Avian eggshell coloration predicts shell-matrix protoporphyrin content

Avian eggshell pigmentation may provide information about a female’s physiological condition, in particular her state of oxidative balance. Previously we found that female house wrens (Troglodytes aedon Vieillot, 1809) with lighter, less-maculated, and redder ground-colored shells were older and produced heavier offspring than females laying darker, browner eggs. The strong pro-oxidant protoporphyrin is responsible for this species’ eggshell pigmentation, so differences in pigmentary coloration may be related to eggshell protoporphyrin content and reflect female oxidative balance and condition during egg-formation. Therefore, we tested the assumption that egg-surface coloration is related to the amount of protoporphyrin in the shell matrix. We analyzed digital photographs of eggs to determine maculation coverage as a measure of the overall ground coloration of the egg and its red-, green-, and blue-channel pixel values. Pigments were then extracted from these same eggs and analyzed using high-performance liquid chromatography. There was a strong, positive relationship between eggshell redness and protoporphyrin content of eggshells, but no relationship between percent maculation and protoporphyrin content. Thus, when older, larger females deposit more protoporphyrin in their eggshells, this may reflect a tolerance for high levels of circulating protoporphyrin or an effective mechanism for off-loading protoporphyrin into the eggshell matrix. Keywords: Eggs, Female quality, HPLC, Eggshell pigmentation, Eggshell speckles

Egg spottiness reflects female condition, physiological stress, and ornament expression in a common rallid species

Lay Summary • There is equivocal support for direct associations between maternal quality and deposition of protoporphyrin pigmentation (dark spots and blotches) in avian eggshells. • Research on protoporphyrin eggshell pigmentation has primarily focused on a single avian order (Passeriformes). • We examined associations of protoporphyrin-based eggshell pigmentation with female phenotypic and genetic traits in a non-passerine species, the Eurasian Coot. • Deposition of protoporphyrin in eggshells (total number per area of egg spots) positively correlated with female condition and expression of a putative bare-part ornament (frontal shield), while it was negatively associated with the level of physiological stress. • Protoporphyrin-based eggshell pigmentation acts as a reliable signal of female phenotypic (but not genetic) traits in the Eurasian Coot. • Signaling properties of protoporphyrin-based egg coloration are likely to largely differ between different evolutionary lineages of birds.

Differences in eggshell pigmentation pattern between Common Moorhen Gallinula chloropus and Eurasian Coot Fulica atra eggs

Facultative avian brood parasites increase their reproductive output by laying eggs in the nests of conspecifics or closely related species. The Common Moorhen Gallinula chloropus is a well-known facultative brood parasite that nests in wetlands, which are utilised also by a wide range of waterfowl, including the Eurasian Coot Fulica atra. The two species breed in similar habitats and have a similar egg pigmentation pattern; thus, the Coot can be a suitable host of brood-parasitic Moorhen. To study whether there is any discernible difference between the spotting pattern of Coot and Moorhen eggs, we compared the density of different sized spots on eggs of the two species. Our results show that Coot eggs have a significantly higher density of small speckles then Moorhen eggs, while the latter species has eggs with more conspicuous larger spots. Therefore, Coots can possibly rely on these differences in eggshell pattern to recognize and eject the brood parasitic Moorhen eggs.

Connecting the dots: avian eggshell pigmentation, female condition and paternal provisioning effort

Differences in avian eggshell pigmentation could be an honest signal of female quality that males use to inform their nestling provisioning effort. We investigated whether among-individual variation in protoporphyrin-based eggshell pigmentation in house wrens (Troglodytes aedon) reflects female fitness-associated traits and whether males use that information. Females laying lighter clutches were older and larger than females laying darker clutches. Nestlings hatching from lighter clutches had greater size-corrected mass on post-hatch day 11, a measure that strongly predicts survival and recruitment to the breeding population. To test whether male provisioning effort responds to clutch pigmentation, we used a reciprocal clutch cross-fostering design, swapping dark with light clutches and light with dark; in controls, we swapped light with light clutches and dark with dark. Shortly before hatching, clutches were returned to their original nest to avoid confounding effects of nestling quality on male provisioning. Contrary to the sexual selection hypothesis, clutch pigmentation had no effect on male provisioning. Males were probably able to observe eggshell pigmentation and thus had information about female quality, but they did not use this information to modulate their nestling provisioning. This may be because of constraints on species-specific reproductive opportunities, or because variation in eggshell protoporphyrin serves other functions.