Patterns of soft tissue and cellular preservation in relation to fossil bone tissue structure and overburden depth at the Standing Rock Hadrosaur Site, Maastrichtian Hell Creek Formation, South Dakota, USA

Recovery of soft tissues and cells from fossil bones is becoming increasingly common, with structures morphologically consistent with vertebrate osteocytes, blood vessels, fibrous/collagenous matrix, and potential intravascular contents now recognized from specimens dating back to the Permian. However, it largely remains unclear how bone tissue structure, early diagenetic regimes, and many other taphonomic variables influence or control the preservation potential of soft tissues in vertebrate fossils. To explore the influence of a few of these factors, we tested a suite of fossils from the Standing Rock Hadrosaur Site, a vast Edmontosaurus annectens bonebed in the Maastrichtian Hell Creek Formation of South Dakota, for preservation of cellular and tissue components. Demineralization of bone samples from each specimen yielded abundant microstructures morphologically consistent with vertebrate osteocytes, blood vessels, and collagenous matrix. This includes the first recovery of osteocytes and vessels from a fossil vertebral centrum and ossified tendons. Perhaps surprisingly, no correlation was found between soft tissue/cellular recovery and either bone tissue structure type (cortical vs. cancellous) or overburden depth at the time of discovery. A traditional taphonomic survey of the site, conducted in parallel and reported previously, affords a clear and detailed history of these remains, both pre- and postburial. Cumulative taphonomic evidence indicates the Edmontosaurus individuals died in a mass mortality event and their disarticulated remains were buried rapidly in a shallow floodplain pond during a crevasse splay event. Oxygenated flood waters and/or groundwater oxidized initially sideritic concretions to goethite during early diagenesis, facilitating rapid cementation of portions of the sediment that likely aided stabilization of soft tissues by shielding regions of the bones from prolonged exposure to pore fluids. Our findings support cancellous bone as a viable target for cellular analyses, corroborate previous propositions that iron-rich environments and rapid burial facilitate soft tissue preservation, and provide new details into early diagenetic environments conducive to such preservation.