Iridescence, the change in hue of a surface with varying observation angles, is used by insects, birds, fish, and reptiles for species recognition and mate selection. We identified iridescence in flowers of Hibiscus trionum and Tulipa species and demonstrated that iridescence is generated through diffraction gratings that might be widespread among flowering plants. Although iridescence might be expected to increase attractiveness, it might also compromise target identification because the object's appearance will vary depending on the viewer's perspective. We found that bumblebees (Bombus terrestris) learn to disentangle flower iridescence from color and correctly identify iridescent flowers despite their continuously changing appearance. This ability is retained in the absence of cues from polarized light or ultraviolet reflectance associated with diffraction gratings.

Insects use several senses to forage, detecting floral cues such as color, shape, pattern, and volatiles. We report a formerly unappreciated sensory modality in bumblebees (Bombus terrestris), detection of floral electric fields. These fields act as floral cues, which are affected by the visit of naturally charged bees. Like visual cues, floral electric fields exhibit variations in pattern and structure, which can be discriminated by bumblebees. We also show that such electric field information contributes to the complex array of floral cues that together improve a pollinator's memory of floral rewards. Because floral electric fields can change within seconds, this sensory modality may facilitate rapid and dynamic communication between flowers and their pollinators.

Floral colour signals are used by pollinators as predictors of nutritional rewards, such as nectar. But as insect pollinators often need to invest energy to maintain their body temperature above the ambient temperature, floral heat might also be perceived as a reward. Here we show that bumblebees (Bombus terrestris) prefer to visit warmer flowers and that they can learn to use colour to predict floral temperature before landing. In what could be a widespread floral adaptation, plants may modulate their temperature to encourage pollinators to visit.

Conical epidermal cells can influence a diverse set of petal properties. The fitness benefits they provide to plants are likely to vary with pollinator and habitat, and models are now required to understand how these different factors interact.

The plant surface is by default flat, and development away from this default is thought to have some function of evolutionary advantage. Although the functions of many plant epidermal cells have been described, the function of conical epidermal cells, a defining feature of petals in the majority of insect-pollinated flowers, has not. The location and frequency of conical cells have led to speculation that they play a role in attracting animal pollinators. Snapdragon (Antirrhinum) mutants lacking conical cells have been shown to be discriminated against by foraging bumblebees. Here we investigated the extent to which a difference in petal surface structure influences pollinator behavior through touch-based discrimination. To isolate touch-based responses, we used both biomimetic replicas of petal surfaces and isogenic Antirrhinum lines differing only in petal epidermal cell shape. We show that foraging bumblebees are able to discriminate between different surfaces via tactile cues alone. We find that bumblebees use color cues to discriminate against flowers that lack conical cells--but only when flower surfaces are presented at steep angles, making them difficult to manipulate. This facilitation of physical handling is a likely explanation for the prevalence of conical epidermal petal cells in most flowering plants.

Enhanced light-harvesting is an area of interest for optimising both natural photosynthesis and artificial solar energy capture 1,2. While iridescence has been shown to exist widely and in diverse forms in plants and other photosynthetic organisms and symbioses 3,4 , there has yet to be any direct link demonstrated between iridescence and photosynthesis. Here we show that epidermal chloroplasts, also known as iridoplasts, in shade-dwelling species of Begonia 5 , notable for their brilliant blue iridescence, have a photonic crystal structure formed from a periodic arrangement of the light-absorbing thylakoid tissue itself. This structure enhances photosynthesis in two ways: by increasing light capture at the predominantly green wavelengths available in shade conditions, and by directly enhancing quantum yield by 10-15% under low light conditions. These findings together imply that the iridoplast is a highly modified chloroplast structure adapted to make best use of the extremely low light conditions in the tropical forest understory in which it is found 5,6. A phylogenetically diverse range of shade-dwelling plant species have been found to produce similarly structured chloroplasts 7-9 , suggesting that the ability to produce chloroplasts whose

Pollinating insects utilise various sensory cues to identify and learn rewarding flower species. One such cue is floral temperature, created by captured sunlight or plant thermogenesis. Bumblebees, honeybees and stingless bees can distinguish flowers based on differences in overall temperature between flowers. We report here that floral temperature often differs between different parts of the flower creating a temperature structure or pattern. Temperature patterns are common, with 55% of 118 plant species thermographed, showing within-flower temperature differences greater than the 2°C difference that bees are known to be able to detect. Using differential conditioning techniques, we show that bumblebees can distinguish artificial flowers differing in temperature patterns comparable to those seen in real flowers. Thus, bumblebees are able to perceive the shape of these within-flower temperature patterns. Floral temperature patterns may therefore represent a new floral cue that could assist pollinators in the recognition and learning of rewarding flowers.

The area of space immediately around the floral display is likely to have an increased level of humidity relative to the environment around it, due to both nectar evaporation and floral transpiration. This increased level of floral humidity could act as a closedistance cue for pollinators or influence thermoregulation, pollen viability and infection of flowers by fungal pathogens. However, with a few exceptions, not much is known about the patterns of floral humidity in flowering plants or the physiological traits that result in its generation. We conducted a survey of 42 radially symmetrical flower species (representing 21 widely spread families) under controlled conditions. Humidity was measured using a novel robot arm technique that allowed us to take measurements along transects across and above the floral surface. The intensity of floral humidity was found to vary between different flower species. Thirty of the species we surveyed presented levels of humidity exceeding a control comparable to background humidity levels, while twelve species did not. Patterns of floral humidity also differed across species. Nevertheless, floral humidity tended to be highest near the center of the flower, and decreased logarithmically with increasing distance above the flower, normally declining to background levels within 30 mm. It remains unclear how physiological traits influence the diversity of floral humidity discovered in this survey, but floral shape seems to also influence floral humidity. These results demonstrate that floral humidity may occur in a wide range of species and that there might be greater level of diversity and complexity in this floral trait than previously known.