The release of biological control agents has been an important means of controlling invasive species for over 150 years. While these releases have led to the sustainable control of over 250 invasive pest and weed species worldwide, a minority have caused environmental harm. A growing recognition of the risks of biological control led to a focus on risk assessment beginning in the 1990s along with a precipitous decline in releases. While this new focus greatly improved the safety of biological control, it came at the cost of lost opportunities to solve environmental problems associated with invasive species. A framework that incorporates benefits and risks of biological control is thus needed to understand the net environmental effects of biological control releases. We introduce such a framework, using native biodiversity as the common currency for both benefits and risks. The model is based on interactions among four categories of organisms: (1) the biological control agent, (2) the invasive species (pest or weed) targeted by the agent, (3) one or more native species that stand to benefit from the control of the target species, and (4) one or more native species that are at risk of being harmed by the released biological control agent. Conservation values of the potentially benefited and harmed native species are incorporated as well, and they are weighted according to three axes: vulnerability to extinction, the ecosystem services provided, and cultural significance. Further, we incorporate the potential for indirect risks to native species, which we consider will result mainly from the ecological process of agent enrichment that may occur if the agent exploits but does not control the target pest or weed. We illustrate the use of this framework by retrospectively analyzing the release of the vedalia beetle, Novius (= Rodolia) cardinalis, to control the cottony cushion scale, Icerya purchasi, in the Galapagos Islands. While the framework is particularly adaptable to biological control releases in natural areas, it can also be used in managed settings, where biological control protects native species through the reduction of pesticide use.
Bacterial symbionts in arthropods are common, vary in their effects, and can dramatically influence the outcome of biological control efforts. Macrolophus pygmaeus (Heteroptera: Miridae), a key component of biological control programs, is mainly predaceous but may also display phytophagy. M. pygmaeus hosts symbiotic Wolbachia, which induce cytoplasmic incompatibility, and two Rickettsia species, R. bellii and R. limoniae, which are found in all individuals tested. To test possible involvement of the two Rickettsia species in the feeding habits of M. pygmaeus, we first showed that the microbiome of the insect is dominated by these three symbionts, and later described the distribution pattern of the two Rickettsia species in its digestive system. Although both Rickettsia species were located in certain gut bacteriocyes, in caeca and in Malpighian tubules of both sexes, each species has a unique cellular occupancy pattern and specific distribution along digestive system compartments. Infrequently, both species were found in a cell. In females, both Rickettsia species were detected in the germarium, the apical end of the ovarioles within the ovaries, but not in oocytes. Although the cause for these Rickettsia distribution patterns is yet unknown, it is likely linked to host nutrition while feeding on prey or plants.
In agricultural landscapes, uncultivated habitat patches may have a focal role in supporting communities of ecosystem service providers. However, little is known on the variances among different types of uncultivated habitat patches in providing resources and maintaining populations of these beneficial organisms. We studied wild bee communities in natural and semi-natural uncultivated patches embedded in semi-arid Mediterranean agricultural landscapes. We investigated the effects of local- and landscape-scale land-use characteristics, as well as their interactions, on bee diversity, functional composition, and forage and nesting resources. Most bee community parameters were affected by both local- and landscape-scale characteristics, but no significant interactions were found among the scales. Local land-use effects were related primarily to overall plant cover, and to the abundance and richness of flowering plants. Landscape effects, mostly limited to a 400 m range, were varied. The abundance of focal crop pollinators varied considerably between patch type and pollinator species. The different types of uncultivated habitats maintain complementary bee and flower communities. Our findings show the important role of uncultivated habitat patches in providing floral and nesting resources for bees, and creating resource-landscapes that can support wild bee communities and crop pollination services in Mediterranean agricultural landscapes.
Animals of a wide range of taxonomic groups mix various food sources to achieve a nutritionally balanced diet. The strategies they adopt to balance multiple nutrients depend on their availability in the environment. Behavioural and physiological adaptations to forage for nutrient-differing food sources have rarely been investigated in respect to nutrient availability in the environment. We developed a simulation model to explore the strategy consumers should adopt in response to the abundance of two nutritionally complementary food types. Results show that (1) consumers should invest more effort in detecting the scarce resource; (2) there is an optimized negative relationship between effort foragers should allocate to find the two types of food; (3) consumers should exhibit higher selectivity when the proportion of food types in the habitat deviates from their optimal ratio in the diet. These findings have important implications for pest control using predators that benefit from plant-based food supplements.