landing and oviposition preferences of the cabbage root fly,Delia radicum (L.). The main factor governing the site most suitable for landing was the conspicuousness of the object and not its shape. Oviposition was influenced considerably by the pre-conditioning of the females. Deprived females laid eggs even when denied access to both host plant chemicals and host-plant models. The dominant role of contact chemical stimuli in host acceptance was reconfirmed, but only a combination of physical and chemical stimuli appeared capable of eliciting normal oviposition. The combination of contact chemical stimuli and the presence of a stem on the test model had a synergistic effect on the numbers of eggs laid in both choice and no-choice situations. In choice bioassays, female cabbage root flies distinguished between models of different shapes, heights and sizes. The size and shape of the models appeared to be perceived in part after the flies had landed.     

Influence of protein hydrolysate baits on oviposition in the apple maggot, Rhagoletis pomonella

Ovipositional responses of apple maggot (AM), Rhagoletis pomonella (Walsh), females were studied in the laboratory on apples (var: Golden Delicious) treated with different rates of four protein hydrolysate baits in choice and no-choice tests. Protein hydrolysate baits at rates of 0.5 and 1% had no significant effect, but oviposition was greatly reduced at higher rates of 5 and 10%. Apple maggot females exposed to apples treated with protein hydrolysate baits at a rate of 10% made 41–71% fewer punctures and laid 41–73% fewer eggs than in untreated control. No oviposition activity was shown on apples treated with 25 and 100% Nulure®. In no-choice tests the AM females laid 75–96% fewer eggs in apples treated with 10 and 25% Nulure compared to controls and no oviposition occurred in apples treated with 100% Nulure. Apple maggot females arrived in similar numbers on apples treated with 10% Nulure and untreated apples, but only 5% of those arriving on Nulure-treated apples showed ovipositor boring with no egg deposition while 60% of females arriving on untreated apples showed ovipositor boring activity and laid an average of 2.5 eggs per apple. In another experiment, individual AM females displayed similar behavioral responses to 10% Nulure-treated apples; none of the 56 females tested on treated apples displayed ovipositor boring activity, but 59% of the females (N=56) tested on untreated apples displayed ovipositor boring within 5 min of their arrival. Ninetyeight percent of AM females stayed and fed on fruit surfaces for 5 min on Nulure-treated apples without ovipositor boring compared to only 2% on untreated apples. Of the females that arrived on untreated apples, 39% flew away within 5 min without ovipositor boring compared to only 2% of those that arrived on Nulure-treated apples. Results of these two behavioral experiments suggest that upon arrival on a protein bait-treated apple, an apparent change of behavior occurs in AM females and instead of attempting to oviposit, they attempt to feed on fruit surfaces resulting in reduced oviposition activity. These results indicate that the feeding and oviposition-related activities of AM females are probably mutually exclusive and that the feeding behavior preempts oviposition activities on host fruits treated with higher rates of protein hydrolysate baits.     

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.     

Oviposition behaviour and response of Monochamus scutellatus (Coleoptera: Cerambycidae) to conspecific eggs and larvae

• 1 The oviposition behaviour of white‐spotted sawyer beetle Monochamus scutellatus (Say) (Coleoptera: Cerambycidae) females was studied under laboratory and field conditions.
• 2 Females walked over the bark surface and stopped to palpate it with their labial and maxillary palpi. Once a potential oviposition site was located, the female excavated a scar in the bark with her mandibles, deposited an egg and brown plug in the scar and inserted her mandibles again into the scar prior to leaving the area.
• 3 Female M. scutellatus preferred to lay eggs on bolts containing no eggs (control) to those containing eggs from other females, but did not show this preference when provided with control bolts and those containing their own eggs. When females were provided with bolts containing their own eggs and bolts containing eggs from other females, they laid more eggs on bolts containing their own eggs. Females laid more eggs on control bolts than on bolts containing their own larvae or larvae from other females.
• 4 The brown plug in the oviposition scar may provide a protective barrier to the egg, a chemical message that the site is occupied, or both.

     

Oviposition choice by the Onopordum capitulum weevil Larinus latus (Coleoptera: Curculionidae) and its effect on the survival of immature stages

Female Larinus latus lay eggs into the capitula of their Onopordum spp. host plants from the onset of capitulum development until the completion of flowering. This tight linkage between insect life-history and plant development optimised larval survival, apart from some initial asynchrony between the readiness of the females to oviposit and the suitability of the very small capitula to physically support eggs. Eggs were laid on bracts and stems of capitula or directly in the florets, when these became available. Both the location of the egg and time of oviposition influenced mortality factors such as egg and larval parasitism, egg desiccation and larval establishment. Overall, survival of eggs laid later in the season into florets was higher, although the adults that emerged were smaller. The change in choice of oviposition site that occurs at the onset of flowering, not only favours survival, but ensures more efficient resource use by the larvae. This occurs because sequential flowering of Onopordum spp. drives a sequential oviposition pattern and spreads the egg load more evenly over the available capitula, reversing a trend earlier in the season to clump eggs that were laid directly on the bracts. Other potential constraints, such as capitulum size, absolute egg density and previous egg-laying do not play a major role in determining the oviposition pattern of L. latus. Larval survival was positively correlated with capitulum size and not strongly influenced by egg density. Competition for resources only appeared to play a role in smaller capitula, and manifested itself in a reduction in the size of emerging adults rather than the death of immatures. In the absence of strong interspecific competition, the oviposition behavior of L. latus has become geared to maximising resource use for larvae (i.e. its own potential competitiveness) rather than minimising interactions with other members of the capitulum endophage guild.