New biogeochemical insights into Mesozoic terrestrial paleoecology and evidence for omnivory in troodontid dinosaurs

The Cretaceous paleocommunities of North America preserve a rich record of biodiversity that suggests many species occupied narrow biogeographic ranges in comparison to their ecological equivalents in extant systems. How taxa in these systems partitioned their niches and structured their communities can be difficult to determine from fossils alone, which has led to a variety of hypotheses concerning diets and habitat use. Here, we examine element ratios (Sr/Ca, Ba/Ca) in the enamel of a suite of co-occurring vertebrate taxa sampled from a spatiotemporally constrained interval in the Oldman Formation of Alberta, Canada, to reconstruct trophic structure, and use delta (super 13) C, delta (super 18) O, and (super 87) Sr/ (super 86) Sr compositions to test for niche partitioning and habitat use among hadrosaurids, ceratopsids, and ankylosaurs. We also test previously proposed dietary hypotheses of troodontid theropods. In large ornithischians, we find Ba/Ca and Sr/Ca ratios that are consistent with herbivory, with hadrosaurs distinct from ceratopsids and ankylosaurids in their (super 87) Sr/ (super 86) Sr ranges, a pattern that is indicative of differences in habitat use/breadth, dietary plant sources, and feeding height. The sampled mammals, varanoid lizards, dromaeosaurids, and tyrannosaurids preserve a gradient of lower Sr/Ca and Ba/Ca ratios that is consistent with animal-dominant omnivorous to faunivorous diets. Troodontids, which have been variably hypothesized as either faunivorous, omnivorous, or herbivorous due to their distinct and unusual dentition, preserve Sr/Ca and Ba/Ca ratios that fall between those of the ornithischians and the dromaeosaurids. From these multi-proxy data, we interpret troodontids as mixed-feeding to plant-dominant omnivores. These proxies represent a valuable tool for understanding the trophic and community ecology of Cretaceous ecosystems and hold enormous potential for future research in paleobiology.