How much calcium to shell out? Eggshell calcium carbonate content is greater in birds with thinner shells, larger clutches and longer lifespans

The avian eggshell is a bio-ceramic structure that protects the embryo. It is composed almost entirely of calcium carbonate and a small amount of organic material. An optimal amount of calcium carbonate in the eggshell is essential for the embryo's development, yet how the ratio of calcium carbonate to organic matter varies between species has not been investigated. Calcium is a limiting resource for most birds, so its investment in their eggs should be optimized for a bird's life history. We measured the relative calcium carbonate content of eggshells in 222 bird species and tested hypotheses for how this trait has evolved with the life-history strategies of these species and other traits of their respective egg physiologies. We found that (i) eggshell calcium carbonate content was positively correlated with species having thinner eggshells and smaller than expected eggs relative to incubating parental mass, (ii) species with small mean clutch sizes had lower calcium carbonate content in their eggshells, and (iii) for species with larger clutch sizes, eggshell calcium carbonate content was negatively correlated with their mean lifespan. The pattern of lower eggshell calcium carbonate in longer lived, larger clutched birds suggests that calcium provision to the eggshell has long-term costs for the individual.