Geographic Mosaics of Fly Pollinators With Divergent Color Preferences Drive Landscape-Scale Structuring of Flower Color in Daisy Communities

The striking variation in flower color across and within Angiosperm species is often attributed to divergent selection resulting from geographic mosaics of pollinators with different color preferences. Despite the importance of pollinator mosaics in driving floral divergence, the distributions of pollinators and their color preferences are seldom quantified. The extensive mass-flowering displays of annual daisy species in Namaqualand, South Africa, are characterized by striking color convergence within communities, but also color turnover within species and genera across large geographic scales. We aimed to determine whether shifts between orange and white-flowered daisy communities are driven by the innate color preferences of different pollinators or by soil color, which can potentially affect the detectability of different colored flowers. Different bee-fly pollinators dominated in both community types so that largely non-overlapping pollinator distributions were strongly associated with different flower colors. Visual modeling demonstrated that orange and white-flowered species are distinguishable in fly vision, and choice experiments demonstrated strongly divergent color preferences. We found that the dominant pollinator in orange communities has a strong spontaneous preference for orange flowers, which was not altered by conditioning. Similarly, the dominant pollinator in white communities exhibited an innate preference for white flowers. Although detectability of white flowers varied across soil types, background contrast did not alter color preferences. These findings demonstrate that landscape-level flower color turnover across Namaqua daisy communities is likely shaped by a strong qualitative geographic mosaic of bee-fly pollinators with divergent color preferences. This is an unexpected result given the classically generalist pollination phenotype of daisies. However, because of the dominance of single fly pollinator species within communities, and the virtual absence of bees as pollinators, we suggest that Namaqua daisies function as pollination specialists despite their generalist phenotypes, thus facilitating differentiation of flower color by pollinator shifts across the fly pollinator mosaic.

The carcinine transporter CarT is required in Drosophila photoreceptor neurons to sustain histamine recycling

Synaptic transmission from Drosophila photoreceptors to lamina neurons requires recycling of histamine neurotransmitter. Synaptic histamine is cleared by uptake into glia and conversion into carcinine, which functions as transport metabolite. How carcinine is transported from glia to photoreceptor neurons remains unclear. In a targeted RNAi screen for genes involved in this pathway, we identified carT, which encodes a member of the SLC22A transporter family. CarT expression in photoreceptors is necessary and sufficient for fly vision and behavior. Carcinine accumulates in the lamina of carT flies. Wild-type levels are restored by photoreceptor-specific expression of CarT, and endogenous tagging suggests CarT localizes to synaptic endings. Heterologous expression of CarT in S2 cells is sufficient for carcinine uptake, demonstrating the ability of CarT to utilize carcinine as a transport substrate. Together, our results demonstrate that CarT transports the histamine metabolite carcinine into photoreceptor neurons, thus contributing an essential step to the histamine–carcinine cycle.

UV reflectance but no evidence for colour mimicry in a putative brood-deceptive orchid Corybas cheesemanii

Rewardless orchids attract pollinators by food, sexual, and brood-site mimicry, but other forms of sensory deception may also operate. Helmet orchids (Corybas, Nematoceras and related genera) are often assumed to be brood-site deceivers that mimic the colours and scents of mushrooms to fool female fungus gnats (Mycetophilidae) into attempting oviposition and pollinating flowers. We sampled spectral reflectances and volatile odours of an endemic terrestrial New Zealand orchid Corybas cheesemanii, and co-occurring wild mushrooms. The orchid is scentless to humans and SPME GC-MS analyses did not detect any odours, but more sensitive methods may be required. The orchids reflected strongly across all visible wavelengths (300-700nm) with peaks in the UV (~320nm), yellow-green (500-600 nm) and red regions (650-700 nm), whereas mushrooms and surrounding leaf Utter reflected predominantly red and no UV. Rather than mimicking mushrooms, these orchids may attract pollinators by exploiting insects’ strong sensory bias for UV. Modelhng spectral reflectances into a categorical fly vision model and a generic tetrachromat vision model provided very different results, but neither suggest any mimicry of mushrooms. However, these models require further assessment and data on fly spectral sensitivity to red wavelengths is lacking – a problem given the predominance of red, fly-pollinated flowers worldwide.