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Certain species of parasites have the apparent ability to alter the behaviour of their host in order to facilitate the completion of their own life cycle. While documented in hairworms (phylum Nematomorpha), the ability for mermithid parasites (from the sister phylum Nematoda) to force hosts to enter water remains more enigmatic. Here, we present the first experimental evidence in a laboratory setting that an insect which normally never enters open water (the European earwig Forficula auricularia) will readily enter the water when infected with a mermithid nematode (Mermis nigrescens). Only adult mermithids appear capable of inducing this polarising shift in behaviour, with mermithid length being a very strong predictor of whether their host enters water. However, mermithid length was only weakly associated with how long it took an earwig to enter water following the beginning of a trial. Considering the evidence presented here and its alignment with a proteomic investigation on the same host-parasite system, this study provides strong evidence for adaptive behavioural manipulation and a foundational system for further behavioural and mechanistic exploration.

The immune response affects host's survival and reproductive success. Insurmountable immune function has not evolved because it is costly and there is a trade-off between other life-history traits. In previous studies several factors such as diet and temperature have been proposed to cause interpopulation differences in immune response. Moreover, the insect immune system may be functionally more protective upon secondary exposure, thus infection history may associate with the immune response. Here we measured how geographical location and parasite burden is related to variation in immune response between populations. We included 13 populations of the Northern Damselfly Coenagrion hastulatum (Odonata: Coenagrionidae) in Finland over a latitudinal range of 880 km to this study. We found that water mites associated strongly with the immune response at interpopulation level: the more the mites, the higher the immune response. Also, in an alternative model based on AIC, latitude and individual size associated with the immune response. In turn, endoparasitic gregarines did not affect the immune response. To conclude, a positive interpopulation association between the immune response and the rate of water mite infection may indicate (i) local adaptation to chronic parasite stress, (ii) effective 'induced' immune response against parasites, or (iii) a combined effect of both of these.

Host discrimination by immature host-seeking endoparasites is a complex and somewhat unexplored topic. In the case of multiple infections, conflicts among conspecifics may occur to monopolize space and resources in the same host. Two or more 1st instar larvae of Xenos vesparum (Strepsiptera, Stylopidae) may enter into a Polistes dominulus (Hymenoptera, Vespidae) larva and develop together until the adult stage of both parasite and host. We carried out a screening of mitochondrial haplotypes in X. vesparum individuals extracted from superparasitized wasps taken in 5 naturally infected nests from different areas of Tuscany (Italy), to assess whether non-sibling parasites may infect the same colony and host. In total, we obtained 12 different haplotypes out of 122 genotyped individuals of both sexes: 17 of 34 superparasitized wasps hosted parasites that originated from females differing in their haplotypes. To date, this is the first described case of superparasitism with non-sibling host-seeking larvae infecting a single individual hymenopteran host. In addition, at least in heavily infected colonies, there is evidence of a male-biased sex-ratio and synchronous development of the parasites, regardless of their haplotypes. Finally, the distribution of haplotypes per nest is consistent with either phoretic infection or larvipositing on nests by means of superparasitized wasps.

Polistes dominulus are social wasps which are the host of the strepsipteran endoparasite Xenos vesparum. In the hibernating phase, unparasitized and parasitized wasps leave natal nests and aggregate together in sheltered quarters. In aggregations, wasps are socially active, and some individuals perform helping behaviour. Here we investigated if castrated parasitized wasps perform worker tasks in mixed aggregations. Moreover, by gas chromatography and mass spectrometry, we examined the cuticular hydrocarbons of unparasitized and parasitized wasps to evaluate if the infection alters the composition of cuticular waxes that are recognition cues in social insects. In clusters, infected females do not perform helping behaviour and they are less active than unparasitized wasps. Cuticular hydrocarbons are slightly differentiated between unparasitized and parasitized wasps but, generally, unparasitized wasps are more similar to wasps infected by Xenos females compared to wasps infected by Xenos males. Wasps infected by Xenos males do not usually survive the winter. This chemical similarity is probably a consequence of the similar physiological condition of unparasitized and female-affected Polistes wasps. At this stage, it is difficult to affirm whether these modifications are a true parasite manipulation or a consequence of infection.

Avian trypanosomes are widespread parasites of birds, the transmission of which remains mostly unclear, with various blood-sucking insects mentioned as possible vectors. A search for vectors of trypanosomes of sparrowhawk (Accipiter nisus), buzzard (Buteo buteo), lesser-spotted eagle (Aquila pomarina) and kestrel (Falco tinnunculus) was performed in Czech and Slovak Republics. Black flies (Eusimulium spp.), hippoboscid flies (Ornithomyia avicularia), mosquitoes (Culex pipiens pipiens) and biting midges (Culicoides spp.), trapped while attempting to feed on raptor nestlings, were found to contain trypanosomatids in their intestine. Trypanosomes from the raptors and blood-sucking insects were isolated, and their 18S rRNA sequences were used for species identification and for the inference of intra- and interspecific relationships. Together with the trypanosome isolated from a black fly, the bird trypanosomes formed a well-supported Trypanosoma avium clade. The isolates derived from hippoboscid flies and mosquitoes are most likely also avian trypanosomes infecting birds other than the studied raptors. Analysis of the kinetoplast, that has features characteristic for the avian trypanosomes (minicircle size; dimensions of the kinetoplast disc), provided further evidence for the identification of vectors. It is suggested that all trypanosomes isolated from raptors included in this study belong to the T. avium complex and are transmitted by the ornithophilic simuliids such as Eusimulium securiforme.

The frequency distribution of first generation, Steinernema feltiae Filipjev parasitic stages was over-dispersed with the majority of hosts containing few or no parasitic stages, whilst a few hosts contained a great many. Because of high extraction efficiency, the frequency distributions of the parasitic stages and the infective stages in the soil were assumed to be directly related. To explain the frequency distribution of the parasites it was therefore necessary to account for the frequency distribution of the S. feltiae infective stages in the soil. The infective stages were spatially aggregated into 30 cm diameter patches at the site of host death. These patches were randomly distributed approximately 1 m apart. At the 1 m scale, the pooled counts of infective stages were randomly distributed. Thus, in contrast to the frequency distributions, the spatial structuring of S. feltiae changed with the spatial scale of the interaction. This dynamic spatial structuring means that the majority of samples taken would contain few or no infective stages, whilst a few soil samples would contain a great many. Thus, the spatial structuring of the infective stages generates the over-dispersed frequency distribution of the S. feltiae in the soil. Hosts, encountering infective stages from this spatial distribution will, therefore, show an over-dispersed frequency distribution of S. feltiae parasitic stages.

Existing models of the basic case reproduction number (R0) for vector-borne diseases assume (i) that the distribution of vectors over the susceptible host species is homogeneous and (ii) that the biting preference for the susceptible host species rather than other potential hosts is a constant. Empirical evidence contradicts both assumptions, with important consequences for disease transmission. In this paper we develop an Ideal Free Distribution (IFD) model of host choice by blood-sucking insects, predicated on the argument that vectors must have evolved to choose the least defensive hosts in order to maximize their feeding success. From a re-analysis of existing data, we demonstrate that the interference constant, m, of the IFD can vary between host species. As a result, the predicted distribution of insects over hosts has 2 desirable and intuitively plausible behaviours: that it is heterogeneous both within and between host species; and that the intensity of heterogeneity varies with host and vector density. When the IFD model is incorporated into R0, the relationship with the vector:host ratio becomes non-linear. If correct, the IFD could add considerable realism to models which seek to predict the effect of these ecological parameters on disease transmission as they vary naturally (e.g. through seasonality in vector density or host population movement) or as a consequence of artificial manipulation (e.g. zooprophylaxis, vector control). It raises the possibility of targeting transmission hot spots with greater accuracy and concomitant reduction in control effort. The robustness of the model to simplifying assumptions is discussed.

Host-parasitoid interactions are abundant in nature and offer great scope for the study of coevolution. A particularly fertile area is the interaction between internal feeding parasitoids and their hosts. Hosts have evolved a variety of means of combating parasitoids, in particular cellular encapsulation, while parasitoids have evolved a wide range of countermeasures. Studies of the evolution of host resistance and parasitoid virulence are reviewed, with an emphasis on work involving Drosophila and its parasitoids. Genetic variation in both traits has been demonstrated using isofemale line and artificial selection techniques. Recent studies have investigated the fitness costs of maintaining the ability to resist parasitoids, the comparative fitness of flies that have successfully defended themselves against parasitoids, and the degree to which resistance and virulence act against one or more species of host or parasitoid. A number of studies have examined geographical patterns, and sought to look for local adaptation; or have compared the traits across a range of species. Finally, the physiological and genetic basis of change in resistance and virulence is being investigated. While concentrating on Drosophila, the limited amount of work on different systems is reviewed, and other possible areas of coevolution in host-parasitoid interactions are briefly discussed.

The literature contains many examples of changes induced by parasites in the behaviour and/or other phenotypic traits of insects. From an evolutionary perspective, the nature of these changes is usually difficult to assess. Parasite-induced changes in host behaviour can be adaptations of either host or parasite, or they can be mere pathological consequences of infection. Of the many criteria and experimental tests necessary to distinguish between adaptations and non-selected consequences, two are particularly important: the demonstration of fitness benefits for either host or parasite associated with the behavioural change, and the elucidation of the proximate mechanism responsible for the behavioural change. Another approach can serve to identify adaptive changes in behavior: mapping specific behavioural alterations on a phylogeny of either hosts or parasites. The usefulness of this approach is illustrated with two examples, acanthocephalan-cockroach associations and insect-fungus associations. The adaptive nature of parasite-induced behavioural changes will always be difficult to evaluate because they are the product of two distinct but interacting genotypes. However, experimental and phylogenetic approaches can provide valuable insights into the evolutionary history of insect-parasite interactions.

Host fertility is often curtailed as a result of parasitic infection. The hypothesis that this may confer an adaptive advantage upon the symbionts if nutrients are directed from reproduction and made available for host/parasite maintenance is explored. The suggestion is made that an understanding of the mechanisms underlying the pathophysiology of fecundity reduction may shed light upon the evolutionary implications of this strategy for both parasite and host. To illustrate this the down-regulation of egg production is explored with reference to a particular model system, the association between metacestodes of the rat tapeworm, Hymenolepis diminuta and the mealworm beetle, Tenebrio molitor. Several aspects of host reproductive behavior and physiology are affected by infection in this association, including vitellogenesis. Metacestodes directly inhibit the fat body synthesis of vitellogenin in a stage-specific, density-dependent manner. This inhibition is likely to be orchestrated by a modulator molecule, produced by the parasite. In the ovarian follicles, juvenile hormone III binding to a specific follicular membrane-binding protein is inhibited in infected beetles, resulting in the down-regulation of a cascade of events which enables vitellogenin to pass into the developing oocyte. Data to support the proposed existence of a parasite-induced antigonadotrophin, of host origin, are discussed. Evidence that similar mechanisms operate in Plasmodium-infected anopheline mosquitoes and Onchocerca-infected blackflies is presented in support of the possibility that a parasite-induced reduction in host reproductive fitness is an adaptive strategy and an assessment of who is manipulating whom is made.

Understanding the temporal and spatial distribution of entomopathogenic nematodes is essential for determining the role of these insect parasites in soil communities and ultimately for their use in suppression of pest insect populations. We measured the vertical and horizontal distribution of endemic populations of entomopathogenic nematodes (Steinernema carpocapsae and Heterorhabditis bacteriophora) in turfgrass. Vertical distribution was determined by taking soil cores every 3 h from 05.00 to 23.00 h, over 4 days, and dividing the cores into 8, 1 cm deep sections. Steinernema carpocapsae was recovered primarily near the soil surface: 50% of positive sections were recovered in the thatch or first 1 cm of soil. S. carpocapsae recovery was lower during the middle of the day and none were recovered in the upper section. H. bacteriophora was recovered uniformly throughout the top 8 cm of soil and its vertical distribution did not change over the course of the day. Horizontal distribution was measured as the number of nematodes recovered from cores taken from 12 randomly selected 0.3 x 0.8 m sections from within four 15.3 x 15.3 m plots. Samples were collected biweekly over a 9-month period. H. bacteriophora had a patchier distribution than S. carpocapsae and both nematode species had more patchy distributions then their potential hosts. Our results support the hypothesis that these two species of nematode utilize different foraging strategies; S. carpocapsae primarily a surface adapted ambusher and H. bacteriophora as a cruise forager.

Three satellite DNAs previously isolated from the entomopathogenic nematodes Steinernema carpocapsae, Heterorhabditis bacteriophora and Heterorhabditis indicus give hybridization signals only with the S. carpocapsae, H. bacteriophora and H. indicus populations tested, indicating that these satellite sequences are species-specific. Because of their reiteration and their variabilities, we have shown that these sequences are able to discriminate at the interspecific level between the Steinernema and Heterorhabditis species, but also at the intraspecific level between S. carpocapsae strains. Furthermore, in simple squashed nematode experiments, we are able to unambiguously identify S. carpocapsae, H. bacteriophora and H. indicus populations. This last procedure is effective even on a single infective juvenile, with the main advantage that it avoids time-consuming DNA extractions.

The role of the larva of the alder fly Sialis lutaria as a host for Neoechinorhynchus rutili (Acanthocephala) has been investigated. Brown trout (Salmo trutta) infected with N. rutili were found to be feeding on S. lutaria larvae which contained juvenile acanthocephalans in their haemocoels. These acanthocephalans have been identified as N. rutili. A previous report of this host-parasite relationship indicated some form of host response to the larval acanthocephalan. Stained wax sections of S. lutaria larvae infected with N. rutili did not show any evidence of either a host response or encapsulation. Samples of S. lutaria larvae revealed an interesting pattern of infection. Only larvae of a particular size range were infected. An experiment to infect commercially obtained rainbow trout (Salmo gairdneri) with N. rutili by feeding them with infected S. lutaria larvae was successful. Since S. lutaria larvae are known to be prey items for trout it appears that this route of infection could commonly occur in the natural environment. Larval S. lutaria is considered as an additional intermediate host to several ostracod species which have been shown to play a role in the transmission of N. rutili.