Abstract. Matches between the bills of hummingbirds and the flowers they visit have been interpreted as examples of coadaptation and feeding specialization. Observations of birds feeding at flowers longer or shorter than their bills combined with a lack of experimental evidence for foraging trade-offs, however, fail to support these interpretations. We addressed these inconsistencies by considering a seldom-studied dimension of hummingbird-flower relationships, the shape of bills and flowers, through experiments on the Purple-throated Carib, Eulampis jugularis, and its major food plant, Heliconia, in the eastern Caribbean. Bills of male E. jugularis are considerably shorter and straighter than bills of females. We examined foraging performances and trade-offs during visits to natural heliconias and 34 artificial flowers of differing length and curvature. Supporting predictions based on matches between bill and flower morphology, handling times of females were significantly shorter than those of males at the long, curved flowers of a green morph of H. bihai. Contrary to predictions, handling times of males were not significantly shorter than handling times of females at the short flowers of H. caribaea. At artificial flowers, maximum extraction depths of females were significantly longer than maximum extraction depths of males at all curved flowers, but not at straight flowers. Handling times of females were significantly shorter than handling times of males at the longest artificial flowers for all curvatures, whereas handling times of males were significantly shorter at short, straight, artificial flowers, but only while hover-feeding without a perch. Within each sex, handling times were inversely related to bill length at long flowers for all shapes. Taken together, these performance trade-offs suggest that the long, curved bills of females are adapted for feeding from long, curved flowers, whereas the short bills of males are adapted for hover-feeding from short, straighter flowers. In addition, the finding that differences in feeding performance occur at the extremes of floral phenotypes suggests that the evolution of bill morphology may be driven by a small subset of the flowers visited by a hummingbird species.

Genes underlying signal reception should evolve to maximize signal detection in a particular environment. In animals, opsins, the protein component of visual pigments, are predicted to evolve according to this expectation. Fireflies are known for their bioluminescent mating signals. The eyes of nocturnal species are expected to maximize detection of conspecific signal colors emitted in the typical low-light environment. This is not expected for species that have transitioned to diurnal activity in bright daytime environments. Here we test the hypothesis that opsin gene sequence plays a role in modifying firefly eye spectral sensitivity. We use genome and transcriptome sequencing in four firefly species, transcriptome sequencing in six additional species, and targeted gene sequencing in 28 other species to identify all opsin genes present in North American fireflies and to elucidate amino acid sites under positive selection. We also determine whether amino acid substitutions in opsins are linked to evolutionary changes in signal mode, signal color, and light environment. We find only two opsins, one long wavelength and one ultraviolet, in all firefly species and identify 25 candidate sites that may be involved in determining spectral sensitivity. In addition, we find elevated rates of evolution at transitions to diurnal activity, and changes in selective constraint on LW opsin associated with changes in light environment. Our results suggest that changes in eye spectral sensitivity are at least partially due to opsin sequence. Fireflies continue to be a promising system in which to investigate the evolution of signals, receptors, and signaling environments.

Firefly species (Lampyridae) vary in the color of their adult bioluminescence. It has been hypothesized that color is selected to enhance detection by conspecifics. One mechanism to improve visibility of the signal is to increase contrast against ambient light. High contrast implies that fireflies active early in the evening will emit yellower luminescence to contrast against ambient light reflected from green vegetation, especially in habitats with high vegetation cover. Another mechanism to improve visibility is to use reflection off the background to enhance the light signal. Reflectance predicts that sedentary females will produce greener light to maximize reflection off the green vegetation on which they signal. To test these predictions, we recorded over 7500 light emission spectra and determined peak emission wavelength for 675 males, representing 24 species, at 57 field sites across the Eastern United States. We found support for both hypotheses: males active early in more vegetated habitats produced yellower flashes in comparison to later-active males with greener flashes. Further, in 2 of the 8 species with female data, female light emissions were significantly greener as compared to males.

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Monarch butterflies (Danaus plexippus) throughout the world are commonly infected by the specialist pathogen Ophryocystis elektroscirrha (OE). This protozoan is transmitted when larvae ingest infectious stages (spores) scattered onto host plant leaves by infected adults. Parasites replicate internally during larval and pupal stages, and adult monarchs emerge covered with millions of dormant spores on the outsides of their bodies. Across multiple monarch populations, OE varies in prevalence and virulence. Here, we examined geographic and genetic variation in OE spore morphology using clonal parasite lineages derived from each of four host populations (eastern and western North America, South Florida and Hawaii). Spores were harvested from experimentally inoculated, captive-reared adult monarchs. Using light microscopy and digital image analysis, we measured the size, shape and color of 30 replicate spores per host. Analyses examined predictors of spore morphology, including parasite source population and clone, parasite load, and the following host traits: family line, sex, wing area, and wing color (orange and black pigmentation). Results showed significant differences in spore size and shape among parasite clones, suggesting genetic determinants of morphological variation. Spore size also increased with monarch wing size, and monarchs with larger and darker orange wings tended to have darker colored spores, consistent with the idea that parasite development depends on variation in host quality and resources. We found no evidence for effects of source population on variation in spore morphology. Collectively, these results provide support for heritable variation in spore morphology and a role for host traits in affecting parasite development.