Colletes hederae bees are equally attracted by visual and olfactory cues of inconspicuous Hedera helix flowers

Wild bees are heavily declining worldwide except for a few species, such as Colletes hederae, which is spreading in its distribution throughout Europe. Colletes hederae mainly forages on ivy (Hedera helix) which is widespread in Europe and the plants' availability is thought to contribute to the successful spread of C. hederae. A rapid location of the plants using visual and/or olfactory floral cues would allow the bee to efficiently forage. Beside bee visitors, the flowers attract a high variety of other insects, such as Vespula wasps that were recently investigated regarding their floral-cue preferences. The aim of this study was to investigate the communication between C. hederae and its H. helix host flowers, and to compare the results with that previously obtained with V. germanica wasps. We identified headspace compounds detectable by the bees using gas chromatography coupled to electroantennography (GC-EAD) and performed behavioral experiments to both compare the attractiveness of visual and olfactory floral cues and to determine the attractiveness of a synthetic mixture composed of physiologically active compounds. In the GC-EAD analyses, bees responded to 15 flower-specific compounds of various chemical classes, of which 4-oxoisophorone, (E)-linalool-oxide furanoid, and acetophenone were the most abundant in the floral scent. In the bioassays, visual and olfactory flower cues were equally attractive for bees, but a combination of both cues was needed to elicit not only approach responses but also landings. A synthetic mixture of the EAD-active compounds was attractive to the bees, but to a lesser extent than the natural scent of H. helix flowers. The bees' integrations of different floral-cue modalities in its search image and its strong antennal responses elicited by various floral scent compounds make C. hederae highly effective in finding its host flowers. In comparison to V. germanica wasps, the bees relied stronger on visual cues than the wasps do, but both species showed the highest attraction when presented with a combination of the cues.