Look and touch: multimodal sensory control of flower inspection movements in the nocturnal hawkmoth Manduca sexta.

A crucial stage in the interaction between pollinators and plants is the moment of physical contact between them, known as flower inspection, or handling. Floral guides - conspicuous colour markings, or structural features of flower corollas - have been shown to be important in the inspecting behaviour of many insects, particularly in diurnal species. For the nocturnal hawkmoth Manduca sexta tactile input has an important role in flower inspection, but there is no knowledge about the use of visual floral guides in this behaviour. I carried out a series of experiments to first, evaluate the putative role of floral guides during flower inspection and second, to explore how simultaneous tactile and visual guides could influence this behaviour. Results show that visual floral guides affect flower inspection by M. sexta. Moths confine proboscis placement to areas of higher light reflectance regardless of their chromaticity, but do not appear to show movements in any particular direction within these areas. I also recorded inspection times, finding that moths can learn to inspect flowers more efficiently when visual floral guides are available. Additionally, I found that some visual floral guides can affect the body orientation that moths adopt while hovering in front of horizontal models. Finally, when presented with flower models offering both visual and tactile guides, the former influenced proboscis placement, whereas the latter controlled proboscis movements. Results show that innate inspection behaviour is under multimodal sensory control, consistent with other components of the foraging task. Fine scale inspection movements (elicited by diverse floral traits) and the tight adjustment between the morphology of pollinators and flowers appear to be adaptively integrated, facilitating reward assessment and effective pollen transfer.