Life tables of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer (Lepidoptera : Thaumetopoeidae) at local and regional scales

A major ongoing debate in population ecology has surrounded the causative factors underlying the abundance of phytophagous insects and whether or not these factors limit or regulate herbivore populations. However, it is often difficult to identify mortality agents in census data, and their distribution and relative importance across large spatial scales are rarely understood. Were, we present life tables for egg batches and larval cohorts of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer, using intensive local sampling combined with extensive regional monitoring to ascertain the relative importance of different mortality factors at different localities. Extinction of entire cohorts (representing the entire reproductive output of one female) at natural localities was high, with 82% of the initial 492 cohorts going extinct. Mortality was highest in the egg and early instar stages due to predation from dermestid beetles, and while different mortality factors (e.g. hatching failure, egg parasitism and failure to establish on the host) were present at many localities, dermestid predation, either directly observed or inferred from indirect evidence, was the dominant mortality factor at 89% of localities surveyed. Predation was significantly higher in plantations than in natural habitats. The second most important mortality factor was resource depletion, with 14 cohorts defoliating their hosts. Egg and larval parasitism were not major mortality agents. A combination of predation and resource depletion consistently accounted for the majority of mortality across localities, suggesting that both factors are important in limiting population abundance. This evidence shows that O. lunifer is not regulated by natural enemies alone, but that resource patches (Acacia trees) ultimately, and frequently, act together to limit population growth.

Role of chemical cues from cotton in mediating host selection and oviposition behaviour in Helicoverpa armigera (Hubner) (Lepidoptera : Noctuidae)

We investigated the role of chemoreception in the host selection and oviposition behaviour of Helicoverpa armigera in the laboratory using five cotton genotypes and synthetic volatile terpenes. Female moths oviposited on substrates treated with methanol, ethanol, acetone and pentane extracts of leaves, squares and flowers of the cotton genotypes. Phytochemicals soluble in pentane were the most efficient in eliciting oviposition behaviour. In a two-way bioassay, pentane extracts of leaves or squares of a Multiple Host-plant Resistance genotype (MHR11), Deltapine commercial (DP90), and Smith Red Leaf (SRL) received significantly more eggs than solvent-treated controls. Extracts of squares of the native genotype Gossypium nelsonii did not receive more eggs. Females preferred DP90 and MHR11 to SRL and G. nelsonii. Female moths also laid more eggs on pentane extracts of MHR11 flowers than MHR11 leaves from preflowering, early flowering and peak-flowering plants. In a flight chamber, female moths used olfactory cues at short range to mediate oviposition and discrimination between host plants. Egg-laying, mated females were attracted at a distance (1.5 m) to volatile compounds released by whole plants and odours emanating from filter papers treated with synthetic volatile terpenes. Individually, the terpenes did not stimulate any significant oviposition response. However, there was a significant oviposition response to a mixture of equal volumes of the terpenes (trans-beta-caryophyllene, alpha-pinene, beta-pinene, myrcene, beta-bisabolol, and alpha-humulene). Conversely, antennectomised (moths with transected antennae), egg-laying, mated females did not stimulate any significant oviposition response. The significance of these findings in relation to H. armigera hostplant selection are discussed.

Analysis of long-term light-trap data for Helicoverpa spp. (Lepidoptera : Noctuidae) in Australia: the effect of climate and crop host plants

Regression analyses of a long series of light-trap catches at Narrabri, Australia, were used to describe the seasonal dynamics of Helicoverpa armigera (Hubner). The size of the second generation was significantly related to the size of the first generation, to winter rainfall, which had a positive effect, and to spring rainfall which had a negative effect. These variables accounted for up to 96% of the variation in size of the second generation from year to year. Rainfall and crop hosts were also important for the size of the third generation. The area and tonnage of many potential host crops were significantly correlated with winter rain. When winter rain was omitted from the analysis, the sizes of both the second and third generations could be expressed as a function of the size of the previous generation and of the areas planted to lucerne, sorghum and maize. Lucerne and maize always had positive coefficients and sorghum a negative one. We extended our analysis to catches of H. punctigera (Wallengren), which declines in abundance after the second generation. Winter rain had a positive effect on the sizes of the second and third generations, and rain in spring or early summer had a negative effect. Only the area grown to lucerne had a positive effect on abundance. Forecasts of pest levels from a few months to a few weeks in advance are discussed, along with the improved understanding of the seasonal dynamics of both species and the significance of crops in the management of insecticide resistance for H. armigera.

Toxicity of biorational insecticides to Helicoverpa spp. (Lepidoptera : Noctuidae) and predators in cotton field

Field trials on upland cotton (Gossypium hirstum L.) during its reproductive phase were used to assess the toxicity of several biorational pesticides and chemicals to Helicoverpa armigera (Hubner) and H. puntigera Wallengren, as well as major predators at Dalby, Queensland, Australia. Moderate rate-dependent control was obtained in plots treated with neem (Azadirachta indica A. Juss) seed extract-azadirachtin (Aza) at rates of 30, 60 and 90 g/ha. Plots treated with Talstar EC (bifenthrin) applications achieved the best results, followed by treatment with alternation of chemicals (methomyl, bifenthrin, thiodicarb and endosulfan) and biorational insecticides (neem oil, azadirachtin and Bacillus thuringiensis kurstaki var. Berliner). Predators, including lady beetles, lacewings, spiders and predatory bugs, were insensitive to Aza, tooseendanin (Tsdn) and BT applications. In contrast, chemicals were very destructive of predators. All treatments provided some protection from infestation of H. armigera and H. puntigera. The effect of Aza on Helicoverpa spp. was reflected in a relatively higher yield of seed cotton harvested from Aza-treated plots compared with the control, but chemical control achieved significantly higher yields than any other treatment.

Survival and development of Helicoverpa armigera (Hubner) (Lepidoptera : Noctuidae) on neem (Azadirachta indica A. Juss) leaves

The survival, development and feeding responses of Helicoverpa armigera (Hubner) on neem (Azadirachta indica A. Juss) leaflets were evaluated under choice and no-choice conditions. Neem leaflets provided to starving fourth-instar larvae were chewed, but larvae expectorated most of the material. The weight of surviving larvae decreased and no fourth instars completed development on neem leaflets alone. Larvae resumed feeding when transferred to cotton leaves after 5 days of feeding on neem leaflets. Fourth instars strongly discriminated between neem leaflets and cotton leaves when offered a choice. Early sixth instars decreased in weight and had delayed development when feet only on neem leaflets. Mure than one-half lived for more than 2 weeks and some completed development to adults. Neem is an unacceptable non-host plant species for H. armigera.

Foraging and vein-cutting behaviour of Euploea core corinna (W.S. Macleay) (Lepidoptera : Nymphalidae) caterpillars feeding on latex-bearing leaves

Caterpillars of Euploea core corinna (W. S. Macleay) sever leaf veins prior to feeding on their latex-bearing host plants, which restricts the flow of latex at feeding sites. The severing of leaf veins by insects feeding on latex-bearing plants is commonly referred to as 'sabotaging' and is thought to be an evolved response by the insect to counter the negative effects of feeding on latex-rich leaves. Sabotaging behaviour is described for all instars of E. core corinna, with particular attention given to neonates. Vein cutting by neonate E. core corinna caterpillars can occur within 2 h of hatching, with most caterpillars establishing feeding sites within 10 h. Commonly, first instars cut an are-shaped row of leaf side-veins parallel to the leaf margin, but they may also cut the leaf mid-rib in a fashion similar to older instar larvae. From a sample of 50 E. core corinna larvae, representing all instars, we found that the diameters of the veins cut by caterpillars are closely correlated to larval head width (r=0.90). Through manipulative experiments, we demonstrate for the first time that sabotaging behaviour in neonate caterpillars imposes no detectable short-term physiological costs on those caterpillars.

Ecology and behavior of first instar larval Lepidoptera

Neonate Lepidoptera are confronted with the daunting task of establishing themselves on a food plant. The factors relevant to this process need to be considered at spatial and temporal scales relevant to the larva and not the investigator. Neonates have to cope with an array of plant surface characters as well as internal characters once the integument is ruptured. These characters, as well as microclimatic conditions, vary within and between plant modules and interact with larval feeding requirements, strongly affecting movement behavior, which may be extensive even for such small organisms. In addition to these factors, there is an array of predators, pathogens, and parasitoids with which first instars must contend. Not surprisingly, mortality in neonates is high but can vary widely. Experimental and manipulative studies, as well as detailed observations of the animal, are vital if the subtle interaction of factors responsible for this high and variable mortality are to be understood. These studies are essential for an understanding of theories linking female oviposition behavior with larval survival, plant defense theory, and population dynamics, as well as modern crop resistance breeding programs.

A comparison of the performance and biological efficiency of new knapsack sprayers and a controlled droplet application (CDA) sprayer for the control of the diamondback moth, Plutella xylostella L. (Lepidoptera : Plutellidae)

Field trials and laboratory bioassays were undertaken to compare the performance and efficacy (mortality of diamondback moth larvae) of insecticides applied to cabbages with three high volume hydraulic knapsack sprayers (NS-16, PB-20 and Selecta 12V) and a controlled droplet application (CDA) sprayer. In field experiments, the high volume knapsack sprayers (application rate 500-600 L ha(-1)) provided better spray coverage on the upper and lower surfaces of inner leaves, the upper surfaces of middle and outer leaves, and greater biological efficacy than the CDA sprayer (application rate 20similar to40 L ha(-1)). The PB-20 provided better spray coverage on the upper surface of middle leaves and both surfaces of outer leaves when compared with the Selecta 12V. However, its biological efficacy in the field was not significantly different from that of the other high volume sprayers. Increasing the application rate from 20 to 40 L ha(-1) for the CDA sprayer significantly increased droplet density but had no impact on test insect mortality. Laboratory evaluations of biological efficacy yielded higher estimates than field evaluations and there was no significant difference between the performance of the PB-20 and the CDA sprayer. Significant positive relationships were detected between insect mortality and droplet density deposited for both the PB-20 and the CDA sprayers.

Larval diet affects number of femoral 'brush' scales in male Helicoverpa punctigera moths (Lepidoptera : Noctuidae)

Males of Helicoverpa punctigera (Wallengren) show considerable variation in the number of femoral scales on the prothoracic legs. Such intraspecific variation in adult morphology could indicate the presence of undetected sibling species, or it may be related to larval diet. Helicoverpa putactigera is polyphagous, and different host plant species are likely to represent diets of different quality. Femoral lengths and the numbers of femoral scales on the prothoracic legs were therefore determined from: (i) individuals that had been collected as larvae from various host species in the field; and (ii) individuals that had been laboratory-reared, in split-family tests, on different diets, namely cotton, lucerne, sowthistle and artificial diet. Host plant species (and therefore presumably diet quality) influenced femoral length of H. punctigera males and, perhaps in conjunction with this, the number of femoral scales on the fore leg. The rearing experiment indicated, in addition, that the effect of host plant quality varies with larval stage, and that the pattern of this variation across the immature stages is dependent on host plant species. The recorded variation in the morphology of field-collected H. punctigera males is therefore most readily explained as a consequence of different individuals developing (at least for most of their larval life) on different host plant species, with diet quality varying significantly with species. The relevance of these results for insect developmental studies and evolutionary interpretations of host relationships is outlined.

Host selection and utilisation of Sonchus oleraceus (Asteraceae) by Helicoverpa armigera (Lepidoptera : Noctuidae): A genetic analysis

The cotton bollworm (Helicoverpa armigera) prefers the common sowthistle (Sonchus oleraceus L.) to cotton (Gossypium hirsutum L.), sorghum (Sorghum bicolor L.) and maize (Zea mays L.) for oviposition in the field in Australia. Using the common sowthistle and cotton as host plants, we carried out this study to evaluate genetic variation in both oviposition preference and larval growth and genetic correlation between maternal preference and larval performance. There was a significant genetic component of phenotypic variation in both characters, and the heritability of oviposition preference was estimated as 0.602. Helicoverpa armigera larvae survived slightly better and grew significantly faster on common sowthistle than on cotton, but genetic correlation between maternal preference and larval growth performance was not detectable. Instead, larval growth performance on the two hosts changed with families, which renders the interaction between family and host plant significant. As a result, the genetic correlation between mean values of larval growth across the two host species was not different from zero. These results are discussed in the context of the relationship between H. armigera and the common sowthistle and the polyphagous behaviour of this insect in general.

Taeniogonalos raymenti Carmean & Kimsey (Hymenoptera : Trigonalidae) reared as a hyperparasite of Sturmia convergens (Weidemann) (Diptera : Tachinidae), a primary parasite of Danaus plexippus (L.) (Lepidoptera : Nymphalidae)

Taeniogonalos raymenti is confirmed as a hyperparasitoid of the tachinid Sturmia convergens which parasitises larval Danaus plexippus. Trigonalids are indirect parasitoids and in this case we have direct evidence that wasp eggs must have been laid on the caterpillar's host plant. Asclepias fruticosa. before the secondary host, but not necessarily before the primary tachinid host, was present. Levels of hyperparasitism during our sampling period were very low at less than two percent.

Field test of a model of migration of moths (Lepidoptera : Noctuidae) in inland Australia

A migration of Helicoverpa punctigera (Wallengren), Heliothis punctifera (Walker) and Agrotis munda Walker was tracked from Cameron Corner (29degrees00'S, 141degrees00'E) in inland Australia to the Wilcannia region, approximately 400 km to the south-east. A relatively isolated source population was located using a distribution model to predict winter breeding, and confirmed by surveys using sweep netting for larvae. When a synoptic weather pattern likely to produce suitable conditions for migration developed, moths were trapped in the source region. The next morning a simulation model of migration using wind-field data generated by a numerical weather-prediction model was run. Surveys using sweep netting for larvae, trapping and flush counts were then conducted in and around the predicted moth fallout area, approximately 400 km to the south-east. Pollen carried on the probosces of moths caught in this area was compared with that on moths caught in the source area. The survey data and pollen comparisons provided evidence that migration had occurred, and that the migration model gave accurate estimation of the fallout region. The ecological and economic implications of such migrations are discussed.

Development of Trichogramma australicum Girault (Hymenoptera : Trichogrammatidae) in Helicoverpa (Lepidoptera : Noctuidae) host eggs

Morphological studies of development of the egg parasitoid Trichogramma australicum Girault in the cotton bollworm, Helicoverpa armigera (Hubner), were conducted to provide benchmarks for assessing developmental rates in both natural hosts and artificial diets. Observations of living embryos and histological sections show that embryos proceed rapidly through cleavage and blastoderm formation and show a characteristic pinching or rotation 8 h after deposition. Eggs progressively increase in volume, primarily by increasing in diameter at the widest point. At 29 rectangle 1 C the duration of the egg stage is 22-24 h, the larval stage 27 h, the prepupal stage 50-52 h, and pupa 85 h. Larvae undergo dramatic shape changes as they ingest food but do not show signs of larval moults, reinforcing observations that there is only one larval instar. Criteria for staging the embryonic and postembryonic development in natural hosts will be used for future studies aimed at developing and refining artificial diets for Trichogramma.