Nectar cardenolides and floral volatiles mediate a specialized wasp pollination system.

Specialization in plant pollination systems can arise from traits that function as filters of flower visitors. This may involve chemical traits such as floral volatiles that selectively attract favoured visitors and non-volatile nectar constituents that selectively deter disfavoured visitors through taste or longer-term toxic effects or both. We explored the functions of floral chemical traits in the African milkweed Gomphocarpus physocarpus, which is pollinated almost exclusively by vespid wasps, despite having nectar that is highly accessible to other insects such as honeybees. We demonstrated that the nectar of wasp-pollinated G. physocarpus contains cardenolides that had greater toxic effects on Apis mellifera honeybees than on Vespula germanica wasps, and also reduced feeding rates by honeybees. Behavioural experiments using natural compositions of nectar compounds showed that these interactions are mediated by non-volatile nectar chemistry. We also identified volatile compounds with acetic acid as a main component in the floral scent of G. physocarpus that elicited electrophysiological responses in wasp antennae. Mixtures of these compounds were behaviourally effective for attraction of V. germanica wasps. The results show the importance of both volatile and non-volatile chemical traits as filters that lead to specialization in plant pollination systems.

Land-use affects pollinator-specific resource availability and pollinator foraging behaviour.

Land-use management is a key factor causing pollinator declines in agricultural grasslands. This decline can not only be directly driven by land-use (e.g., habitat loss) but also be indirectly mediated through a reduction in floral resource abundance and diversity, which might in turn affect pollinator health and foraging. We conducted surveys of the abundance of flowering plant species and behavioural observations of two common generalist pollinator species, namely the bumblebee Bombus lapidarius and the syrphid fly Episyrphus balteatus, in managed grasslands of variable land-use intensity (LUI) to investigate whether land-use affects (1) resource availability of the pollinators, (2) their host plant selection and (3) pollinator foraging behaviour. We have found that the floral composition of plant species that were used as resource by the investigated pollinator species depends on land-use intensity and practices such as mowing or grazing. We have also found that bumblebees, but not syrphid flies, visit different plants depending on LUI or management type. Furthermore, LUI indirectly changed pollinator behaviour via a reduction in plot-level flower diversity and abundance. For example, bumblebees show longer flight durations with decreasing flower cover indicating higher energy expenditure when foraging on land-use intensive plots. Syrphid flies were generally less affected by local land use, showing how different pollinator groups can differently react to land-use change. Overall, we show that land-use can change resource composition, abundance and diversity for pollinators, which can in turn affect pollinator foraging behaviour and potentially contribute to pollinator decline in agricultural grasslands.

Floral Trait Preferences of Three Common wild Bee Species.

The interaction between bees and flowering plants is mediated by floral cues that enable bees to find foraging plants. We tested floral cue preferences among three common wild bee species: Lasioglossum villosulum, Osmia bicornis, and Bombus terrestris. Preferences are well studied in eusocial bees but almost unknown in solitary or non-eusocial generalist bee species. Using standardized artificial flowers altered in single cues, we tested preferences for color hue, achromatic contrast, scent complexity, corolla size, and flower depth. We found common attractive cues among all tested bees. Intensively colored flowers and large floral displays were highly attractive. No preferences were observed in scent complexity experiments, and the number of volatiles did not influence the behavior of bees. Differing preferences were found for color hue. The specific behaviors were probably influenced by foraging experience and depended on the flower choice preferences of the tested bee species. In experiments testing different flower depths of reward presentation, the bees chose flat flowers that afforded low energy costs. The results reveal that generalist wild bee species other than well-studied honeybees and bumblebees show strong preferences for distinct floral cues to find potential host plants. The diverse preferences of wild bees ensure the pollination of various flowering plants.

Forest structure, plants, arthropods, scale, or birds' functional groups: What key factor are forest birds responding to?

Forest birds respond to a diverse set of environmental factors, including those altered by forest management intensity, such as resource and habitat availability in the form of food or nesting sites. Although resource/habitat availability and bird traits likely mediate responses of bird diversity to global change drivers, no study has assessed the direct and indirect effects of changes in forest management and traits on bird assemblages jointly at large spatial scales. In this context the questions remain whether (1) the birds' response to forest management changes through alterations in structural properties and/or food availability, or (2) if birds' eco-morphological traits act as environmental filters in response to environmental factors. We audio-visually recorded birds at 150 forest plots in three regions of Germany and assessed the forest structure (LiDAR) as well as the diversity of the herbaceous layer and diversity and biomass of arthropods. We further assessed eco-morphological traits of the birds and tested if effects on bird assemblages are mediated by changes in eco-morphological traits' composition. We found that abundance and species numbers of birds are explained best by models including the major environmental factors, forest structure, plants, and arthropods. Eco-morphological traits only increased model fit for indirect effects on abundance of birds. We found minor differences between the three regions in Germany, indicating spatial congruency of the processes at the local and regional scale. Our results suggest that most birds are not specialized on a particular food type, but that the size, diversity and species composition of arthropods are important. Our findings question the general view that bird traits adapt to the resources available.