Incubation behaviours of oviraptorosaur dinosaurs in relation to body size.

Most birds sit on their eggs during incubation, a behaviour that likely evolved among non-avian dinosaurs. Several 'brooding' specimens of smaller species of oviraptorosaurs and troodontids reveal these non-avian theropods sat on their eggs, although little is known of incubation behaviour in larger theropod species. Here we examine egg clutches over a large body size range of oviraptorosaurs in order to understand the potential effect of body size on incubation behaviour. Eggshell porosity indicates that the eggs of all oviraptorosaurs were exposed in the nest, similar to brooding birds. Although all oviraptorosaur clutches consist of radially arranged eggs in a ring configuration, clutch morphology varies in that the central opening is small or absent in the smallest species, becomes significantly larger in larger species, and occupies most of the nest area in giant species. Our results suggest that the smallest oviraptorosaurs probably sat directly on the eggs, whereas with increasing body size more weight was likely carried by the central opening, reducing or eliminating the load on the eggs and still potentially allowing for some contact during incubation in giant species. This adaptation, not seen in birds, appears to remove the body size constraints of incubation behaviour in giant oviraptorosaurs.