Ankylosaurian body armor function and evolution with insights from osteohistology and morphometrics of new specimens from the Late Cretaceous of Antarctica

The body armor of ankylosaurians is a unique morphological feature among dinosaurs. While ankylosaurian body armor has been studied for decades, paleohistological analyses have only started to uncover the details of its function. Yet there has been an overall bias toward sampling ankylosaurian remains from the Northern Hemisphere, with limited quantitative studies on the morphological and functional evolution of the osteoderms composing their body armor. Here, we describe new ankylosaurian materials recovered from the Late Cretaceous of Antarctica that, in combination with data compiled from the literature, reveal new insights into the evolution of the ankylosaurian body armor. Based on histological microstructure and phylogenetic results, the new Antarctic material can be assigned to Nodosauridae. This group shares the absence/poor development of their osteodermal basal cortex and highly ordered sets of orthogonal structural fibers in the superficial cortex. Our morphospace analyses indicate that large morphological diversity is observed among both nodosaurids and ankylosaurids, but osteoderms became more functionally specialized in late-diverging nodosaurids. Besides acting as effective protection against predation, osteoderms also exhibit highly ordered structural fibers in nodosaurids, enabling a decrease in cortical bone thickness (as in titanosaurs), which could have been co-opted for secondary functions, such as calcium remobilization for physiological balance. The latter may have played a key role in nodosaurid colonization of high-latitude environments, such as Antarctica and the Arctic Circle.