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Agricultural intensification can impact agrobiodiversity in several ways such as in terms of population densities, community composition and food web interactions across all trophic levels. This effect can be investigated at two scales: field-scale and landscape scale. Here it was assessed how the impact of fertilisation (within field) and landscape complexity (within landscape) impact cereal aphid-primary parasitoid-secondary parasitoid systems in winter wheat in Germany. A newly developed molecular technique was used to quantify species-specific linkages between aphids, primary parasitoids and secondary parasitoids sampled in fertilised and unfertilised plots in either simple or complex structured landscapes. The results show a stronger effect of fertilisation than landscape complexity on the groups: fertilisation positively affected the crop plants while it negatively affected both the density of the cereal aphid Sitobion avenae and its primary parasitism rates whereas no effect on the level of secondary parasitism rates was observed. Landscape complexity had no effect on plants, aphids, as well as on primary parasitism rate. In case of secondary parasitism rate there was an effect in interaction with sampling date. Field identity accounted for the strongest effect on parasitoid community composition (10.6% of the variance) from all tested variables, while fertilisation and landscape complexity had almost no effect (1.1% and no effect). Nevertheless, a weak cascading effect of both environmental factors could be observed as the primary-secondary parasitoid network structure responded to both. However, these observed effects on food webs strongly depended on species identity, highlighting the need of species-level food web assessment.

Facultative bacterial endosymbionts can protect their aphid hosts from natural enemies such as hymenopteran parasitoids. As such, they have the capability to modulate interactions between aphids, parasitoids and hyperparasitoids. However, the magnitude of these effects in natural aphid populations and their associated parasitoid communities is currently unknown. Moreover, environmental factors such as plant fertilization and landscape complexity are known to affect aphid–parasitoid interactions but it remains unclear how such environmental factors affect the interplay between aphids, parasitoids and endosymbionts.Here, we tested whether facultative endosymbionts confer protection to parasitoids in natural populations of the English grain aphid, Sitobion avenae, and if this is affected by plant fertilization and landscape complexity. Furthermore, we examined whether the effects of facultative endosymbionts can cascade up to the hyperparasitoid level and increase primary‐hyperparasitoid food web specialization.Living aphids and mummies were collected in fertilized and unfertilized plots within 13 wheat fields in Central Germany. We assessed the occurrence of primary parasitoid, hyperparasitoid and endosymbiont species in aphids and mummies using a newly established molecular approach.Facultative endosymbiont infection rates were high across fields (~80%), independent of whether aphids were parasitized or unparasitized. Aphid mummies exhibited a significantly lower share of facultative endosymbiont infection (~38%). These findings suggest that facultative endosymbionts do not affect parasitoid oviposition behaviour, but decrease parasitoid survival in the host. Facultative endosymbiont infection rates were lower in mummies collected from fertilized compared to unfertilized plants, indicating that plant fertilization boosts the facultative endosymbiont protective effect. Furthermore, we found strong evidence for species‐specific and negative cascading effects of facultative endosymbionts on primary and hyperparasitoids, respectively. Facultative endosymbionts impacted parasitoid assemblages and increased the specialization of primary‐hyperparasitoid food webs: these effects were independent from and much stronger than other environmental factors.The current findings strongly suggest that facultative endosymbionts act as a driving force in aphid–parasitoid–hyperparasitoid networks: they shape insect community composition at different trophic levels and modulate, directly and indirectly, the interactions between aphids, parasitoids and their environment.

As food items can vary in nutrient composition, diet selectivity can be an important trait to optimize nutrition. Most parasitoids depend as adults on various sugar-rich food sources such as nectar and honeydew. Nectar is often superior as a food source to honeydew, yielding higher parasitoid longevity. Little is known, however, of a preference parasitoids may have for one of these sugar sources and how this affects variation in their life-history traits. In this study, a combined empirical and modelling approach was adopted. First, it was investigated whether parasitoids show a feeding preference for nectar or honeydew in the laboratory and whether feeding history affects the preference of parasitoids. Second, using a process-based spatially explicit simulation model, it was evaluated how preference for nectar affects parasitoid life-history variables in fields with varying flower densities. The laboratory study revealed that naive parasitoids showed no preference for either nectar or honeydew. However, when parasitoids had fed previously on honeydew and were then given the choice between nectar and honeydew, they preferred feeding on nectar to honeydew (with marginal statistical significance). Model simulations indicated that when nectar sources were present in the field, parasitoids that showed preference for nectar had a 1.3-1.8 times longer life span and parasitised up to 1.4 times more hosts than parasitoids that did not distinguish between honeydew and nectar. These combined empirical and modelling data suggest that aphid parasitoids may be able to discriminate between nectar and honeydew, and that a preference for nectar may enhance their longevity and reproduction in fields where nectar sources are available.

The potential impact of nectar availability for biological pest control is of increasing relevance due to the disappearance of non-crop plants in intensified agricultural landscapes. However, little is known about how floral resources and their spatial distribution in and around crop fields influence cereal aphid parasitoid (Hymenoptera, Aphidiidae) survival and population dynamics. Cereal aphid parasitoids may be less dependent on flowers as they can use honeydew from their aphid hosts as an alternative food resource. However, honeydew appears to be less suitable as a food source for parasitoids when compared to nectar. Here the effects of flower distribution (nectar), aphid abundance (honeydew quantity) and honeydew quality on parasitoid life history parameters were explored using a spatially explicit, individual-based simulation model. In addition, effects of the availability of floral resources in field margins on parasitism levels in winter wheat were investigated in a field study. In the simulation study three flower distributions were compared: (i) flowers restricted to the field margin (most common in agricultural landscapes) or (ii) flowers in strips within the crop, and (iii) random flower distribution in the field. Simulations suggest that nectar feeding increases parasitoid longevity and parasitism at low aphid densities, but only in fields with randomly distributed flowers. In contrast, the spatial distribution of flowers did not affect the parasitoid longevity and parasitism rate at high aphid densities, where parasitoid feeding was restricted to the abundant honeydew resources. In simulations, parasitoid survival and the number of aphids parasitized were also affected by the nutritional quality of the honeydew, with low quality honeydew having a less positive effect on parasitoid longevity and parasitism rate. The field study did not indicate evidence of a significant relationship between flower cover in field margins and parasitism rates in wheat near the margins. This finding is compatible with results from simulations and suggests that honeydew feeding is more prevalent in cereal aphid parasitoids than nectar feeding. In conclusion, nectar may be an important factor for the potential biocontrol success of cereal aphid parasitoids, but most likely only at low aphid densities and especially when flowers are broadly distributed in the field.

Insect parasitoids play a major role in terrestrial food webs as they are highly diverse, exploit a wide range of niches and are capable of affecting host population dynamics. Formidable difficulties are encountered when attempting to quantify host-parasitoid and parasitoid-parasitoid trophic links in diverse parasitoid communities. Here we present a DNA-based approach to effectively track trophic interactions within an aphid-parasitoid food web, targeting, for the first time, the whole community of parasitoids and hyperparasitods associated with a single host. Using highly specific and sensitive multiplex and singleplex polymerase chain reaction, endoparasitism in the grain aphid Sitobion avenae (F) by 11 parasitoid species was quantified. Out of 1061 aphids collected during 12 weeks in a wheat field, 18.9% were found to be parasitized. Parasitoids responded to the supply of aphids, with the proportion of aphids parasitized increasing monotonically with date, until the aphid population crashed. In addition to eight species of primary parasitoids, DNA from two hyperparasitoid species was detected within 4.1% of the screened aphids, with significant hyperparasitoid pressure on some parasitoid species. In 68.2% of the hyperparasitized aphids, identification of the primary parasitoid host was also possible, allowing us to track species-specific parasitoid-hyperparasitoid links. Nine combinations of primary parasitoids within a single host were found, but only 1.6% of all screened aphids were multiparasitized. The potential of this approach to parasitoid food web research is discussed.