Cortical growth marks reveal extended juvenile development in New Zealand moa

Cyclical growth marks in cortical bone, deposited before attainment of adult body size, reflect osteogenetic changes caused by annual rhythms and are a general phenomenon in non-avian ectothermic and endothermic tetrapods. However, the growth periods of ornithurines (the theropod group including all modern birds) are usually apomorphically shortened to less than a year, so annual growth marks are almost unknown in this group. Here we show that cortical growth marks are frequent in long bones of New Zealand's moa (Aves: Dinornithiformes), a recently extinct ratite order. Moa showed the exaggerated K-selected life-history strategy formerly common in the New Zealand avifauna, and in some instances took almost a decade to attain skeletal maturity. This indicates that reproductive maturity in moa was extremely delayed relative to all extant birds. The two presently recognized moa families (Dinornithidae and Emeidae) also showed different postnatal growth rates, which were associated with their relative differences in body size. Both species of giant Dinornis moa attained their massive stature (up to 240 kg live mass) by accelerating their juvenile growth rate compared to the smaller emeid moa species, rather than by extending the skeletal growth period.