The Acrolepiopsis assectella silk cocoon: kairomonal function and chemical characterisation

Two soluble sericin-like polypeptides, B1 and B2, from leek moth (Acrolepiopsis assectella) cocoons trigger host-acceptance behaviour in the parasitoid, Diadromus pulchellus (Proc. Roy. Soc. London B 269 (2002) 1879). We found that these polypeptides were particularly cysteine-rich and lost their ability to trigger host-acceptance behaviour after being denatured and purified. This suggests that inter-disulphide bonds and the secondary structure of B1 and B2 are important for their biological activity. We also isolated six insoluble polypeptides (or polypeptides of low solubility) from A. assectella cocoons. At least four of these polypeptides triggered host-acceptance behaviour. The strongest responses were observed with P22, a light-chain fibroin or a seroin-peptide, and P100, a sericin-like polypeptide that is probably more strongly associated with the silk core than are B1 and B2. In conclusion, several polypeptides from different parts of the A. assectella silk-cocoon (the insoluble core and coating of the silk thread) are able to elicit host-acceptance behaviour in D. pulchellus females. These polypeptides belong to different silk protein families and are used as kairomones by this specialist parasitoid.     

Trap cropping effect on oviposition behaviour of the leek moth Acrolepiopsis assectella and the diamondback moth Plutella xylostella

The potential of trap cropping as a pest control strategy was explored for two specialist moth species, where oviposition behaviour determines larval distribution. Oviposition behaviour was studied at three different scales: Individual plants in the laboratory, small scale field experiments with and without cages, and medium scale field trials. The study insects were the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), a specialist on Brassica species and the leek moth, Acrolepiopsis assectella (Z.) (Lepidoptera: Acrolepiidae), a specialist on Allium species. The diamondback moth preferred Indian mustard, Brassica juncea (L.) over white cabbage, B. oleracea var alba (L.). The leek moth did not show preference when offered leek, Allium porrum (L.) and chives, A. schoenoprasum (L.). However, the leek moth did prefer to oviposit on plants of a larger biomass over smaller plants. In both the small scale and the medium scale field studies the trap crops reduced the number of eggs laid on the primary crop. The total number of eggs laid in the monoculture and the trap crop treatments were the same in the small scale field cage study, but the females distributed their eggs differently. More eggs were laid on trap crops compared with primary crops and this pattern was also validated in the medium scale field trials. In this study only the direct effects of trap cropping were considered. Indirect effects of vegetation diversity such as enhancement of natural enemies could potentially increase the efficacy of trap cropping.     

Trap cropping effect on oviposition behaviour of the leek moth Acrolepiopsis assectella and the diamondback moth Plutella xylostella

The potential of trap cropping as a pest control strategy was explored for two specialist moth species, where oviposition behaviour determines larval distribution. Oviposition behaviour was studied at three different scales: Individual plants in the laboratory, small scale field experiments with and without cages, and medium scale field trials. The study insects were the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), a specialist on Brassica species and the leek moth, Acrolepiopsis assectella (Z.) (Lepidoptera: Acrolepiidae), a specialist on Allium species. The diamondback moth preferred Indian mustard, Brassica juncea (L.) over white cabbage, B. oleracea var alba (L.). The leek moth did not show preference when offered leek, Allium porrum (L.) and chives, A. schoenoprasum (L.). However, the leek moth did prefer to oviposit on plants of a larger biomass over smaller plants. In both the small scale and the medium scale field studies the trap crops reduced the number of eggs laid on the primary crop. The total number of eggs laid in the monoculture and the trap crop treatments were the same in the small scale field cage study, but the females distributed their eggs differently. More eggs were laid on trap crops compared with primary crops and this pattern was also validated in the medium scale field trials. In this study only the direct effects of trap cropping were considered. Indirect effects of vegetation diversity such as enhancement of natural enemies could potentially increase the efficacy of trap cropping.     

Volatiles from damaged plants as major cues in long-range host-searching by the specialist parasitoidCotesia rubecula

The role of volatile stimuli in the long-range host-searching behaviour of the specialist parasitoidCotesia rubecula Marshall (Hymenoptera: Braconidae) was studied. Components from the plant-host-complex Brussels sprouts (Brassica oleracea L. var.gemmifera (DC.) Schulz. cv. ‘Titurel’)-Pieris rapae L. (Lepidoptera: Pieridae) were compared for their attractiveness in dual choice tests in a windtunnel. Stimuli from cabbage plants that were mechanically damaged or damaged byP. rapae caterpillars were more attractive to this parasitoid species than stimuli emitted by the host larvae or their faeces. Parasitoids preferred leaves from the plant-host-complex over artificially damaged leaves. Undamaged cabbage plants were the least attractive to the foraging females. These results indicate that in-flight searching behaviour ofC. rubecula is guided by plant-derived information and that for this specialist species more reliable and specific host-derived cues play a minor role at longer distances.     

Oviposition and feeding preference of Acrolepiopsis assectella Zell. (Lep., Acrolepiidae)

Abstract:  The leek moth, Acrolepiopsis assectella (Zell.), is a recently discovered exotic species in eastern Ontario and western Quebec. This Allium spp. (Asparagales, Alliaceae) specialist can cause up to 40% crop damage. A no-choice experiment was used to determine the relationship between oviposition behaviour and larval survival of the leek moth as the phylogenetic distance from the preferred host Allium ampeloprasum var. porrum L. increased. Results indicate that oviposition preference and larval survival of the leek moth declined as the phylogenetic distance from the preferred host increased. These results support the conclusion that the leek moth is a specialist feeder on closely related Allium spp. although the strength of this preference may decline as the motivation to oviposit increases. This may indicate that the leek moth is able to use closely related novel hosts as temporary refuges if the preferred host plant is unavailable.     

Responses of ovipositing moths to host plant deprivation: life history aspects and implications for intercropping

1. When considering intercropping as a strategy to reduce pest oviposition, knowledge about the insect’s oviposition behaviour is very important. Physiological effects on the insect because of difficulties in finding a suitable oviposition site may also be important. 2. In the present study, the effects that delays in access to host plants have on lifetime fecundity on diamondback moth and leek moth were examined. The ability to postpone egg laying, fecundity and lifetime oviposition are discussed in relation to intercrop/cover crop as a strategy to reduce oviposition on crop plants. 3. When faced with host plant deprivation, the diamondback moth is relatively more dependent upon host plant stimuli for the onset of egg production. By contrast, leek moth is able to postpone egg production for a longer time. There even appeared to be a tendency for leek moth females to extend their lifetime when faced with host plant deprivation. 4. We conclude that leek moths have the ability to postpone production of eggs and lay them later in life when finally encountering host plants after a period of host plant deprivation. Therefore, the use of intercropping as a strategy to reduce oviposition is questionable. For such an insect, use of a trap crop might be a better option because the female will lay her eggs in the trap crop and not get the opportunity to lay them later in life when finally encountering crop plants.     

Role of Caterpillar Silk Thread in Location of Host Pupae by the Parasitoid <Emphasis Type="Italic">Diadromus pulchellus</Emphasis>

Diadromus pulchellus is a solitary ichneumonid parasitoid. Its only known host is the pupa of Acrolepiopsis assectella, a specialist herbivore of Allium species. D. pulchellus females parasitize A. assectella pupae within 48 h after the caterpillars spin their cocoon and begin to pupate. Having observed that the cocoon produced by the leek moth caterpillar stimulates parasitoid egg-laying and that caterpillar leaves a silk thread, we studied the hypothesis that silk thread might be involved in host-finding by the parasitoid. Behavioral tests showed that when D. pulchellus females encounter a host silk thread, they change directions, follow the thread, and quickly locate the host. These findings show that pupal parasitoids can use signals produced by their hosts at the developmental instar preceding the one that they parasitize.     

Effects of specialist parasitoids on oviposition preference of phytophagous insects: encounter–dilution effects in a tritrophic interaction

Abstract.  1. Host plant preferences of the female diamondback moth Plutella xylostella were studied.
2. Female moths preferred conspecific-damaged cabbage plants over undamaged cabbage plants. The performance of P. xylostella larvae on conspecific-infested plants did not differ significantly from that of larvae on undamaged plants.
3.  Cotesia plutellae , the specialist parasitoid wasp of P. xylostella larvae, displayed equal preference for plants with differing levels of host-larvae damage, and the wasp attacked only one or two hosts on average before leaving an infested plant, irrespective of the number of hosts on the plant. It is hypothesised that the oviposition preferences of P. xylostella females for host plants already damaged by conspecific larvae demonstrate an encounter–dilution effect against C. plutellae .     

Effects of floral resources on the efficacy of a primary parasitoid and a facultative hyperparasitoid

Resources added to agroecosystems to enhance biological control are potentially available to multiple members of the resident insect community—not only the biological control agents for which the resources are intended. Many studies have examined the effects of sugar feeding on the efficacy of biological control agents. However, such information is lacking for other, interacting species such as facultative hyperparasitoids, which may contribute to pest suppression but can also interfere with introduced biological control agents. Under greenhouse conditions, we tested the direct effects of sugar and nectar provisioning on the longevity, host‐killing impact and offspring production of two pupal parasitoids associated with leek moth, Acrolepiopsis assectella: the introduced biological control agent, Diadromus pulchellus, and the native facultative hyperparasitoid, Conura albifrons. Adding sucrose, buckwheat or a combination of buckwheat and common vetch to a sugar‐deprived system (potted leek plants in cages) increased parasitoid longevity and resulted in higher leek moth parasitism and mortality compared to water or common vetch treatments. However, the two parasitoid species exhibited a distinct temporal response to the treatments, likely influenced by differences in their life histories. This study provides insight into how integrating conservation biological control techniques could affect the success of a classical biological control programme.     

The effects of periodic warming on the survival and fecundity of Diadromus pulchellus during long-term storage

Biological control strategies capitalise on natural mechanisms such as predation and parasitism to reduce the need for chemical applications to control insect pests. In Canada, the parasitic wasp Diadromus pulchellus Wesmael (Hymenoptera: Ichneumonidae) is being investigated for its use in the biological control of an invasive crop pest, the leek moth, Acrolepiopsis assectella (Zeller) (Lepidoptera: Acrolepiidae). Large numbers of insects will be needed for releases to ensure that populations of D. pulchellus establish quickly and impact leek moth populations. Since the current culture is not producing the number of insects required for large-scale releases, the accumulation and storage of D. pulchellus might be a viable option to obtain ideal numbers. Currently, little is known about the optimal conditions for the long-term storage and release of D. pulchellus, which overwinter in nature as adults. Using insects from the active culture, the effect of intermittent, short-term warming on cold-stored adults was evaluated for survivorship and fecundity. In accordance with previous findings, females survived cold storage more readily than males. However, the warming regimes employed had no significant impacts on overall survivorship. Cold-stored females had reduced fecundity compared to females maintained in the culture, though no significant differences were noted between the treatments. In addition, the offspring sex-ratio for all treatments was male skewed. Thus, the warming procedures utilised provided no advantages over current techniques for the long-term storage of D. pulchellus intended for release.     

Non-host plant extracts reduce oviposition of Plutella xylostella (Lepidoptera: Plutellidae) and enhance parasitism by its parasitoid Cotesia plutellae (Hymenoptera: Braconidae)

Botanical preparations, usually from non-host plants, can be used to manipulate the behaviour of insect pests and their natural enemies. In this study, the effects of extracts of Chrysanthemum morifolium, a non-host plant of the diamondback moth, Plutella xylostella (Linnaeus), on the olfactory and oviposition responses of this phytophagous insect and on levels of parasitism by its specialist parasitoid Cotesia plutellae (Kurdjumov) were examined, using Chinese cabbage Brassica campestris L. ssp. pekinensis as the test host plant. Olfactometer tests showed that volatiles of chrysanthemum extract-treated host plants were less attractive to P. xylostella females than those from untreated host plants; and in contrast, volatiles of the chrysanthemum extract-treated host plants were more attractive to females of its parasitoid C. plutellae than those from untreated host plants. Oviposition preference tests showed that P. xylostella females laid only a small proportion of their eggs on chrysanthemum extract-treated host plants, while ovipositing parasitoid females parasitized a much higher proportion of host larvae feeding on the treated host plants than on untreated host plants. These results suggest that certain non-host plant compounds, when applied onto a host plant, may render the plant less attractive to a phytophagous insect but more attractive to its parasitoids. Application of such non-host plant compounds can be explored to develop push-pull systems to reduce oviposition by a pest insect and at the same time enhance parasitism by its parasitoids in crops.     

The contribution of a pollinating seed predator to selection on Silene latifolia females

Interactions, antagonistic or mutualistic, can exert selection on plant traits. We explored the role of Hadena bicruris, a pollinating seed predator, as a selective agent on its host, the dioecious plant Silene latifolia. We exposed females from artificial-selection lines (many, small flowers (SF) vs. few, large flowers (LF)) to this moth. Infestation did not differ significantly between lines, but the odds of attacked fruits aborting were higher in SF females. We partitioned selection between that caused by moth attack and that resulting from all other factors. In both lines, selection via moth attack for fewer, smaller flowers contrasted with selection via other factors for more flowers. In LF females, selection via the two components was strongest and selection via moth attack also favoured increased fruit abortion. This suggests that the moths act as more of a selective force on flower size and number via their predating than their pollinating role.     

Diamondback moth performance and preference for leaves of Brassica oleracea of different ages and strata

The preference–performance hypothesis predicts that moth behaviour links plant variations with caterpillar attack and distribution, and the plant‐age hypothesis states that specialist herbivores are more successful in exploring younger plant tissue. We integrated these predictions to investigate underlying mechanisms by which moths and caterpillars of Plutella xylostella L. (Lepidoptera: Plutellidae) track and exploit within‐plant variability of leaf age and stratification. We measured leaf proteins, glucosinolates and fibre, as well as larval choice, developmental performance, and moth oviposition preference with regard to leaf age classes (young, mature and senescent) of three varieties (collard, cauliflower and cabbage) of the main host plant Brassica oleracea L. Larvae consistently fit the prediction that specialist herbivores prefer and perform better on young, upper leaves that have the highest protein level, despite the highest content of defence compounds. Conversely, moths laid more eggs on fibrous and less nutritious leaves from the lower and senescent stratum. We argue that the leaf stratification of host plants imposes conflicting selective pressures concerning offspring feeding and protection on adult females. If egg mortality is catastrophic on the upper nutritious leaves in a particular microclimatic context (e.g. sun, heat, winds, drought or rain‐washing), then oviposition preference will remain for the suboptimal lower and senescent leaves. The ability of larvae to spread upwards over the plant to access the more nutritious leaf stratum is critical when eggs are preferentially laid on the protective low‐quality leaves.     

Herbivore-induced plant volatiles mediate host selection by a root herbivore

In response to herbivore attack, plants mobilize chemical defenses and release distinct bouquets of volatiles. Aboveground herbivores are known to use changes in leaf volatile patterns to make foraging decisions, but it remains unclear whether belowground herbivores also use volatiles to select suitable host plants. We therefore investigated how above- and belowground infestation affects the performance of the root feeder Diabrotica virgifera virgifera, and whether the larvae of this specialized beetle are able to use volatile cues to assess from a distance whether a potential host plant is already under herbivore attack. Diabrotica virgifera larvae showed stronger growth on roots previously attacked by conspecific larvae, but performed more poorly on roots of plants whose leaves had been attacked by larvae of the moth Spodoptera littoralis. Fittingly, D. virgifera larvae were attracted to plants that were infested with conspecifics, whereas they avoided plants that were attacked by S. littoralis. We identified (E)-β-caryophyllene, which is induced by D. virgifera, and ethylene, which is suppressed by S. littoralis, as two signals used by D. virgifera larvae to locate plants that are most suitable for their development. Our study demonstrates that soil-dwelling insects can use herbivore-induced changes in root volatile emissions to identify suitable host plants.     

Diadromus pulchellus in North America: field release against leek moth and new characters to distinguish it from Diadromus subtilicornis,a native diamondback moth parasitoid

We report successful overwintering of Diadromus pulchellus in North America (Ontario) following introduction of this species from Europe to control the leek moth, Acrolepiopsis assectella, a recently established alien species. Field rearing revealed that the native Diadromus subtilicornis emerged only from diamondback moth, Plutella xylostella, whereas D. pulchellus was reared almost exclusively from leek moth. The single D. pulchellus reared from diamondback moth was anticipated because host range studies found this species could develop on both leek moth and diamondback moth in the laboratory, although, it had not been previously reported from diamondback moth in the field in Europe. DNA barcoding of specimens of both Diadromus spp. confirmed their species status and novel morphological characters are presented to distinguish D. pulchellus from D. subtilicornis. In addition, DNA from specimens of D. subtilicornis from Europe clustered with DNA from specimens across Canada, confirming that it is a single Holarctic species. Finally, a new host association for D. subtilicornis is recorded from the dame's rocket moth: Pseudoplutella porrectella.     

Herbivore-Specific,Density-Dependent Induction of Plant Volatiles: Honest or “Cry Wolf” Signals?

Plants release volatile chemicals upon attack by herbivorous arthropods. They do so commonly in a dose-dependent manner: the more herbivores, the more volatiles released. The volatiles attract predatory arthropods and the amount determines the probability of predator response. We show that seedlings of a cabbage variety (Brassica oleracea var. capitata, cv Shikidori) also show such a response to the density of cabbage white (Pieris rapae) larvae and attract more (naive) parasitoids (Cotesia glomerata) when there are more herbivores on the plant. However, when attacked by diamondback moth (Plutella xylostella) larvae, seedlings of the same variety (cv Shikidori) release volatiles, the total amount of which is high and constant and thus independent of caterpillar density, and naive parasitoids (Cotesia vestalis) of diamondback moth larvae fail to discriminate herbivore-rich from herbivore-poor plants. In contrast, seedlings of another cabbage variety of B. oleracea (var. acephala: kale) respond in a dose-dependent manner to the density of diamondback moth larvae and attract more parasitoids when there are more herbivores. Assuming these responses of the cabbage cultivars reflect behaviour of at least some genotypes of wild plants, we provide arguments why the behaviour of kale (B. oleracea var acephala) is best interpreted as an honest signaling strategy and that of cabbage cv Shikidori (B. oleracea var capitata) as a “cry wolf” signaling strategy, implying a conflict of interest between the plant and the enemies of its herbivores: the plant profits from being visited by the herbivore''s enemies, but the latter would be better off by visiting other plants with more herbivores. If so, evolutionary theory on alarm signaling predicts consequences of major interest to students of plant protection, tritrophic systems and communication alike.     

<Emphasis Type="Italic">Catalpa bignonioides</Emphasis> alters extrafloral nectar production after herbivory and attracts ant bodyguards

Inducible anti-herbivore defenses are found within many plant taxa, but there are fewer examples of inducible indirect defenses that incorporate the third trophic level. This study links caterpillar foraging, herbivore-induced changes in extrafloral nectar production, and the attraction of ants to vulnerable leaves and plants. Catalpa bignonioides Walter (Bignoniaceae) uses extrafloral nectar to attract ant (Forelius pruinosus(Roger)) bodyguards in response to Ceratomia catalpae (Boisduval)(Lepidoptera: Sphingidae) herbivory. Ant density per leaf increased with the sugar content of extrafloral nectar excreted by sampled leaves, suggesting that increased nectar production could attract or retain beneficial arthropods. The masses of sucrose, fructose, glucose and all three sugars combined in the extrafloral nectar increased two- to three-fold on attacked leaves within 36 h of the experimental addition of caterpillars. Production rates for neighboring non-attacked leaves and non-attacked leaves on adjacent plants did not differ over the same time period. Ant attendance at caterpillar-attacked leaves increased two- to three-fold within 24 h of herbivory, relative to attendance at neighboring, undamaged leaves. These attacked leaves attracted the fewest ants prior to the onset of herbivory, suggesting the specialist caterpillar may avoid or be excluded from leaves with more bodyguards. The removal of leaf tissue with scissors did not alter ant attendance at damaged leaves. Mean ant attendance per leaf on attacked plants increased 6- to 10-fold after caterpillar introduction, relative to adjacent unattacked plants. The plant's biotic defense thus operates at two scales; the number of bodyguards (ant workers) on the plant increases after attack, and this increased workforce is biased towards attacked leaves within plants. Fewer caterpillars remained on plants that attracted greater numbers of ants, suggesting these bodyguards benefit the plant.     

Behavior of the tachinid parasitoid <Emphasis Type="Italic">Exorista japonica</Emphasis> (Diptera: Tachinidae) on herbivore-infested plants

Odors from corn plants infested with the larvae of the noctuid herbivore Mythimna separata (Walker) attract tachinid fly, Exorista japonica Townsend, females; and odors from corn plants artificially damaged also attract this fly. We investigated the responses of flies to herbivore-infested, artificially damaged, and undamaged plants before and after arrival at the target plants to measure timing of the behavioral sequence. The behavior of the flies for 10 min after takeoff from a release point was observed in a wind-tunnel bioassay. The percentage of flies attracted to the plants was higher in infested and artificially damaged plants than in undamaged plants. Latency before takeoff was longer in undamaged plants, and time from takeoff to arrival at the plant was also longer in undamaged plants. Moreover, flies stayed longer on infested and artificially damaged plants. Flies walked longer on infested plants than on artificially damaged and undamaged plants. In this paper, we summarize the behavioral data and discuss the host-searching behavior of E. japonica females.     

The role of female search behaviour in determining host plant range in plant feeding insects: a test of the information processing hypothesis

Recent theoretical studies have suggested that host range in herbivorous insects may be more restricted by constraints on information processing on the ovipositing females than by trade-offs in larval feeding efficiency. We have investigated if females from polyphagous species have to pay for their ability to localize and evaluate plants from different species with a lower ability to discriminate between conspecific host plants with differences in quality. Females of the monophagous butterflies Polygonia satyrus, Vanessa indica and Inachis io and the polyphagous P. c-album and Cynthia cardui (all in Lepidoptera, Nymphalidae) were given a simultaneous choice of stinging nettles (Urtica dioica) of different quality. In addition, the same choice trial was given to females from two populations of P. c-album with different degrees of specificity. As predicted from the information processing hypothesis, all specialists discriminated significantly against the bad quality nettle, whereas the generalists laid an equal amount of eggs on both types of nettle. There were no corresponding differences between specialist and generalist larvae in their ability to utilize poor quality leaves. Our study therefore suggests that female host-searching behaviour plays an important role in determining host plant range.     

Chemotype of <Emphasis Type="Italic">Senecio jacobaea</Emphasis> affects damage by pathogens and insect herbivores in the field

The evolution of the diversity of plant secondary metabolites is still poorly understood. To determine whether natural enemies could exert selection on plant secondary chemistry, pathogen infestation and invertebrate herbivory were measured on 10 genotypes of Senecio jacobaea (Tansy Ragwort) at two experimental field sites during a 2-year period. The genotypes represented two chemotypes based on the presence of the pyrrolizidine alkaloids (PA) jacobine and erucifoline. At one site, Heteren, mainly generalist herbivores were present. Here, damage was limited and did not differ among genotypes or chemotypes. At the other site, Meijendel, several specialists attacked the plants. Damage increased during the year, with a peak in July when most damage was caused by the specialist moth Tyria jacobaeae. At this peak there was no difference in damage among chemotypes. In the months prior to T. jacobaeae damage, chemotypes with jacobine were more severely attacked by specialists than the chemotypes without jacobine. Total damage during that period was positively correlated with both total PA concentration and jacobine concentration. Probably plant vigor also played a role in host preference since damage per individual plant was positively correlated with plant size. Our results suggest that total PA concentration and specifically jacobine had a positive effect on specialist feeding, indicating ecological costs involved in the production of PAs. Ecological costs related to plant secondary compounds could explain why not all individuals produce high levels of these compounds. In addition, differences in specialist herbivore pressures among sites may contribute to the variation in secondary metabolites among populations.