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Showing papers in "Entomologia Experimentalis Et Applicata in 2000"


Journal ArticleDOI
TL;DR: The induction mechanism as well as the ecological consequences in a multitrophic and evolutionary context, and empirical evidence has been obtained that shows that the plant's response can increase plant fitness, in terms of seed production, due to a reduced consumption rate of parasitized herbivores.
Abstract: Herbivorous and carnivorous arthropods use plant volatiles when foraging for food. In response to herbivory, plants emit a blend that may be quantitatively and qualitatively different from the blend emitted when intact. This induced volatile blend alters the interactions of the plant with its environment. We review recent developments regarding the induction mechanism as well as the ecological consequences in a multitrophic and evolutionary context. It has been well established that carnivores (predators and parasitoids) are attracted by the volatiles induced by their herbivorous victims. This concerns an active plant response. In the case of attraction of predators, this is likely to result in a fitness benefit to the plant, because through consumption a predator removes the herbivores from the plant. However, the benefit to the plant is less clear when parasitoids are attracted, because parasitisation does usually not result in an instantaneous or in a complete termination of consumption by the herbivore. Recently, empirical evidence has been obtained that shows that the plant's response can increase plant fitness, in terms of seed production, due to a reduced consumption rate of parasitized herbivores. However, apart from a benefit from attracting carnivores, the induced volatiles can have a serious cost because there is an increasing number of studies that show that herbivores can be attracted. However, this does not necessarily result in settlement of the herbivores on the emitting plant. The presence of cues from herbivores and/or carnivores that indicate that the plant is a competitor- and/or enemy-dense space, may lead to an avoidance response. Thus, the benefit of emission of induced volatiles is likely to depend on the prevailing faunal composition. Whether plants can adjust their response and influence the emission of the induced volatiles, taking the prevalent environmental conditions into account, is an interesting question that needs to be addressed. The induced volatiles may also affect interactions of the emitting plant with its neighbours, e.g., through altered competitive ability or by the neighbour exploiting the emitted information. Major questions to be addressed in this research field comprise mechanistic aspects, such as the identification of the minimally effective blend of volatiles that explains the attraction of carnivores to herbivore-infested plants, and evolutionary aspects such as the fitness consequences of induced volatiles. The elucidation of mechanistic aspects is important for addressing ecological and evolutionary questions. For instance, an important tool to address ecological and evolutionary aspects would be to have plant pairs that differ in only a single trait. Such plants are likely to become available in the near future as a result of mechanistic studies on signal-transduction pathways and an increased interest in molecular genetics.

558 citations


Journal ArticleDOI
TL;DR: The ‘appropriate/inappropriate landings’ theory is discussed to indicate the type of work needed in future studies to improve the understanding of how intercropping, undersowing and companion planting can be used to optimum effect in crop protection.
Abstract: Seven hypotheses, including the ‘Resource Concentration Hypothesis’ and the ‘Enemies Hypothesis’, have been put forward to explain why fewer specialist insects are found on host plants growing in diverse backgrounds than on similar plants growing in bare soil. All seven hypotheses are discussed and discounted, primarily because no one has used any of them to produce a general theory of host plant selection, they still remain as hypotheses. However, we have developed a general theory based on detailed observations of insect behaviour. Our theory is based on the fact that during host plant finding the searching insects land indiscriminately on green objects such as the leaves of host plants (appropriate landings) and non-host plants (inappropriate landings), but avoid landing on brown surfaces, such as soil. The complete system of host plant selection involves a three-link chain of events in which the first link is governed by cues from volatile plant chemicals, the central link by visual stimuli, and the final link by cues from non-volatile plant chemicals. The previously ‘missing’ central link, which is based on what we have described as ‘appropriate/inappropriate landings’, is governed by visual stimuli. Our theory explains why attempts to show that olfaction is the crucial component in the central link of host plant selection proved intractable. The ‘appropriate/inappropriate landings’ theory is discussed to indicate the type of work needed in future studies to improve our understanding of how intercropping, undersowing and companion planting can be used to optimum effect in crop protection. The new theory is used also to suggest how insect biotypes could develop and to describe why pest insects do not decimate wild host plants growing in ‘natural’ situations.

484 citations


Journal ArticleDOI
TL;DR: It is suggested that spiders tend to remain in diversified patches and that extending the diversification throughout the whole crop (as in interspersed diversification) offers the best prospects for improving pest control.
Abstract: A review of the literature showed that spider abundance was increased by diversification in 63% of studies. A comparison of diversification modes showed that spider abundance in the crop was increased in 33% of studies by ‘aggregated diversification’ (e.g. intercropping and non-crop strips) and in 80% of studies by ‘interspersed diversification’ (e.g., undersowing, partial weediness, mulching and reduced tillage). It is suggested that spiders tend to remain in diversified patches and that extending the diversification throughout the whole crop (as in interspersed diversification) offers the best prospects for improving pest control. There is little evidence that spiders walk in significant numbers into fields from uncultivated field edges, but diversification at the landscape level serves to foster large multi-species regional populations of spiders which are valuable as a source of aerial immigrants into newly planted crops. There are very few manipulative field studies where the impact of spiders on pests has been measured in diversified crops compared with undiversified controls. It is encouraging, however, that in those few studies an increased spider density resulted in improved pest control. Future work needs are identified.

334 citations


Journal ArticleDOI
TL;DR: Alate production in pea aphids was significantly increased in large populations of two PASS‐ and PAR‐positive subclones relative to their parent clones, and this ameliorative effect of facultative bacteria at 25 °C was not found for two other sets of parent clones and their derived sub clones.
Abstract: The effects of two bacterial endosymbionts, designated PASS and PAR, were evaluated on the pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera:Aphididae), in which they occur facultatively, and on the blue alfalfa aphid, A. kondoi Shinji, in which these bacteria have not been found in natural populations. Subclones of pea aphids and blue alfalfa aphids, derived from parent aphid clones that did not contain PASS or PAR, were infected with one or both bacteria, generating PASS- and/or PAR-positive subclones with minimal genetic differences from the parent clones. Under laboratory conditions at 20 °C, PAR consistently reduced the fecundity (by between 19 and 60%) of subclones derived from three different parent pea aphid clones on bur clover, Medicago hispida Gaertn. PAR had intermediate effects on pea aphids reared on sweet pea, Lathyrus odoratus L., and had no significant effect on pea aphids on alfalfa, Medicago sativa L. The effect of PASS was either neutral or negative, depending on parent clone as well as host plant. Also at 20 °C, PASS reduced fecundity (70–77%) and longevity (40–48%), and increased the age of first reproduction (by up to 1.5 days) of blue alfalfa aphid reared on alfalfa and clover. PAR had a less dramatic effect (e.g., 30–39% reduction in fecundity) on these traits of blue alfalfa aphid. In contrast, PAR and PASS increased the fitness of pea aphid subclones of one parent clone reared for three generations at 25 °C on each of the three test plants. Without facultative bacteria, fecundity of the parent clone was reduced to a mean total of 35. However, this ameliorative effect of facultative bacteria at 25 °C was not found for two other sets of parent clones and their derived subclones. Alate production in pea aphids was significantly increased in large populations of two PASS- and PAR-positive subclones relative to their parent clones. Attempts to transmit PASS or PAR horizontally, i.e., from aphid to aphid via feeding on host plants (bur clover), were unsuccessful.

278 citations


Journal ArticleDOI
TL;DR: It is concluded that intraguild interactions have a primary effect in driving fluctuations in aphid parasitoid populations and can substantially alter the effectiveness of aphid Parasitoids as biological‐control agents.
Abstract: We reviewed the literature on aphid parasitoids to determine the occurrence, nature and outcome of intraguild interactions. Intraguild interactions were described for larval, pupal and adult aphid parasitoids and by the type of natural enemy (fungus, predator, or parasitoid). They appear to be prevalent in most aphid parasitoid systems and, except for parasitoid-parasitoid interactions, they are mostly asymmetric, with aphidophagous predators and pathogens killing parasitoids. The limited experimental evidence from field studies is insufficient to provide a comprehensive pattern of the consequences of intraguild interactions for aphid parasitoid populations in general and, more specifically, for the efficacy of biological control. However, because intraguild interactions are widespread in aphid-natural enemy communities and mostly detrimental to aphid parasitoids, we conclude that intraguild interactions have a primary effect in driving fluctuations in aphid parasitoid populations. Drawing on case studies, we further argue that intraguild interactions can substantially alter the effectiveness of aphid parasitoids as biological-control agents.

225 citations


Journal ArticleDOI
TL;DR: It is concluded that parasitization of P. rapae by C. rubecula potentially confers a considerable fitness benefit for A. thaliana plants when compared to plants exposed to feeding damage by unparasitized P.Rapae larvae, justifying the use of the term mutualism for this parasitoid‐plant interaction.
Abstract: We tested whether a plant's life time seed production is increased by parasitization of herbivores in a tritrophic system, Arabidopsis thaliana (Brassicaceae) plants, Pieris rapae (Lepidoptera: Pieridae) caterpillars and the solitary endoparasitoid Cotesia rubecula (Hymenoptera: Braconidae). We established seed production for intact A. thaliana plants, plants that were mechanically damaged, plants fed upon by parasitized caterpillars and plants fed upon by unparasitized caterpillars. In the first experiment, with ecotype Landsberg (erecta mutant), herbivory by unparasitized P. rapae caterpillars resulted in a strongly reduced seed production compared to undamaged plants. In contrast, damage by P. rapae caterpillars that had been parasitized by C. rubecula did not result in a significant reduction in seed production. For the second experiment with the ecotype Columbia, the results were identical. Plants damaged by unparasitized caterpillars only produced seeds on regrown shoots. Seed production of plants that had been mechanically damaged was statistically similar to that of undamaged plants. Production of the first ripe siliques by plants fed upon by unparasitized caterpillars was delayed by 18–22 days for Landsberg and 9–10 days for Columbia. We conclude that parasitization of P. rapae by C. rubecula potentially confers a considerable fitness benefit for A. thaliana plants when compared to plants exposed to feeding damage by unparasitized P. rapae larvae. Plants that attract parasitoids and parasitoids that respond to herbivore-induced plant volatiles will both experience selective advantage, justifying the use of the term mutualism for this parasitoid-plant interaction. This type of mutualism is undoubtedly very common in nature.

219 citations


Journal ArticleDOI
TL;DR: It is concluded that plant chemical cues play a major role in determining initial preferences through a process of emergence conditioning but that learning processes, involving cues encountered during oviposition in or contact with the host, can modify these initial preferences.
Abstract: The host preference behaviour of the generalist aphid parasitoid Aphidius colemani was investigated using a Y-tube olfactometer. Female A. colemani showed a preference for the host-plant complex on which they had been reared, even though the same aphid host was involved, demonstrating a host plant preference. This preference was not evident when the parasitoids were dissected from their mummies prior to adult emergence. Host plant preference exhibited during host selection appeared to be induced by chemical cues encountered on the mummy case at the time of emergence, but preferences could be changed by subsequent foraging experiences. It is concluded that plant chemical cues play a major role in determining initial preferences through a process of emergence conditioning but that learning processes, involving cues encountered during oviposition in or contact with the host, can modify these initial preferences.

171 citations


Journal ArticleDOI
TL;DR: Interactions among these attributes of flowers and butterflies were investigated, using data from a field study in Cornwall and from Porter et al. (1992), and findings may help to inform habitat management for butterfly nectaring flowers.
Abstract: The profitability of butterfly foraging depends in part on the corolla depth and clustering of flowers, and the tongue length, body mass and wing loading of butterflies. Interactions among these attributes of flowers and butterflies were investigated, using data from a field study in Cornwall and from Porter et al. (1992). The maximum corolla depth from which a butterfly can feed depends on tongue length, which correlates with the more easily measured attributes of body mass and wing loading. Small, short-tongued butterflies did not visit deep flowers. The quantity of nectar sugar per flower necessary for profitable foraging depends on foraging costs, which are expected to correlate with wing loading. Butterfly species with a high wing loading generally confined their visits to flowers that were clustered or very nectar-rich. Butterfly species with a low wing loading included solitary and less nectar-rich flowers in their diet. Body mass and wing loading affect a butterfly's load-carrying capacity (limiting the distance between fuelling stops) and cooling rate (limiting the distance between stops for basking or endothermic warming), and will therefore influence the capacity for floral selectivity and for migration and dispersal. Body mass, wing loading and tongue length characterised families or subfamilies of butterflies. For example vanessine nymphalids, with their long tongues and high wing loading, visited the deep, massed flowers of Buddleja davidii, but lycaenids, with their short tongues and low wing loading, did not. These often visited members of the Asteraceae. Eupatorium cannabinum, with massed flowers offering abundant and accessible nectar, was visited by butterflies of all tongue lengths and both high and low wing loading. These findings may help to inform habitat management for butterfly nectaring flowers.

126 citations


Journal ArticleDOI
TL;DR: Effects of nitrogen deficiency in hydroponically grown barley seedlings on the development and reproduction of the aphid Rhopalosiphum padi (L.) (Hemiptera: Aphididae) were investigated.
Abstract: Effects of nitrogen deficiency in hydroponically grown barley seedlings (Hordeum vulgare L.) on the development and reproduction of the aphid Rhopalosiphum padi (L.) (Hemiptera: Aphididae) were investigated. Plant growth was significantly reduced in seedlings grown without nitrogen. Aphid intrinsic rate of increase (r m ) was also significantly lower on these plants compared with that on plants grown with 8 mol m -3 nitrogen. Phloem sap was collected from seedling stems by aphid stylectomy and amino acids quantified by HPLC. There was a significant reduction in the concentration of non-essential amino acids as a group, but not of essential amino acids. Electrical penetration graphs (EPG) indicated that aphids reached the phloem more quickly and fed for longer on plants grown with nitrogen. This is the first reported study in which this combination of techniques has been used to understand the interactions of an aphid and plant under different environmental conditions.

112 citations


Journal ArticleDOI
TL;DR: According to the reductions in trap catch and of mating frequency measured by tethered codling moth females, efficiency of the attract and kill droplets lasted 5–7 weeks, after which it decreased slowly, and the hibernating population stayed at a low level.
Abstract: The attract and kill technique has been formulated in a product under the trade name ‘Sirene CM®’. It consists of a viscous paste containing 0.16% codlemone to attract the male moths and 6.0% permethrin to kill them. The formulation is applied by hand twice per season using a specially developed system which can be calibrated for application of the paste to the host plant in small droplets of either 100 μl or 50 μl. Between 1995 and 1997, 15 trials on control of the codling moth were conducted in isolated orchards in the Lake Geneva region. In each plot, depending on tree size, two applications varying between 52 and 537 g ha−1 of Sirene CM were made. In 14 trials, the larval attack of the codling moth on fruit was below the economic threshold of 1% and the hibernating population stayed at a low level. One single plot (0.4 ha) had to be treated with a curative spray in 1995, because the initial population was much too high. According to the reductions in trap catch and of mating frequency measured by tethered codling moth females, efficiency of the attract and kill droplets lasted 5–7 weeks, after which it decreased slowly.

105 citations


Journal ArticleDOI
TL;DR: The major factor responsible for synergistic interactions between imidacloprid and entomopathogenic nematodes appears to be the general disruption of normal nerve function due to imidcloprid resulting in drastically reduced activity of the grubs, which facilitates host attachment of infective juvenile nematode infection in white grubs.
Abstract: Entomopathogenic nematodes and the chloronicotinyl insecticide, imidacloprid, interact synergistically on the mortality of third-instar white grubs (Coleoptera: Scarabaeidae). The degree of interaction, however, varies with nematode species, being synergistic for Steinernema glaseri (Steiner) and Heterorhabditis bacteriophora Poinar, but only additive for Steinernema kushidai Mamiya. The mechanism of the interaction between imidacloprid and these three entomopathogenic nematodes was studied in the laboratory. In vials with soil and grass, mortality, speed of kill, and nematode establishment were negatively affected by imidacloprid with S. kushidai but positively affected with S. glaseri and H. bacteriophora. In all other experiments, imidacloprid had a similar effect for all three nematode species on various factors important for the successful nematode infection in white grubs. Nematode attraction to grubs was not affected by imidacloprid treatment of the grubs. Establishment of intra-hemocoelically injected nematodes was always higher in imidacloprid-treated grubs but the differences were small and in most cases not significant. The major factor responsible for synergistic interactions between imidacloprid and entomopathogenic nematodes appears to be the general disruption of normal nerve function due to imidacloprid resulting in drastically reduced activity of the grubs. This sluggishness facilitates host attachment of infective juvenile nematodes. Grooming and evasive behavior in response to nematode attack was also reduced in imidacloprid-treated grubs. The degree to which different white grub species responded to entomopathogenic nematode attack varied considerably. Untreated Popillia japonica Newman (Coleoptera: Scarabaeidae) grubs were the most responsive to nematode attack among the species tested. Untreated Cyclocephala borealis Arrow (Coleoptera: Scarabaeidae) grubs showed a weaker grooming and no evasion response, and untreated C. hirta LeConte (Coleoptera: Scarabaeidae) grubs showed no significant response. Chewing/biting behavior was significantly increased in the presence of nematodes in untreated P. japonica and C. borealis but not in C. hirta and imidacloprid-treated P. japonica and C. borealis. Our observations, however, did not provide an explanation for the lack of synergism between imidacloprid and S. kushidai.

Journal ArticleDOI
TL;DR: Despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male‐biased sex ratio in early‐instar hosts.
Abstract: In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented We compared the fitness of male and female A nigripes adults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125-175 foundresses plus 100-180 sons to the next generation However, when expressed as the innate capacity for increase (r m ), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts

Journal ArticleDOI
TL;DR: The results suggest that it is important to suppress the growth of M. stramonium and S. nigrum in order to avoid the risk of further expansion of whitefly populations where these weeds and both B‐ an Q‐biotypes are present.
Abstract: Significant differences in some reproductive parameters of the B and Q biotypes of Bemisia tabaci (Gennadius) with regard to four species of winter weeds were determined in a no-choice assay. The highest fecundity (eggs) and fertility (pupae and adults) were obtained with Malva parviflora L. as host, followed by Capsella bursa-pastoris L., Brassica kaber (DC) and Lactuca serriola L. The percentage of adult emergence (from egg to adult) was significantly higher (P < 0.001) with M. parviflora, C. bursa-pastoris and B. kaber than with L. serriola. Except on L. serriola, the mean values of the reproductive parameters for the Q-biotype were significantly (P < 0.05) greater than those for the B-biotype. On the other hand, in choice experiments with summer weeds, both B- and Q-biotypes preferred Datura stramoniumL. and Solanum nigrumL. to Amaranthus retroflexus L., Chenopodium album L. and Echinochloa crus-galliL. Significantly more males and females, and more pupae and empty pupal cases per plant were found on D. stramoniumthan on S. nigrum. Adult abundance on A. retroflexus, C. albumand E. crus-galli plants was very low. At a constant temperature of 26 1C, the Q-biotype developed significantly faster than the B-biotype on S. nigrum and D. stramonium. Both biotypes required, for this development, significantly more days on D. stramoniumthan on S. nigrum. These results suggest that it is important to suppress the growth of M. parviflora, C. bursa-pastoris, B. kaber, D. stramonium and S. nigrum in order to avoid the risk of further expansion of whitefly populations where these weeds and both B- an Q-biotypes are present.

Journal ArticleDOI
TL;DR: The susceptibility of immature stages of the legume flower thrips, Megalurothrips sjostedti, to the entomopathogenic fungus Metarhizium anisopliae, was investigated and the adult stage was found to be more susceptible to infection than the larval and pupal stages.
Abstract: The susceptibility of immature stages of the legume flower thrips, Megalurothrips sjostedti, to the entomopathogenic fungus Metarhizium anisopliae, was investigated under laboratory conditions. The adult stage was found to be more susceptible to infection than the larval and pupal stages. Mortality at all stages was dose-dependent, with the highest concentration of 1 10 8 conidia ml 1 producing the highest mortality (26, 46 and 100% for larvae, pupae and adults, respectively) at 8 days post-inoculation. At the same concentration, daily pollen consumption was significantly reduced at 2 days after treatment in infected adults but more slowly in infected larvae. Fecundity, egg fertility and longevity in adults surviving infection as larvae were significantly reduced compared to the control.

Journal ArticleDOI
TL;DR: A tentative physiological model describing the host regulation role played by different parasitoid‐derived factors is proposed.
Abstract: The biochemical profile and metabolism of Acyrthosiphon pisum (Harris) (Homoptera, Aphididae) are markedly altered and redirected in response to parasitization by the endophagous braconid Aphidius ervi Haliday (Hymenoptera, Braconidae). In the present study, the role played in the host regulation process by teratocytes, cells deriving from the dissociation of the embryonic membrane of the parasitoid, is taken into consideration. The protein synthesis activity of these cells of embryonic origin is analysed in vitro and an essential characterization of those proteins de novo synthesized and released in the incubation medium is provided. Teratocytes, obtained by dissecting parasitized host aphids, 3, 4 and 5 days after parasitoid oviposition, were incubated in vitro and, at the end of the incubation period, were separated from the medium for SDS-PAGE analysis of both cellular and secreted proteins. Various cellular proteins were more abundant as the time between parasitization and teratocyte collection increased. Furthermore, two proteins, showing an approximate molecular mass of 15 kD (p15) and 45 kD (p45) respectively, were abundantly secreted in the incubation medium by 5 day-old teratocytes. Incubations in presence of 35S radiolabelled amino acids indicated that p15 and p45 are both synthesized by A. ervi teratocytes. The amino acid composition of these two proteins was similar to that reported for other insect proteins with a demonstrated nutritional function. The p45 protein was found to be glycosylated. A tentative physiological model describing the host regulation role played by different parasitoid-derived factors is proposed.

Journal ArticleDOI
TL;DR: This work confirms the existence of an aphid cuticular kairomone, and shows that host exuviae, which also elicited strong and persistent oviposition reactions from A. ervi females, appear to be a good alternative source of ovipositional k airomone(s).
Abstract: The oviposition behaviour of the aphid parasitoid Aphidius ervi Haliday is influenced by both chemical and physical cues. Oviposition attack responses were elicited by paint pigments sealed into the tip of a glass capillary tube. Parasitoids reacted to yellow pigments with repeated oviposition attack responses, but they did not react to green pigments. The spectrum of reflected light from the yellow pigments was very similar to that from the `green' natural host Acyrthosiphon pisum (Harris), with a high proportion of the total radiation energy being emitted in the yellow-orange wavebands (580-660 nm). Pea aphid cornicle secretion also elicited oviposition attack responses, which were not exclusively induced by its pale yellow-green colour. In fact, the oviposition attack response to capillary tips coated with cornicle secretion remained evident under red light conditions, which, in contrast, nearly completely suppressed the response to yellow pigments. Chemical compounds from cornicle secretion do not appear to be involved in parasitoid orientation, even though they stimulate intense oviposition attack responses. Olfactometer experiments showed that the putative kairomone involved acts only at very short range or on contact. Host exuviae, which also elicited strong and persistent oviposition reactions from A. ervi females, appear to be a good alternative source of ovipositional kairomone(s). This work confirms the existence of an aphid cuticular kairomone.

Journal ArticleDOI
TL;DR: The oviposition behaviour of Episyrphus balteatus DeGeer (Diptera: Syrphidae) was investigated in four‐choice experiments in screen cages and in experiments with direct behavioural observations.
Abstract: The oviposition behaviour of Episyrphus balteatus DeGeer (Diptera: Syrphidae) was investigated in four-choice experiments in screen cages and in experiments with direct behavioural observations. Females laid hardly any eggs in the absence of aphids and preferred larger aphid colonies to small ones. Honeydew attracted the females via olfaction and elicited oviposition. Females were able to perceive the presence of conspecific eggs and oviposited less often in aphid colonies in which eggs were present. The oviposition deterring stimulus was also active when the eggs were removed, hence, the deterrent did not act visually, but probably olfactorily.

Journal ArticleDOI
TL;DR: The data suggest that stimulation of reproduction by these insecticides probably does not play a major role in cotton aphid outbreaks or resurgence, and that modifying aphid population structure and growth through the use of juvenoids such as pyriproxifen may be an effective proactive approach to pest control without adversely impacting beneficial organisms or causing pest resurgence.
Abstract: Insecticides are often implicated in causing outbreaks of the cotton aphid, Aphis gossypii (Glover) (Homoptera: Aphididae), through stimulation of reproduction. In this study we report the sublethal effects of dosages of bifenthrin, acephate, carbofuran or pyriproxifen on cotton aphid reproduction. We could not detect any increase or decrease in the intrinsic rate of increase of cotton aphids exposed to bifenthrin, acephate or carbofuran. However, we did detect some increases in the net reproductive rate of aphids treated with bifenthrin justifying further investigation of the effect on reproduction by this insecticide. Trends based on simple linear regression models suggest that sublethal dosages of bifenthrin or carbofuran have a negative impact on aphid population growth as dosages increase. These data suggest that stimulation of reproduction by these insecticides probably does not play a major role in cotton aphid outbreaks or resurgence. Pyriproxyfen is a juvenoid currently used for control of whiteflies in cotton. It demonstrated significant activity towards cotton aphid reared on treated cotton in our bioassays. Pyriproxyfen caused sterility in most aphids exposed to dosages exceeding 1 ppm, and reduced aphid longevity by approximately 50%. However, it did not appear to greatly influence the reproductive potential or longevity of reproductively mature aphids. A field study indicates that pyriproxyfen affects cotton aphid population structure and may have potential in managing cotton aphid outbreaks. Modifying aphid population structure and growth through the use of juvenoids such as pyriproxifen may prove to be an effective proactive approach to pest control without adversely impacting beneficial organisms or causing pest resurgence.

Journal ArticleDOI
TL;DR: The movements of insects which underlie their population distributions are discussed, the value of integrating spatial information into improved management strategies for C. assimilis is discussed and the potential for the spatial targeting of insecticides to reduce the amount applied and to conserve T. perfectus is discussed.
Abstract: The spatio-temporal distribution of Ceutorhynchus assimilis Payk. (Coleoptera: Curculionidae) adults caught in a rectangular grid of flight traps in a crop of winter oilseed rape (Brassica napus L.) was mapped and was analysed using Spatial Analysis by Distance IndicEs (SADIE). Their distribution was compared to that of their larvae and that of their parasitoid Trichomalus perfectus (Walker) (Hymenoptera: Pteromalidae) in pods. The distribution of immigrating C. assimilis adults was consistent with their arrival at the crop boundaries and movement within the crop towards its centre. Adult C. assimilis were aggregated at all times, invasion being on two fronts, leading to the formation of two major clusters within the crop. Large areas of the crop remained relatively unpopulated. During the emigration phase, numbers declined simultaneously in all parts of the crop. The distributions of adult and larval C. assimilis and of larval T. perfectus were spatially associated. The distribution of the parasitoid did not show a density dependent relationship with that of its host. We discuss the movements of insects which underlie their population distributions, the value of integrating spatial information into improved management strategies for C. assimilis and the potential for the spatial targeting of insecticides to reduce the amount applied and to conserve T. perfectus.

Journal ArticleDOI
TL;DR: In both years, parasitoids were predicted to have emerged from overwintering mummies well in advance of the onset of aphid infestation, and more than a month earlier than the first parasitized aphids were found in winter wheat.
Abstract: Temperature dependencies were established for the egg-to-mummy and mummy-to-adult phases, for mummy mortality, and for parasitism of Aphidius ervi Haliday, Aphidius rhopalosiphi De Stefani-Perez, and Praon volucre (Haliday) (Hymenoptera, Aphidiidae), three parasitoids of Sitobion avenae (Fabricius) (Homoptera, Aphididae), at 8 °C, 12 °C, 16 °C, 20 °C, and 25 °C on winter wheat (cv. Haven). A physiological model described temperature-dependent development over the full temperature range, whereas a linear model was fitted for data above 8 °C and used to estimate the lower temperature thresholds and day-degrees (° D) required for development. The thresholds for A. ervi were 2.2 °C for egg-mummy development and 6.6 °C for mummy-adult development, those for A. rhopalosiphi were 4.5 °C and 7.2 °C, and those for P. volucre were 3.8 °C and 5.5 °C. The time to develop into mummies and adults differed significantly between the three species: A. ervi development into mummies required an average of 159 ° D, while development into adults took an average of 73 ° D. The corresponding average times required for A. rhopalosiphi and P. volucre to develop mummies were 124° D and 126° D, while their development into adults required an average of 70° D and 150° D, respectively. Mummy mortality was 25–35% at 8 °C and less at the higher temperatures tested, but began to increase again at 25 °C, showing a quadratic relationship between mortality and temperature. Parasitization was very low or, in the case of P. volucre, absent up to 12 °C and thereafter increased with increasing temperature. The relationship between parasitization, recorded as percent aphids mummified, and temperature was linear at the temperatures tested and depended on species. A. ervi superparasitized 11.1% aphids at 20 °C and 16.6% aphids at 25 °C, whereas superparasitism was low in A. rhopalosiphi and absent in P. volucre. From 16 °C to 25 °C the P. volucre sex ratio increased. For A. ervi and A. rhopalosiphi there was no trend with temperature, but at 20 °C and 25 °C it was close to even. Field data for 1996 and 1997 allowed for a comparison of actual and expected emergence of overwintering mummies. In both years, parasitoids were predicted to have emerged from overwintering mummies well in advance of the onset of aphid infestation, and more than a month earlier than the first parasitized aphids were found in winter wheat. Observations from trap plants in other crops supported the predictions of the models. Other factors that can affect biological control by cereal aphid parasitoids are discussed.

Journal ArticleDOI
TL;DR: The consequences of constitutive and induced gland expression on two types of herbivores, spider mites (cell content feeders) and noctuid caterpillars (leaf chewers) are determined.
Abstract: Key words: Density-dependent induction, genotype-by-environment interaction, pigment glands, herbivory,induced plant defense, plant-insect interactions, plasticityAbstractCotton plants contain suites of phytochemicals thought to be important in defense against herbivores, some ofwhich are localized in pigment glands which contain gossypol and other terpenoid aldehydes. The simple geneticbasis for the expression of these glands has led to the development of near-isogenic glanded and glandless geno-types. Glands may also be phenotypically induced by herbivory. We determined the consequences of constitutiveandinducedgland expressionontwo types ofherbivores, spidermites (cell contentfeeders)andnoctuidcaterpillars(leaf chewers).Induction of glands was strongly dependent on the density of attackers. Spider mite herbivory on cotyledons(1) increased the density (but not total number) of glands on cotyledons linearly, (2) increased the density andtotal number of glands on the first true leaf linearly, and (3) affected the density and total number of glands on thesecond true leaf non-linearly, compared to controls. Neither constitutive nor induced expression of glands affectedmite population growth. An equal reductionof mite population size on induced glanded and glandless plants (50%)relative to uninduced controls indicated that factors other than glands were associated with induced resistance tomites. Constitutive gland expression had a strong negative impact on caterpillar performance, reducing growth by45%. Induced resistance to caterpillars was three times stronger in glanded genotypes than in glandless genotypes,indicating that factors associated with induced resistance to caterpillars are strongly associated with glands. Threecotton varieties were highly variable in their constitutive and induced resistance to mites and caterpillars.Thus, defense of cotton plants against herbivores can be roughly categorized as constitutive and induciblefactors associated with terpenoid aldehyde containing pigment glands that are effective against caterpillars, andfactors not associated with glands that are effective against mites.IntroductionPlants are the basal resource for organisms in manyfood webs. As a consequence, most plants are con-sumed by numerous herbivores. Where herbivoresreduce plant fitness, natural selection has presum-ably favored the evolution of defensive mechanismsin plants. Plant defenses can be expressed constitu-tively (all of the time) and facultatively (induced bysome environmental cue) (Karban & Baldwin, 1997;Agrawal, 1998). Genotypically determined constitu-tive and induced expression of phytochemicals canmediate either specific or general resistance to at-tackers. Furthermore, different resistance mechanismshave the potential to interact with each other to in-crease (or decrease) defense (Berenbaum & Zangerl,1996). Some plant defenses such as the tetracyclictriterpenoid cucurbitacins of the cucurbitaceae are ef-fective against many attackers including spider mites,roaches, beetles, lepidopteran larvae, mice and ver-tebrate grazers (Da Costa & Jones, 1971; Metcalf L Tallamy et al., 1997; Agrawal et al.,

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TL;DR: The number of host and parasitoid extinctions described in this study exceeds the number of extinctions usually observed in field studies of host‐parasitoid metapopulations.
Abstract: Metapopulation theory makes a number of predictions concerning the effects of dispersal on the persistence of predator-prey or host-parasitoid systems. While the stabilising effects of dispersal have been shown in a number of laboratory studies, evidence from field studies remains scarce due to a lack of suitable model systems. I describe a host-parasitoid system that shows a classical metapopulation structure with frequent extinctions and colonisations consisting of the aphidiid Lysiphlebus hirticornis and the aphid Metopeurum fuscoviride. Both the parasitoid and the aphid are specialists on their respective hosts. I followed the dynamics of host and parasitoid on individually marked tansy (Tanacetum vulgare) plants, the host of M. fuscoviride. Dynamics of host and parasitoid populations were characterized by frequent extinctions and colonisations. Mean longevity of aphid colonies was only 3.1 weeks. Parasitism by L. hirticornis was a main cause of extinction for the aphid as rates of parasitism often reached 100%, in particular towards the end of the field season. Patchiness in this system occurs at two spatial scales. Aphid colonies form on single tansy ramets = shoots but movements of aphid individuals among ramets within a particular tansy genet are frequent. Because aphids can persist on a genet for a large numer of generations, it is argued that local populations form on genets rather than ramets. The number of host and parasitoid extinctions described in this study exceeds the number of extinctions usually observed in field studies of host-parasitoid metapopulations. It is suggested that aphid-parasitoid systems such as the one studied in this paper may be good models to test the predictions of metapopulation theory.

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TL;DR: Olfactometry showed that the response of spring migrants of the bird cherry‐oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid.
Abstract: Olfactometry showed that the response of spring migrants of the bird cherry-oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid. Between three and four days after becoming adult, and having left the winter host Prunus padus L., aphids lost their negative response to the chemical. The change in response was not associated with contact with a summer host, oats. In a settling choice bioassay, migrants avoided oats which had been exposed to volatile methyl salicylate. Aphids with removed antennal tips did not avoid the exposed plant, indicating that plant choice was influenced by cues from the plant surface. The results are discussed in relation to the use of methyl salicylate in integrated control.

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TL;DR: Predictions of prey weight consumption rate by Coleomegilla maculata lengi was higher at intermediate prey size leading to a convex prey utilization curve were fulfilled and it was demonstrated that it is not the optimal size range for this predator.
Abstract: Prey utilization by the generalist predator Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) of three sympatric lepidopterous species was quantified in relation with prey size (age) and prey species. Based on optimal foraging theory, we argued that costs associated with the utilization of small and large prey are higher than those of intermediate prey size. As a result, we expected a higher prey consumption rate on intermediate prey size leading to a convex prey consumption curve. Laboratory experiments showed that, within a given prey instar, Coleomegilla maculata lengi preyed more on Plutella xylostella (L.) compared to Artogeia rapae (L.) and Trichoplusia ni (Hubner). Generally, prey consumption rate by Coleomegilla maculata lengi on the three prey species decreased with increasing immature prey size (age). The predation efficacy of Coleomegilla maculata lengi adults and fourth instar larvae was higher compared to younger coccinellids (L2). Although, Coleomegilla maculata lengi showed a higher level of predation on smaller immature prey, we demonstrated that it is not the optimal size range for this predator. As predicted, prey weight consumption rate by Coleomegilla maculata lengi was higher at intermediate prey size leading to a convex prey utilization curve. The beneficial impact of Coleomegilla maculata lengi predation on the host plant was also estimated by using a Protection Index that considers the differential predation caused by the coccinellids and the relative importance of each pest species in terms of plant injury. Coleomegilla maculata lengi has a more significant beneficial impact when it preys on T. ni immatures.

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TL;DR: The results indicate that the attraction of secondary pests is facilitated by broken grain kernels, which resulted from either mechanical damage during harvesting and/or binning procedures, or the feeding activity of primary insect pests.
Abstract: Observations on behavioural activity involved in the host selection by secondary pests of stored grains, Oryzaephilus surinamensis (Linnaeus), Tribolium castaneum (Herbst) and Tribolium confusum J. du Val, with respect to intact and mechanically or naturally damaged kernels are reported. Our results indicate that the attraction of secondary pests is facilitated by broken grain kernels, which resulted from either mechanical damage during harvesting and/or binning procedures, or the feeding activity of primary insect pests. Insect damaged kernels were more attractive to O. surinamensis, T. castaneum and T. confusum than whole kernels; in addition insect damaged kernels elicit more attractiveness than mechanically split kernels. The damage caused by primary pests, such as Rhyzopertha dominica (Fabricius) and Sitophilus oryzae (Linnaeus), on whole kernels may facilitate colonization by secondary pests, which continue damaging the cereals. O. surinamensis, T. castaneum and T. confusum utilize the grain volatile odours to distinguish whether the grain kernels of the stored cereals are damaged mechanically or by insects.

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TL;DR: The facts suggest that Wolbachia causes thelytoky and Telenomus spec.
Abstract: Two egg parasitoid species, Telenomus nawai and Telenomus spec. are similar morphologically but they have been treated as different species because of their different reproductive forms, arrhenotoky and thelytoky. Male progeny were produced from the thelytokous colony of Telenomus spec. (TT) by heat and antibiotic treatments. These males mated successfully with females of arrhenotokous colony of T. nawai (AT) and the females produced on average 19.1% males. This percentage did not differ from that obtained from AT females mated with AT males (19.4%). Diagnostic PCR indicated that TT is infected with Wolbachia; antibiotic treatments eliminated Wolbachia from TT. These facts suggest that Wolbachia causes thelytoky and Telenomus spec. (TT) is conspecific with T. nawai (AT).

Journal ArticleDOI
TL;DR: The egg parasitoid studied is known to recognize host eggs of X. luteola by contact kairomones extractable from the egg shell, and clearly differentiated between host eggs and eggs of another closely related chrysomelid species, Galerucella lineola L., as was shown by comparing duration of antennal drumming on host eggs or eggs of G. lineola.
Abstract: Plant synomones and host kairomones are known to guide the egg parasitoid Oomyzus gallerucae to its specific host, the elm leaf beetle Xanthogaleruca luteola (= Pyrrhalta) (Muller) feeding upon elm leaves (Ulmus spp.). In this study, we investigated whether the activities of these plant synomones and kairomones are specific for the plant and herbivore species, respectively. Olfactometer and contact bioassays were used. In habitat location, O. gallerucae (Fonscolombe) is known to use synomones from Ulmus minor (Miller) that are induced by egg depositions of X. luteola. The attractiveness of such induced volatiles was shown to be specific both for the Ulmus species and the herbivore species depositing eggs. Neither leaves of U. glabra Hudson (= U. montana) carrying eggs of X. luteola nor leaves of U. minor (= U. campestris = U. procera) carrying eggs of the chrysomelid species Galeruca tanaceti L. emitted attractive synomones. O. gallerucae is also known to be attracted by volatile kairomones from faeces of X. luteola feeding on U. minor and to show prolonged antennal drumming when contacting substrates contaminated with these faeces. The kairomonal activity of the faeces was proved to be independent of the Ulmus species, since also faeces from elm leaf beetles feeding upon U. glabra emitted attractive volatiles. However, the faecal kairomones were specific for the herbivorous species, since faeces from a lepidopteran larva (Opisthograptis luteolata L.) feeding upon elm hardly elicited any antennal drumming in O. gallerucae. The egg parasitoid studied is known to recognize host eggs of X. luteola by contact kairomones extractable from the egg shell. O. gallerucae clearly differentiated between host eggs and eggs of another closely related chrysomelid species, Galerucella lineola L., as was shown by comparing duration of antennal drumming on host eggs and eggs of G. lineola.

Journal ArticleDOI
TL;DR: Courtship behaviour of the polyphagous ectoparasitoid Lariophagus distinguendus Först was studied and wing fanning and arrestment of the males was elicited by paper discs treated with dichloromethane extracts from virgin females showing the existence of a female‐derived sex pheromone.
Abstract: Courtship behaviour of the polyphagous ectoparasitoid Lariophagus distinguendus Forst. (Hymenoptera, Pteromalidae) was studied. The initial behavioural element of the courtship sequence is a characteristic wing fanning shown by the males at encounters with females. Wing fanning and arrestment of the males was elicited by paper discs treated with dichloromethane extracts from virgin females showing the existence of a female-derived sex pheromone. The pheromone is only active at a distance between 0–5 mm suggesting low volatility of the active compound(s). Females mate only once, and the pheromone is still perceived by males at least 5 days after female mating. Males exposed to dissected female heads, thoraces and abdomens, showed wing fanning towards all segments. However, extracts from female abdomens were significantly more active than those from heads or thoraces suggesting the pheromone source to be located in the abdomen.

Journal ArticleDOI
TL;DR: Data show that after the introduction of a new crop plant into a region, significant changes during the recruitment process in a pestiferous insect may take place, contributing to the future pest status of the insect.
Abstract: Populations of the rapeseed pollen beetle Meligethes aeneus F. (Col., Nitidulidae) from areas with 0-16 years of history of intensive rapeseed growing were compared for key ecological characters. During the first 16 years of rapeseed cultivation the reproductive success of M. aeneus increased 200-300% over that of the beetles living on the natural host plants, cruciferous weeds. The increase was linear over time and statistically highly significant, and it did not appear to be related to food quality or to the size of the beetles. During the same period the tolerance to intraspecific competition decreased, possibly due to the relative absence of such competition on the new crop. Furthermore, the optimum population density for M. aeneus to maximize the size of its next generation on summer turnip rape was determined to be 0.5-1.0 beetles/plant, which is slightly below the economic threshold for chemical control (1 beetle/plant). Therefore the practical protection of the rapeseed yield also ensures the highest possible pest population size for the next year. These mechanisms may in part explain the particular noxiousness of the species as a pest all over Europe. In general these data show that after the introduction of a new crop plant into a region, significant changes during the recruitment process in a pestiferous insect may take place, contributing to the future pest status of the insect. It is suggested that such genetic and ecological changes in insects may be a more common mechanism than previously thought in initiating and sustaining pest outbreaks, and that conventional pest management methods may enhance that effect.

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TL;DR: The performance, as measured by daily rate of parasitism and total parasitism, of five aphelinid species found in Australia parasitising Bemisia tabaci were compared on cotton, hibiscus, rockmelon, soybean and tomato and found that more whiteflies were parasitised when cotton was the source host or rockmelon the test host.
Abstract: The performance, as measured by daily rate of parasitism and total parasitism, of five aphelinid species found in Australia parasitising Bemisia tabaci were compared on cotton, hibiscus, rockmelon, soybean and tomato. Two Eretmocerus spp., both indigenous to Australia, gave the highest levels of parasitism on each of the plant host species tested. The tritrophic interactions between B. tabaci, host plant species and Eret. mundus (Australian parthenogenetic form) (APF) were also examined. In general, more whiteflies were parasitised when cotton was the source host or rockmelon the test host. Parasitism was always low when tomato was either the source or test host. When parasitoids were transferred from rockmelon to cotton, parasitism declined. In contrast, parasitism increased when parasitoids were transferred from cotton to rockmelon. Parasitism also increased when parasitoids were transferred from soybean to rockmelon, yet failed to do so when shifted from soybean to cotton despite cotton normally being a better host. However, when parasitoids were transferred from cotton to soybean there was a marked increase in parasitism. Possible causes are discussed. The field cage trial demonstrated that parasitism by both Eretmocerus spp. increased with increasing whitefly density. Further, the increase in parasitism was not due to the presence of more parasitoids as neither the parasitoid-whitefly ratio nor the total number of parasitoids present had a significant effect on parasitism. The combination of the two species, gave similar levels of parasitism to that achieved by Eret. mundus (APF) alone. Subsequent identification of the emerged individuals indicated that over 50% of the parasitism was due to this species suggesting that it out-competed Eret. queenslandensis. Despite this competition, there was no evidence that overall control was compromised.