Lynn Siefferman

EVIDENCE FOR SEXUAL SELECTION ON STRUCTURAL PLUMAGE COLORATION IN FEMALE EASTERN BLUEBIRDS (SIALIA SIALIS)

Abstract Although the function of ornamental traits in males has been the focus of intensive research for decades, expression of such traits in females has received much less study. Eastern bluebirds (Sialia sialis) display structurally based ultraviolet/blue and melanin‐based chestnut plumage, and in males this plumage coloration is related to both reproductive success and competitive ability. Compared to males, female bluebirds show a subdued expression of blue and chestnut ornamental coloration, and we used a combination of an aviary nutritional‐stress experiment and four years of field data to test the hypothesis that coloration functions as a signal of female quality. First, we tested the effect of food intake on expression of structural and melanin coloration in female eastern bluebirds to determine whether structural or melanin coloration are condition‐dependent traits. Females that were given ad libitum access to food displayed more ornamented structural coloration than females on a food‐restricted diet, but there was no effect of the experiment on melanin ornamentation. Second, we used field data to assess whether female ornamentation correlated with measures of mate quality and parental effort. The structural coloration of females predicted first egg date, maternal provisioning rates, and measures of reproductive success. These data indicate that structural coloration is dependent on nutritional condition and suggest that sexual selection is acting on structurally based plumage coloration in female eastern bluebirds.

Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour †

Evidence suggests that structural plumage colour can be an honest signal of individual quality, but the mechanisms responsible for the variation in expression of structural coloration within a species have not been identified. We used full–spectrum spectrometry and transmission electron microscopy to investigate the effect of variation in the nanostructure of the spongy layer on expression of structural ultraviolet (UV)–blue coloration in eastern bluebird (Sialia sialis) feathers. Fourier analysis revealed that feather nanostructure was highly organized but did not accurately predict variation in hue. Within the spongy layer of feather barbs, the number of circular keratin rods significantly predicted UV–violet chroma, whereas the standard error of the diameter of these rods significantly predicted spectral saturation. These observations show that the precision of nanostructural arrangement determines some colour variation in feathers.

UV-blue structural coloration and competition for nestboxes in male eastern bluebirds

Recent studies suggest that structural plumage coloration can indicate male quality and is used in female mate choice decisions. Whether or not structural coloration functions as a signal in male–male competitive interactions, however, has not been studied. Male eastern bluebirds, Sialia sialis, have brilliant ultraviolet-blue plumage on the head, back, wings and tail that is correlated with both reproductive effort and reproductive success. Bluebirds cannot excavate their own nest cavities, and as a consequence of limited nest sites, often engage in intense competition for nestboxes. We experimentally tested the hypothesis that structural coloration reflects male competitive ability by manipulating the number of available nestboxes. We erected a limited number of nestboxes in early spring and, after birds has established residency in those nestboxes, we added more nestboxes to the study site. We found that the reflective properties of the ultraviolet-blue plumage differed between males that acquired nestboxes early versus late in the spring, indicating that more colourful birds won competitions for access to nest sites. We also found that more colourful males fledged more offspring. These observations support the hypothesis that structural plumage colour is a condition-dependent trait in male eastern bluebirds that could be used to accurately assess the fighting ability of competitors.

Bacteria as an agent for change in structural plumage color: correlational and experimental evidence.

Recent studies have documented that a diverse assemblage of bacteria is present on the feathers of wild birds and that uropygial oil affects these bacteria in diverse ways. These findings suggest that birds may regulate the microbial flora on their feathers. Birds may directly inhibit the growth of harmful microbes or promote the growth of other harmless microbes that competitively exclude them. If keratinolytic (i.e., feather‐degrading) bacteria degrade colored feathers, then plumage coloration could reveal the ability of individual birds to regulate microbial flora. We used field‐ and lab‐based methods to test whether male eastern bluebirds (Sialia sialis) with brighter blue structural plumage coloration were better able to regulate their microbial flora than duller males. When we sampled bluebirds in the field, individuals with brighter color had higher bacterial loads than duller individuals. In the lab, we tested whether bacteria could directly alter feather color. We found that keratinolytic bacteria increased the brightness and purity, decreased the ultraviolet chroma, and did not affect the hue of structural color. This change in spectral properties of feathers may occur through degradation of the cortex and spongy layer of structurally colored barbs. These data suggest that bacteria can alter structural plumage color through degradation.

YOU CAN'T JUDGE A PIGMENT BY ITS COLOR: CAROTENOID AND MELANIN CONTENT OF YELLOW AND BROWN FEATHERS IN SWALLOWS, BLUEBIRDS, PENGUINS, AND DOMESTIC CHICKENS

Abstract The two main pigment types in bird feathers are the red, orange, and yellow carotenoids and the black, gray, and brown melanins. Reports conflict, however, regarding the potential for melanins to produce yellow colors or for carotenoids to produce brown plumages. We used high-performance liquid chromatography to analyze carotenoids and melanins present in the yellow and brown feathers of five avian species: Eastern Bluebirds (Sialia sialis), Barn Swallows (Hirundo rustica), King Penguins (Aptenodytes patagonicus), Macaroni Penguins (Eudyptes chrysolophus), and neonatal chickens (Gallus domesticus). In none of these species did we detect carotenoid pigments in feathers. Although carotenoids are reportedly contained in the ventral plumage of European Barn Swallows (Hirundo rustica rustica), we instead found high concentrations of both eumelanins and phaeomelanins in North American Barn Swallows (H. r. erythrogaster). We believe we have detected a new form of plumage pigment that gives penguin and domestic- chick feathers their yellow appearance. No Puedes Juzgar un Pigmento por su Color: Contenido de Carotenoide y Melanina de Plumas Amarillas y Marrones en Golondrinas, Azulejos, Pingüinos y Gallinas Domésticas Resumen. Los dos tipos principales de pigmentos que las aves incorporan en sus plumas son carotenoides, para desarrollar plumajes rojo, naranja o amarillo, y melaninas, para adquirir coloración negra, marrón, gris o tonalidades color tierra. Sin embargo, existe información conflictiva sobre la potencial coloración de plumas amarillas basadas en melanina y la presencia de caroteniodes en el plumaje marrón de ciertas especies. En este estudio, usamos cromatografía líquida de alto rendimiento para analizar los tipos y cantidades de carotenoides y melaninas presentes en las plumas amarillas y marrones de cinco especies de aves: el azulejo Sialia sialis y la golondrina Hirundo rustica, los pingüinos Aptenodytes patagonicus y Eudyptes chrysolophus y el plumón natal amarillo de la gallina doméstica Gallus domesticus. En ninguna de estas especies detectamos pigmentos carotenoides en las plumas. A pesar de que los carotenoides han sido encontrados en el plumaje ventral de la golondrina Hirundo rustica rustica, nosotros en cambio encontramos altas concentraciones de eumelaninas y feomelaninas en H. r. erythrogaster y en azulejos que variaron entre individuos y regiones de plumaje. Creemos que hemos detectado una nueva forma de pigmento de plumaje que le da a las plumas de pingüinos y pollos domésticos su apariencia amarilla.

Male eastern bluebirds trade future ornamentation for current reproductive investment

Life-history theory proposes that organisms must trade-off investment in current and future reproduction. Production of ornamental display is an important component of reproductive effort that has rarely been considered in tests of allocation trade-offs. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet-blue plumage that is correlated with mate acquisition and male competitive ability. To investigate trade-offs between current reproductive effort and the future expression of a sexually selected ornament, we manipulated the parental effort of males by changing their brood sizes. We found that parents provisioned experimentally enlarged broods more often than reduced broods. As predicted by life-history theory, the change in parental effort had a significant effect on the relative plumage ornamentation of males in the subsequent year: males with reduced broods significantly increased in plumage brightness. Moreover, this change in plumage coloration had a direct effect on the timing of breeding in the following season: males that displayed brighter plumage in the year following the manipulation mated with females that initiated egg laying earlier in the season. These data indicate that male bluebirds must trade-off conserving energy for production of future ornamentation versus expending energy for current reproduction.

The effect of rearing environment on blue structural coloration of eastern bluebirds (Sialia sialis)

We used a brood-size manipulation to test the effect of rearing environment on structural coloration of feathers grown by eastern bluebird (Sialia sialis) nestlings. Ultraviolet (UV)-blue structural coloration has been shown to be sexually selected in this species. Our experimental design took advantage of the growth of UV-blue wing feathers in nestlings that are retained as part of the first nuptial plumage. We cross-fostered nestlings to create enlarged and reduced broods with the purpose of manipulating parental feeding rates and measured the effect on nestling growth and plumage coloration. Brood size influenced feeding rates to offspring, but the effect varied with season. In general, male nestlings reared in reduced broods were fed more often, weighed more, and displayed brighter structural plumage compared to nestlings reared in enlarged broods. Female nestlings appeared to experience less adverse affects of brood enlargement, and we did not detect an effect of brood-size manipulation on the plumage coloration of female nestlings. Measures of plumage coloration in both males and females, however, were correlated to hatching date and nestling mass during feather development. These data provide empirical evidence that environmental quality can influence the development of the blue structural coloration of feathers and that males may be more sensitive to environmental fluctuations than females.

An experimental test of female choice relative to male structural coloration in eastern bluebirds

Several experimental studies have shown that female birds use ornamental melanin and carotenoid plumage coloration as criteria in mate choice. Whether females choose mates based on natural variation in structural coloration, however, has not been well established. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet (UV)-blue plumage coloration on their head, back, wings, and tail, which is positively correlated with condition, reproductive effort, and reproductive success. We experimentally tested the hypothesis that female eastern bluebirds prefer as mates males that display brighter structural coloration by presenting breeding-condition females with males of variable coloration. We conducted two types of mate-choice experiments. First, females chose between males whose coloration was manipulated within the natural range of variation in the population; feathers were either brightened with violet marker or dulled with black marker. Second, females chose between males with naturally dull or bright plumage coloration. In both manipulated and unmanipulated coloration trials, female choice did not differ significantly from random with respect to structural coloration. We found no support for the hypothesis that the UV–blue coloration of male eastern bluebirds functions as a criterion in female mate choice.

Mechanisms of evolutionary change in structural plumage coloration among bluebirds (Sialia spp.)

Combinations of microstructural and pigmentary components of barbs create the colour displays of feathers. It follows that evolutionary changes in colour displays must reflect changes in the underlying production mechanisms, but rarely have the mechanisms of feather colour evolution been studied. Among bluebirds in the genus Sialia, male rump colour varies among species from dark blue to light blue while breast colour varies from blue to rusty. We use spectrometry, transmission electron microscopy and Fourier analysis to identify the morphology responsible for these divergent colour displays. The morphology of blue rump barbs is similar among the three species, with an outer keratin cortex layer surrounding a medullary ‘spongy layer’ and a basal row of melanin granules. A spongy layer is also present in blue breast barbs of mountain bluebirds Sialia currucoides and in rusty breast barbs of western Sialia mexicana and eastern bluebirds Sialia sialis. In blue barbs melanin is basal to the spongy layer, but is not present in the outer cortex or spongy layer, while in rusty barbs, melanin is present only in the cortex. The placement of melanin in the cortex masks expression of structural blue, creating a rusty display. Such shifts in microstructures and pigments may be widespread mechanisms for the evolutionary changes in the colours of feathers and other reflective structures across colourful organisms.

Interspecific competition influences fitness benefits of assortative mating for territorial aggression in eastern bluebirds (Sialia sialis).

Territorial aggression influences fitness and, in monogamous pairs, the behavior of both individuals could impact reproductive success. Moreover, territorial aggression is particularly important in the context of interspecific competition. Tree swallows and eastern bluebirds are highly aggressive, secondary cavity-nesting birds that compete for limited nesting sites. We studied eastern bluebirds at a field site in the southern Appalachian Mountains that has been recently colonized (<40 yr) by tree swallows undergoing a natural range expansion. The field site is composed of distinct areas where bluebirds compete regularly with tree swallows and areas where there is little interaction between the two species. Once birds had settled, we measured how interspecific competition affects the relationship between assortative mating (paired individuals that behave similarly) and reproductive success in eastern bluebirds. We found a strong tendency toward assortative mating throughout the field site. In areas of high interspecific competition, pairs that behaved the most similarly and displayed either extremely aggressive or extremely non-aggressive phenotypes experienced higher reproductive success. Our data suggest that interspecific competition with tree swallows may select for bluebirds that express similar behavior to that of their mate. Furthermore, animal personality may be an important factor influencing the outcome of interactions between native and aggressive, invasive species.