Embryos and eggs for the Cretaceous theropod dinosaur Troodon formosus

Abstract

ABSTRACT Elongate and asymmetric eggs of the oospecies Prismatoolithus levis occur regularly in the Upper Cretaceous Two Medicine Formation of western Montana. These eggs had previously been assigned to the ornithischian Orodromeus makelai, for both juvenile and adult remains are typically associated with these eggs. Reexamination of the embryos shows them to exhibit at least 24 apomorphies of the clades Dinosauria, Theropoda and Paraves. The embryos also display a pneumatic quadrate, closely placed basal tubera, a high tooth count, a metatarsal II much narrower than IV and a strongly constricted metatarsal III, all possible synapomorphies of the Troodontidae. Presence of large basal tubera and a broadly rounded anterior border of the maxillary fenestra permit assignment to Troodon formosus. Most but not all bones appear ossified, suggesting a developmental level comparable to stages 35–38 of avian embryos and a time approaching hatching. Embryos show a consistent level of development from one egg to another indicating synchronous hatching of the clutch. Embryonic Troodon exhibit long distal segments and radically different hindlimb proportions in comparison to adults. Orodromeus and other small vertebrate remains associated with Troodon egg horizons may represent prey of the adults during egg-laying and brooding. Troodon eggs show several aspects either shared or convergent with some birds, and further demonstrate the close relationship of Troodontidae and Aves. These features include: asymmetric egg form, non-branching angusticanaliculate pores, distinct structural differentiation of the mammillary and overlying prismatic layer, barrel-shaped mammillary cones with a blocky calcite cleavage, and prismatic structure visible throughout the second structural layer.