Biliverdin-based egg coloration is enhanced by carotenoid supplementation

Avian eggshell color is remarkably variable among and within species and its possible function has long been controversial. Female birds use biliverdin pigment to color their eggs blue and green. Although evidence is accumulating that blue-green eggshell coloration is costly to produce, the exact mechanisms underlying its expression are little studied. Biliverdin pigment is involved in important maintenance functions such as immune and antioxidant defenses and shows similar properties to carotenoid pigments. Carotenoids play a role as immunoenhancers and have the potential to ameliorate the impact of oxidative processes, although their antioxidant function could be minor for some bird species. Important life-history components can be limited by carotenoid availability, including sexual displays and fecundity. Here we explored if biliverdin-based eggshell pigmentation was affected by carotenoid availability in a seabird, by performing a carotenoid supplementation experiment in female blue-footed boobies (Sula nebouxii) after laying their first egg. In this species, blue eggshell color is associated with incubation patterns, and carotenoid availability affects the crucial trade-off between female foot ornamentation and fecundity. We found that under natural conditions, there was a decline in eggshell color with laying order, suggesting pigment limitation for females. However, carotenoid-supplemented females had enhanced second egg coloration compared to controls, and the color increase was independent of their previous pigment allocation to the first egg. Our results suggest that biliverdin-based eggshell coloration is costly to produce and can be alleviated by current carotenoid availability.