Functional diversity of ground beetles improved aphid control but did not increase crop yields on European farms.

Land-use intensification is often associated with a decline in functional diversity, potentially undermining the provision of ecosystem services. However, how changes in traits affect ecosystem processes remains poorly understood. Variation in trait values among species in a community may drive ecosystem processes. Alternatively, the mass ratio hypothesis proposes that trait values of the dominant species in a local community are related to ecosystem processes. Using data from 159 farms in six European countries, we quantified the impact of local and landscape-level land-use intensity on ground beetles as pest control agents. We then assessed the extent to which functional diversity and community-weighted mean trait values relate to pest control and cereal yield. In addition, we assessed how the responses to land use and the effects of different species on pest control and yield varied with their traits to compare the relative impact of the traits studied. Functional diversity of ground beetles improved aphid removal, but did not translate into higher crop yields. Pest control of aphids was enhanced by a higher proportion of smaller, mobile ground beetles with a preference for the vegetation layer. Smaller, predatory ground beetles in communities improved crop yield. The magnitude of responses to land-use intensification and the effects on pest control and yield were more strongly influenced by body size than other traits. Our study provides evidence that reduced management intensity can improve pest control by supporting small-sized, macropterous ground beetles. In contrast to the claims of ecological intensification, our joint analysis of the direct effects of land use on yield and indirect effects via functional diversity of ground beetles and pest control suggests that ecosystem services by ground beetles cannot compensate for the yield gap due to a reduction in land-use intensity.

Contrasting effects of aquatic subsidies on a terrestrial trophic cascade.

Subsidies from adjacent ecosystems can alter recipient food webs and ecosystem functions, such as herbivory. Emerging aquatic insects from streams can be an important prey in the riparian zone. Such aquatic subsidies can enhance predator abundances or cause predators to switch prey, depending on the herbivores. This can lead to an increase or decrease of in situ herbivores and herbivory. We examined the effects of aquatic subsidies on a simplified terrestrial food web consisting of two types of herbivores, plants and predators (spiders). In our six-week experiment, we focused on the prey choice of the spiders by excluding predator immigration and reproduction. In accordance with predator switching, survival of leafhoppers increased in the presence of aquatic subsidies. By contrast, the presence of aquatic subsidies indirectly reduced weevils and herbivory. Our study shows that effects of aquatic subsidies on terrestrial predators can propagate through the food web in contrasting ways. Thereby, the outcome of the trophic cascade is determined by the prey choice of predators.

Risk of spider predation alters food web structure and reduces local herbivory in the field.

Predators can indirectly enhance plant performance via herbivore suppression, with both prey consumption and changes in prey traits (e.g. changes in foraging behaviour) contributing to the reduction in herbivory. We performed a field experiment to determine the extent of such non-consumptive effects which consisted of repeatedly placing spiders (Pisaura mirabilis) on enclosed plants (Urtica dioica) for cue deposition. Control plants were enclosed in the same way but without spiders. After cue deposition, the enclosures were removed to allow arthropods to colonize the plants and feed on them. Arthropods were removed from the plants before the subsequent spider deposition or control enclosure. During six cycles of enclosure, we quantified leaf damage on the plants. After a seventh cycle, the colonizing arthropods were sampled to determine community composition in relation to the presence/absence of spider cues. We found that the presence of chemotactile spider cues reduced leaf damage by 50 %. In addition, spider cues led to changes in the arthropod community: smaller spiders avoided plants with spider cues. In contrast, the aphid-tending ant Myrmica rubra showed higher recruitment of workers on cue-bearing plants, possibly to protect aphids. Our results show that the risk of spider predation can reduce herbivory on wild plants and also demonstrate that non-consumptive effects can be particularly strong within the predator guild.

Comparison of native and non-native predator consumption rates and prey avoidance behavior in North America and Europe.

Novel predator-prey interactions can contribute to the invasion success of non-native predators. For example, native prey can fail to recognize and avoid non-native predators due to a lack of co-evolutionary history and cue dissimilarity with native predators. This might result in a competitive advantage for non-native predators. Numerous lady beetle species were globally redistributed as biological control agents against aphids, resulting in novel predator-prey interactions. Here, we investigated the strength of avoidance behavior of the pea aphid (Acyrthosiphon pisum) toward chemical cues of native lady beetles and non-native Asian Harmonia axyridis and European Coccinella septempunctata and Hippodamia variegata in North America, hypothesizing that cues of non-native lady beetles induce weaker avoidance behavior than cues of co-evolved native lady beetles. Additionally, we compared aphid consumption of lady beetles, examining potential predation advantages of non-native lady beetles. Finally, we compared cue avoidance behavior between North American and European pea aphid populations and aphid consumption of native and non-native lady beetles in North America and Europe. In North America, pea aphids avoided chemical cues of all ladybeetle species tested, regardless of their origin. In contrast to pea aphids in North America, European pea aphids did not avoid cues of the non-native H. axyridis. The non-native H. axyridis and C. septempunctata were among the largest and most voracious lady beetle species tested, on both continents. Consequently, in North America non-native lady beetle species might have a competitive advantage on shared food resources due to their relatively large body size, compared to several native American lady beetle species. In Europe, however, non-native H. axyridis might benefit from missing aphid cue avoidance as well as a large body size. The co-evolutionary time gap between the European and North American invasion of H. axyridis likely explains the intercontinental differences in cue avoidance behavior and might indicate evolution in aphids toward non-native predators.

Sublethal effects of imidacloprid on interactions in a tritrophic system of non-target species.

Imidacloprid is one of the most used insecticides worldwide, but is highly toxic to non-target arthropods. Effects of sublethal imidacloprid intoxication can potentially propagate in food webs, yet little is known about the impact on non-target populations and communities. We investigated short-term sublethal toxicity of imidacloprid in a tritrophic model system of wild strawberry Fragaria vesca, wood cricket Nemobius sylvestris and nursery web spider Pisaura mirabilis. Strawberries were treated two times with 0mg (control), 1mg (low rate) and 10mg (high rate) of Confidor® WG 70 and crickets were allowed to feed on them. In four lab experiments, we quantified the impact of imidacloprid on leaf damage, growth, behaviour and survival of crickets. The high imidacloprid rate reduced feeding, mass gain, thorax growth and mobility in crickets compared to the control, while mortality was similarly low in all treatments. The low rate reduced mass gain only. Cricket survival of spider predation was higher in the low rate treatment than in the control. Overall, herbivory and predation were reduced at sublethal imidacloprid rates in a non-target organism, three-level food chain, which demonstrates possible propagation of sublethal effects through trophic interactions.