The effects of the insecticide chlorpyrifos on spider and Collembola communities

The effects of chlorpyrifos on aquatic systems are well documented. However, the consequences of the pesticide on soil food webs are poorly understood. In this field study, we hypothesised that the addition of a soil insecticide to an area of upland grassland would impact spider and Collembola communities by decreasing numbers of spiders, consequently, causing an increase in detritivore numbers and diversity. Chlorpyrifos was added to plots on an upland grassland in a randomised block design. Populations of Collembola and spiders were sampled by means of pitfall traps (activity density) and identified to species. Twelve species of Collembola were identified from the insecticide-treated and control plots. Species diversity, richness and evenness were all reduced in the chlorpyrifos plots, although the total number of Collembola increased ten-fold despite the abundance of some spider species being reduced. The dominant collembolan in the insecticide-treated plots was Ceratophysella denticulata, accounting for over 95% of the population. Forty-three species of spider were identified. There were a reduced number of spiders in insecticide-treated plots due mainly to a lower number of the linyphiid, Tiso vagans. However, there was no significant difference in spider diversity between the control and insecticide treatments. We discuss possible explanations for the increase in abundance of one collembolan species in response to chlorpyrifos and the consequences of this. The study emphasises the importance of understanding the effects of soil management practices on soil biodiversity, which is under increasing pressure from land development and food production. It also highlights the need for identification of soil invertebrates to an 'appropriate' taxonomic level for biodiversity estimates. (C) 2007 Elsevier GrnbH. All rights reserved.

Suitability of Arabidopsis thaliana as a model for host plant-Plutella xylostella-Cotesia plutellae interactions

Successful pest management is often hindered by the inherent complexity of the interactions of a pest with its environment. The use of genetically characterized model plants can allow investigation of chosen aspects of these interactions by limiting the number of variables during experimentation. However, it is important to study the generic nature of these model systems if the data generated are to be assessed in a wider context, for instance, with those systems of commercial significance. This study assesses the suitability of Arabidopsis thaliana (L.) Heynh. (Brassicaceae) as a model host plant to investigate plant-herbivore-natural enemy interactions, with Plutella xylostella (L.) (Lepidoptera: Plutellidae), the diamondback moth, and Cotesia plutellae (Kurdjumov) (Hymenoptera: Braconidae), a parasitoid of P. xylostella. The growth and development of P. xylostella and C. plutellae on an A. thaliana host plant (Columbia type) were compared to that on Brassica rapa var. pekinensis (L.) (Brassicaceae), a host crop that is widely cultivated and also commonly used as a laboratory host for P. xylostella rearing. The second part of the study investigated the potential effect of the different A. thaliana background lines, Columbia and Landsberg (used in wider scientific studies), on growth and development of P. xylostella and C. plutellae. Plutella xylostella life history parameters were found generally to be similar between the host plants investigated. However, C. plutellae were more affected by the differences in host plant. Fewer adult parasitoids resulted from development on A. thaliana compared to B. rapa, and those that did emerge were significantly smaller. Adult male C. plutellae developing on Columbia were also significantly smaller than those on Landsberg A. thaliana.

Learning and natal host influence host preference, handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, naive female R vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female R vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining R vindemiae attack rates. mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness.

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra-Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in large colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV2 of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV2>1 for A. funebris between plant patches, while for T. centaureae CV2>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50-200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Differential susceptibilities of peach-potato aphid, Myzus persicae (Homoptera: Aphididae) and its parasitoid, Aphidius matricariae (Hymenoptera: Aphidiidae) to foliar insecticides on partially resistant and susceptible potato cultivars

In a field experiment the effects of Sumicidin (super) 5EC (fenitrothion), Metasystox EC25 (oxydemeton-methyl) and Tamaron SL600 (methamidophos), applied at different dosages, were evaluated against peach-potato aphid, Myzus persicae (Sulzer) and its parasitoid Aphidius matricariae Haliday on Cardinal and Desiree (respectively partially resistant and susceptible potato cultivars to M. persicae). Sumicidin (super) 5EC was found about 30% more effective in reducing aphid populations than the other insecticides tested. The highest doses of each insecticide caused maximum aphid mortality; in general aphid mortality appeared dose dependent. Almost all the higher and lower doses of the tested insecticides were about 19% more effective on Cardinal than on Desiree. The most significant result was the synergistic interaction at the lower doses with plant resistance, so that the same level of control was recorded with second highest dose on Cardinal as with the highest dose on Desiree. Also the same control level was achieved at the lowest dosage rate on Cardinal compared with the next higher dose on the Desiree. Sumicidin (super) 5EC was found least toxic to the parasitoid, A. matricariae in terms of percent parasitism, emergence of parasitoids and number of mature eggs in the emerging female parasitoids; increase of about 22, 67 and 47% respectively were found in parasitoid performance with Tamaron SL600 which was found comparatively highly toxic. The highest doses of all insecticides were found clearly toxic to the parasitoid. In general, effects on the parasitoid were dose dependent. Maximum yield was obtained from the second highest dose of Sumicidin (super) 5EC.

Selective toxicity of different granular insecticides to the peach-potato aphid, Myzus persicae (Homoptera: Aphididae) and its parasitoid, Aphidius matricariae (Hymenoptera: Aphidiidae) in two differentially resistant potato cultivars

Field studies were conducted in Pakistan to examine the effects and the interaction of two differentially resistant potato cultivars i.e. Cardinal and Desiree (one partially resistant and one susceptible to Myzus persicae (Sulzer), respectively) with different dosage rates of granular insecticides, at different time intervals after application in relation to percent kill of M. persicae and effects on the parasitoid Aphidius matricariae Haliday (i.e. the third trophic level) within the aphid mummies, percent parasitism and Thimet 10G (phorate) was found about 30% more effective in reducing aphid population than the Furadan 3G (carbofuran). The highest doses of each insecticide caused maximum aphid mortality, in general aphid mortality appeared dose dependent. Mostly all the higher and lower doses of the tested insecticides were about 10% more effective on Cardinal than on Desiree. The most significant result was the synergistic interaction at the lower doses with plant resistance, so that the same level of control was recorded with the second highest dose on Cardinal as with the highest dose on Desiree. Also the same level of control was observed at the lowest dose on Cardinal as with the second last lowest dose on Desiree. Furadan 3G was found least toxic to the A. matricariae in terms of percent parasitism, emergence of parasitoids and number of mature eggs in the emerging females. Furadan 3G gave 13, 15 and 6% higher figures, respectively from the parasitoid characteristics than Thimet 10G. The highest doses of both insecticides were clearly toxic to the parasitoid. In general, the effects on the parasitoid were dose dependent. The second highest dose of Thimet 10G, gave the maximum yield

The costs and consequences of parasitoid attack for the predatory hoverfly, Episyrphus balteatus

Question: What are the life-history costs for a predatory insect of surviving parasitoid attack, and can parasitoid attack alter predator-prey interactions? Hypotheses: Survivorship is influenced by host age. Hosts that suffer parasitoid attack grow more slowly and consume fewer prey. Those that survive attack are smaller as adults and show reduced survivorship. Organisms: The aphidophagous hoverfly Episyrphus balteatus, its endoparasitoid wasp Diplazon laetatorius and its prey, the pea aphid, Acyrthosiphon pisum. Site of experiments: All experiments were conducted in controlled temperature rooms and chambers in the laboratory. Methods: Episyrphus balteatus larvae of each instar were exposed to attack by Diplazon laetatorius, then dissected to measure the encapsulation response (a measure of immunity). Second instar larvae were either attacked or not attacked by D. laetatorius. Their development rates and numbers of prey consumed were noted. The size and survivorship of surviving (immune) and control hoverflies were compared following eclosion. Conclusions: Successful immune response increased with larval age (first instar 0%, second instar 40%, third instar 100% survival). Second instar larvae that successfully resisted parasitoid attack were larger as pupae (but not as adults) and showed reduced adult survivorship. Female adult survivors were more likely than male survivors to have died within 16 days of eclosion, but there was no difference between unattacked male and female control hoverflies. Attacked larvae, irrespective of immune status, consumed fewer aphids than unattacked individuals. Episyrphus balteatus suffers significant costs of resisting parasitoid attack, and parasitoid attack can reduce the top-down effects of an insect predator, irrespective of whether the host mounts an immune response or not.

Coping with multiple enemies - the evolution of resistance and host-parasitoid community structure

Recent work has shown that the evolution of Drosophila melanogaster resistance to attack by the parasitoid Asobara tabida is constrained by a trade-off with larval competitive ability. However, there are two very important questions that need to be answered. First, is this a general cost, or is it parasitoid specific? Second, does a selected increase in immune response against one parasitoid species result in a correlated change in resistance to other parasitoid species? The answers to both questions will influence the coevolutionary dynamics of these species, and also may have a previously unconsidered, yet important, influence on community structure.

Pupal parasitoid attack influences the relative fitness of Drosophila that have encapsulated larval parasitoids

1. The evolution of host resistance to parasitoid attack will be constrained by two factors: the costs of the ability to defend against attack, and the costs of surviving actual attack. These factors have been investigated using Drosophila melanogaster and its parasitoids as a model system. The costs of defensive ability are expressed as a trade-off with larval competitive ability, whereas the costs of actual defence are exhibited in terms of reduced adult fecundity and size. 2. The costs of actual defence may be ameliorated by the host-choice decisions made by Pachycrepoideus vindemiae, a pupal parasitoid. If larvae that have successfully encapsulated a parasitoid develop into poorer quality hosts, then these may be rejected by ovipositing pupal parasitoids. 3. Pupae developing from larvae that have encapsulated the parasitoid Asobara tabida are smaller and have relatively thinner puparia. Thinner puparia are likely to be associated with a reduction in mechanical strength and possibly with a decrease in desiccation tolerance. 4. Pachycrepoideus vindemiae that develop in capsule-bearing pupae are smaller than those that emerge from previously unattacked hosts. This supports the prediction that ovipositing female P. vindemiae should avoid attacking capsule-bearing hosts. However, in choice experiments with 1-day-old pupae, P. vindemiae females oviposited preferentially in hosts containing a capsule, whereas there was no preference found with 4-day-old hosts. This appears to be a maladaptive host choice decision, as the female pupal parasitoids are preferentially attacking hosts that will result in a reduction of their own fitness. 5. The increased likelihood of attack by a pupal parasitoid is another cost of actual defence against larval parasitoid attack.

The relative fitness of Drosophila melanogaster (Diptera, Drosophilidae) that have successfully defended themselves against the parasitoid Asobara tabida (Hymenoptera, Braconidae)

Drosophila melanogaster larvae defend themselves against parasitoid attack via the process of encapsulation. However, flies that successfully defend them selves have reduced fitness as adults. Adults which carry an encapsulated parasitoid egg are smaller and females produce significantly fewer eggs than controls. Capsule-bearing males allowed repeated copulations with females do not show a reduction in their number of offspring, but those allowed to copulate only once did. No differences were found in time to first oviposition in females, or in time to first copulation in males. We interpret the results as arising from a trade-off between investing resources in factors promoting fecundity and mating success, and in defence against parasitism. The outcome of this investment decision influences the strength of selection for defence against parasitism.

Linking the coevolutionary and population dynamics of host-parasitoid interactions

The interplay between coevolutionary and population or community dynamics is currently the focus of much empirical and theoretical consideration. Here, we develop a simulation model to study the coevolutionary and population dynamics of a hypothetical host-parasitoid interaction. In the model, host resistance and parasitoid virulence are allowed to coevolve. We investigate how trade-offs associated with these traits modify the system's coevolutionary and population dynamics. The most important influence on these dynamics comes from the incorporation of density-dependent costs of resistance ability. We find three main outcomes. First, if the costs of resistance are high, then one or both of the players go extinct. Second, when the costs of resistance are intermediate to low, cycling population and coevolutionary dynamics are found, with slower evolutionary changes observed when the costs of virulence are also low. Third, when the costs associated with resistance and virulence are both high, the hosts trade-off resistance against fecundity and invest little in resistance. However, the parasitoids continue to invest in virulence, leading to stable host and parasitoid population sizes. These results support the hypothesis that costs associated with resistance and virulence will maintain the heritable variation in these traits found in natural populations and that the nature of these trade-offs will greatly influence the population dynamics of the interacting species.

Tritrophic effects of organic and conventional fertilisers on a cereal-aphid-parasitoid system

The impact of parasitoids on pests varies between conventional and low-intensity agricultural systems. Although the impacts on parasitoid natural enemies of many practices within these agricultural systems are well understood, the role of fertilisers has been less well studied. The effects of organic-based and conventional fertilisers on Hordeum vulgare L. (Poaceae), the aphid Metopolophium dirhodum Walker (Hemiptera: Aphididae), and its parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) was investigated using cage release experiments and measures of aphid and parasitoid fitness were taken. Barley tiller number and aphid weight were increased by fertilisers, particularly under conventional treatments. Adult parasitoid size correlated positively with that of the host, M. dirhodum, whereas percentage parasitism was not affected by fertiliser treatment or host size. The results suggest that the increased parasitoid impact observed in some low-intensity or organic systems is not a direct result of fertiliser treatment. Our results indicate that fertiliser treatments that improve cereal-aphid fitness will improve parasitoid fitness as measured by parasitoid size but may not influence percentage parasitism.

Effects of organic and conventional fertiliser treatments on host selection by the aphid parasitoid Diaeretiella rapae

The type and quantity of fertilizer supplied to a crop will differ between organic and conventional farming practices. Altering the type of fertilizer a plant is provided with can influence a plant’s foliar nitrogen levels, as well as the composition and concentration of defence compounds, such as glucosinolates. Many natural enemies of insect herbivores can respond to headspace volatiles emitted by the herbivores’ host plant in response to herbivory. We propose that manipulating fertilizer type may also influence the headspace volatile profiles of plants, and as a result, the tritrophic interactions that occur between plants, their insect pests and those pests’ natural enemies. Here, we investigate a tritrophic system consisting of cabbage plants, Brassica oleracea, a parasitoid, Diaeretiella rapae, and one of its hosts, the specialist cabbage aphid Brevicoryne brassicae. Brassica oleracea plants were provided with either no additional fertilization or one of three types of fertilizer: Nitram (ammonium nitrate), John Innes base or organic chicken manure. We investigated whether these changes would alter the rate of parasitism of aphids on those plants and whether any differences in parasitism could be explained by differences in attractivity of the plants to D. rapae or attack rate of aphids by D. rapae. In free-choice experiments, there were significant differences in the percentage of B. brassicae parasitized by D. rapae between B. oleracea plants grown in different fertilizer treatments. In a series of dual-choice Y-tube olfactometry experiments, D. rapae females discriminated between B. brassicae-infested and undamaged plants, but parasitoids did not discriminate between similarly infested plants grown in different fertilizer treatments. Correspondingly, in attack rate experiments, there were no differences in the rate that D. rapae attacked B. brassicae on B. oleracea plants grown in different fertilizer treatments. These findings are of direct relevance to sustainable and conventional farming practices.

Complex latitudinal variation in the morphology of the kleptoparasitic spider Argyrodes kumadai associated with host use and climatic conditions

We examined complex geographical patterns in the morphology of a kleptoparasitic spider, Argyrodes kumadai, across its distributional range in Japan. To disentangle biotic and abiotic factors underlying morphological variation, latitudinal trends were investigated in two traits, body size and relative leg length, across separate transition zones for host use and voltinism. Statistical analyses revealed complex sawtooth clines. Adult body size dramatically changed at the transition zones for host use and voltinism, and exhibited a latitudinal decline following the converse to Bergmann’s cline under the same host use and voltinism in both sexes. A similar pattern was observed for relative leg length in females but not in males. A genetic basis for a part of observed differences in morphology was supported by a common-garden experiment. Our data suggest that local adaptation to factors other than season length such as resource availability (here associated with host use) obscures underlying responses to latitude.