Flower colour within communities shifts from overdispersed to clustered along an alpine altitudinal gradient.

Altitudinal gradients are interesting models to test the effect of biotic and abiotic drivers of floral colour diversity, since an increase in UV irradiance, decrease of pollinator availability and shifts from bee- to fly-pollination in high relative to low altitudes are expected. We tested the effect of altitude and phylogeny, using several chromatic and achromatic colour properties, UV reflectance and pollinators' discrimination capacity (Apis mellifera, Bombus terrestris, Musca domestica and Eristalis tenax), to understand the floral colour diversity in an alpine altitudinal gradient. All colour properties were weakly related to phylogeny. We found a shift from overdispersed floral colours and high chromatic contrast with the background (for bees) in the low altitude, to clustered floral colours (UV and green range for bees and flies) and clustered chromatic and achromatic properties in the high altitude. Different from flies, bees could discriminate floral colours in all altitudinal ranges. Low altitudes are likely to exhibit suitable conditions for more plant species, increasing competition for pollinators and floral colour divergence. Conversely, the increase in UV irradiance in high altitudes may filter plants with specific floral UV-reflectance patterns. Overall, floral colour diversity suggests that both biotic (pollinator fauna) and abiotic (UV irradiance) drivers shape floral communities, but their importance changes with altitude.

Insect vision models under scrutiny: what bumblebees (Bombus terrestris terrestris L.) can still tell us.

Three contending models address the ability of bees to detect and discriminate colours: the colour opponent coding (COC) model, the colour hexagon (CH) model and the receptor noise-limited (RN) model, but few studies attempt to determine which model fits experimental data best. To assess whether the models provide an accurate description of bumblebee colour space, we trained bees to discriminate four colour pairs. The perceptual distance between the colours of each pair was similar according to the CH model but varied widely according to the COC and RN models. The time that bees required to select a flower and the proportion of correct choices differed between groups: decision times decreased as achromatic contrast increased, and the proportion of correct choices increased with achromatic contrast and perceptual distance, as predicted by the COC and RN models. These results suggest that both chromatic and achromatic contrasts affected the discriminability of colour pairs. Since flower colour affects the foraging choices of bees and foraging choices affect the reproductive success of plants, a better understanding of which model is more accurate under each circumstance is required to predict bee behaviour and the ecological implications of flower choice and colour.

Out of the blue: the spectral sensitivity of hummingbird hawkmoths.

The European hummingbird hawkmoth Macroglossum stellatarum is a diurnal nectar forager like the honeybee, and we expect similarities in their sensory ecology. Using behavioural tests and electroretinograms (ERGs), we studied the spectral sensitivity of M. stellatarum. By measuring ERGs in the dark-adapted eye and after adaptation to green light, we determined that M. stellatarum has ultraviolet (UV), blue and green receptors maximally sensitive at 349, 440 and 521 nm, and confirmed that green receptors are most frequent in the retina. To determine the behavioural spectral sensitivity (action spectrum) of foraging moths, we trained animals to associate a disk illuminated with spectral light, with a food reward, and a dark disk with no reward. While the spectral positions of sensitivity maxima found in behavioural tests agree with model predictions based on the ERG data, the sensitivity to blue light was 30 times higher than expected. This is different from the honeybee but similar to earlier findings in the crepuscular hawkmoth Manduca sexta. It may indicate that the action spectrum of foraging hawkmoths does not represent their general sensory capacity. We suggest that the elevated sensitivity to blue light is related to the innate preference of hawkmoths for blue flowers.

The normalized segment classification model: A new tool to compare spectral reflectance curves.

Color patterns are complex traits under selective pressures from conspecifics, mutualists, and antagonists. To evaluate the salience of a pattern or the similarity between colors, several visual models are available. Color discrimination models estimate the perceptual difference between any two colors. Their application to a diversity of taxonomic groups has become common in the literature to answer behavioral, ecological, and evolutionary questions. To use these models, we need information about the visual system of our beholder species. However, many color patterns are simultaneously subject to selective pressures from different species, often from different taxonomic groups, with different visual systems. Furthermore, we lack information about the visual system of many species, leading ecologists to use surrogate values or theoretical estimates for model parameters.Here, we present a modification of the segment classification method proposed by Endler (Biological Journal of the Linnean Society, 1990 41, 315-352): the normalized segment classification model (NSC). We explain its logic and use, exploring how NSC differs from other visual models. We also compare its predictions with available experimental data.Even though the NSC model includes no information about the visual system of the receiver species, it performed better than traditional color discrimination models when predicting the output of some behavioral tasks. Although vision scientists define color as independent of stimulus brightness, a likely explanation for the goodness of fit of the NSC model is that its distance measure depends on brightness differences, and achromatic information can influence the decision-making process of animals when chromatic information is missing.Species-specific models may be insufficient for the study of color patterns in a community context. The NSC model offers a species-independent solution for color analyses, allowing us to calculate color differences when we ignore the intended viewer of a signal or when different species impose selective pressures on the signal.

Intrafloral Color Modularity in a Bee-Pollinated Orchid.

Flower color has been studied in different ecological levels of organization, from individuals to communities. However, it is unclear how color is structured at the intrafloral level. In bee-pollinated flowers, the unidirectional gradient in color purity and pollen mimicry are two common processes to explain intrafloral color patterns. Considering that floral traits are often integrated, usually reflecting evolutionary modules under pollinator-mediated selection, we hypothesize that such intrafloral color patterns are structured by intrafloral color modules as perceived by bee color vision system. Here, we studied the tropical bee-pollinated orchid Cattleya walkeriana, given its intrafloral color complexity and variation among individuals. Considering bee color vision, we investigated if intrafloral color modules arose among intrafloral patches (tip or base of the sepals, petals, and labellum). We expected a separate color module between the labellum patches (the main attractive structure in orchids) and petals and sepals. We measured the color reflectance and calculated the photoreceptor excitation, spectral purity, hue, and the chromatic contrast of the floral structures in the hexagon color model. Spectral purity (saturation) was higher in the labellum tip in comparison to petals and sepals, generating a unidirectional gradient. Labellum base presented a less saturated yellow UV-absorbing color, which may reflect a pollen mimicry strategy. C. walkeriana presented three intrafloral color modules corresponding to the color of petals and sepals, the color of the labellum tip, and the color of labellum base. These color modules were unrelated to the development of floral structures. Given the importance of intrafloral color patterns in bee attraction and guidance, our results suggest that intrafloral patterns could be the outcome of evolutionary color modularization under pollinator-mediated selection.

Multimodal cues provide redundant information for bumblebees when the stimulus is visually salient, but facilitate red target detection in a naturalistic background.

Our understanding of how floral visitors integrate visual and olfactory cues when seeking food, and how background complexity affects flower detection is limited. Here, we aimed to understand the use of visual and olfactory information for bumblebees (Bombus terrestris terrestris L.) when seeking flowers in a visually complex background. To explore this issue, we first evaluated the effect of flower colour (red and blue), size (8, 16 and 32 mm), scent (presence or absence) and the amount of training on the foraging strategy of bumblebees (accuracy, search time and flight behaviour), considering the visual complexity of our background, to later explore whether experienced bumblebees, previously trained in the presence of scent, can recall and make use of odour information when foraging in the presence of novel visual stimuli carrying a familiar scent. Of all the variables analysed, flower colour had the strongest effect on the foraging strategy. Bumblebees searching for blue flowers were more accurate, flew faster, followed more direct paths between flowers and needed less time to find them, than bumblebees searching for red flowers. In turn, training and the presence of odour helped bees to find inconspicuous (red) flowers. When bees foraged on red flowers, search time increased with flower size; but search time was independent of flower size when bees foraged on blue flowers. Previous experience with floral scent enhances the capacity of detection of a novel colour carrying a familiar scent, probably by elemental association influencing attention.