Titanosaurian sauropod dinosaurs were the most diverse and abundant large-bodied herbivores in the southern continents during the final 30 million years of the Mesozoic Era. Several titanosaur species are regarded as the most massive land-living animals yet discovered; nevertheless, nearly all of these giant titanosaurs are known only from very incomplete fossils, hindering a detailed understanding of their anatomy. Here we describe a new and gigantic titanosaur, Dreadnoughtus schrani, from Upper Cretaceous sediments in southern Patagonia, Argentina. Represented by approximately 70% of the postcranial skeleton, plus craniodental remains, Dreadnoughtus is the most complete giant titanosaur yet discovered, and provides new insight into the morphology and evolutionary history of these colossal animals. Furthermore, despite its estimated mass of about 59.3 metric tons, the bone histology of the Dreadnoughtus type specimen reveals that this individual was still growing at the time of death.
Substantial changes in bone histology accompany the secondary adaptation to life in the water. This transition is well documented in several lineages of mammals and non-avian reptiles, but has received relatively little attention in birds. This study presents new observations on the long bone microstructure of penguins, based on histological sections from two extant taxa (Spheniscus and Aptenodytes) and eight fossil specimens belonging to stem lineages (†Palaeospheniscus and several indeterminate Eocene taxa). High bone density in penguins results from compaction of the internal cortical tissues, and thus penguin bones are best considered osteosclerotic rather than pachyostotic. Although the oldest specimens sampled in this study represent stages of penguin evolution that occurred at least 25 million years after the loss of flight, major differences in humeral structure were observed between these Eocene stem taxa and extant taxa. This indicates that the modification of flipper bone microstructure continued long after the initial loss of flight in penguins. It is proposed that two key transitions occurred during the shift from the typical hollow avian humerus to the dense osteosclerotic humerus in penguins. First, a reduction of the medullary cavity occurred due to a decrease in the amount of perimedullary osteoclastic activity. Second, a more solid cortex was achieved by compaction. In extant penguins and †Palaeospheniscus, most of the inner cortex is formed by rapid osteogenesis, resulting an initial latticework of woven-fibered bone. Subsequently, open spaces are filled by slower, centripetal deposition of parallel-fibered bone. Eocene stem penguins formed the initial latticework, but the subsequent round of compaction was less complete, and thus open spaces remained in the adult bone. In contrast to the humerus, hindlimb bones from Eocene stem penguins had smaller medullary cavities and thus higher compactness values compared with extant taxa. Although cortical lines of arrested growth have been observed in extant penguins, none was observed in any of the current sampled specimens. Therefore, it is likely that even these 'giant' penguin taxa completed their growth cycle without a major pause in bone deposition, implying that they did not undergo a prolonged fasting interval before reaching adult size.
Endogenous biomolecules and soft tissues are known to persist in the fossil record. To date, these discoveries derive from a limited number of preservational environments, (e.g., fluvial channels and floodplains), and fossils from less common depositional environments have been largely unexplored. We conducted paleomolecular analyses of shallow marine vertebrate fossils from the Cretaceous–Paleogene Hornerstown Formation, an 80–90% glauconitic greensand from Jean and Ric Edelman Fossil Park in Mantua Township, NJ. Twelve samples were demineralized and found to yield products morphologically consistent with vertebrate osteocytes, blood vessels, and bone matrix. Specimens from these deposits that are dark in color exhibit excellent histological preservation and yielded a greater recovery of cells and soft tissues, whereas lighter-colored specimens exhibit poor histology and few to no cells/soft tissues. Additionally, a well-preserved femur of the marine crocodilian Thoracosaurus was found to have retained endogenous collagen I by immunofluorescence and enzyme-linked immunosorbent assays. Our results thus not only corroborate previous findings that soft tissue and biomolecular recovery from fossils preserved in marine environments are possible but also expand the range of depositional environments documented to preserve endogenous biomolecules, thus broadening the suite of geologic strata that may be fruitful to examine in future paleomolecular studies.
Maastrichtian–Danian sediments of the Navesink and Hornerstown formations at the Jean and Ric Edelman Fossil Park of Rowan University in Mantua Township, New Jersey, have long intrigued paleontologists. Within the basal Hornerstown Formation occurs the Main Fossiliferous Layer (MFL), a regionally well-known and diverse bonebed. The lithostratigraphic and chronostratigraphic position of this fossil layer have been debated for more than 50 years, fueling debate over its origin. Herein, we present the results of a microstratigraphic analysis of the fossil composition and distribution of the MFL undertaken to rectify these discrepancies. Through methodical top-down excavation, we recorded the three-dimensional position of every fossil encountered. Three-dimensional visualization and analyses of these data in ArcGIS Pro yielded an unprecedented view of this bonebed. Most reported discrepancies about the stratigraphic placement and thickness of the MFL can be explained by the presence of two distinct fossil assemblages within this interval that are occasionally combined into a single bonebed. The stratigraphically-lower assemblage, herein termed an “oyster layer”, is geometrically-tabular and exhibits low taxonomic diversity, high abundance of the oyster Pycnodonte, and moderate taxonomic richness. The stratigraphically-higher assemblage, the MFL, occurs approximately 9 cm higher in section and exhibits high values of taxonomic diversity, fossil abundance, and taxonomic richness. Sedimentological homogeneity throughout this interval suggests that these faunal contrasts arise from the two assemblages having formed via independent taphonomic pathways. Specifically, prevalence of Pycnodonte in the oyster layer implies formation by a selective mortality event, whereas the diversity of the MFL appears to reflect a more universal agent of mortality. Spatial variations in the stratigraphic distribution of fossils within the MFL in our excavation area indicate this assemblage does not form a simple, tabular layer as previously thought and may, in part, record original bathymetry. Importantly, our definition of the MFL and detailed characterization of its stratigraphic placement are essential for future studies on the taphonomic origin and chronostratigraphy of this bonebed. Universal use of this definition would allow researchers to confidently elucidate the exact lithostratigraphic positions of precise chronostratigraphic indicators within the MFL and accurately estimate the degree of time averaging of its fossils.
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