Calvarius

MEANING: Suffering

PERIOD: Late Cretaceous

CONTINENT: Europe

Calvarius is an ornithopod from the Late Cretaceous of Spain. The modified metatarsal that it is known from shows convergent evolution, resembling those of non-iguanodontian ornithopods more than closer relatives. This is likely due to fulfilling its island habitat niche, resulting in Calvarius becoming a small-bodied animal, capable of rapid locomotion.

Abstract from paper: A metatarsal IV from a hitherto unknown endemic small-bodied styracosternan is described from uppermost Maastrichtian strata of NE Spain, part of the Ibero-Armorican island of the Late Cretaceous European Archipelago. This element is unique among the fourth metatarsals of all other ornithopods in combining the greatly elongated proportions of basally branching members of the clade with the prominent medial flange seen in other styracosternans. This specimen becomes the holotype of a new genus and species. Histological data indicate that this individual was a late subadult at the time of death, but not far from reaching somatic maturity. By analogy with the elongate metatarsals of lightly built, small-bodied cursorial ornithopods, it is likely that this animal was also capable of rapid locomotion. This is consistent with the differential distribution of intense bone remodeling in metatarsal IV, likely resulting from biomechanical stress produced during rapid hindlimb propelling. Notably, this styracosternan represents an exception to the mediportality and large body size that characterize all other members of the clade. The elongation of metatarsal IV, inferred cursoriality, and small body size constitute a case of evolutionary convergence with the distantly related non-iguanodontian ornithopods and dryosaurid and elasmarian iguanodontians. Unlike other regions of the world where, during the latest stages of the Cretaceous, the small-bodied cursorial herbivore ecological niche was represented by non-iguanodontians and non-hadrosauriforms, in the Ibero-Armorican island this niche was likely occupied by a styracosternan.

Calvarius is from the Late Cretaceous. The Cretaceous is the third and final geological period of the Mesozoic Era, with the Late Cretaceous making up roughly the second half of it, lasting from about 100 to 66 million years ago. It was a time of significant evolutionary change, with dinosaurs reaching their greatest diversity before the mass extinction at the end of the Cretaceous.

The Cretaceous was a period with a relatively warm climate, though the Late Cretaceous experienced a global cooling trend, caused by falling levels of carbon dioxide in the atmosphere. The continents were nearing their present positions, but high sea levels flooded low-lying regions, turning Europe into an archipelago, and forming the Western Interior Seaway in North America. These seas were home to a variety of marine reptiles, including mosasaurs and plesiosaurs, while pterosaurs and birds shared the skies.

On land, dinosaurs continued to thrive and diversify during the Late Cretaceous, producing many of the most well-known goups, including tyrannosaurs, hadrosaurs, and pachycephalosaurs. Established Cretaceous dinosaur clades like the ceratopsians, ankylosaurs, and dromaeosaurs continued to flourish. Sauropod species consisted almost exclusively of titanosaurs, which seemed to be confined to the Southern Hemisphere for much of the Late Cretaceous. Flowering plants and grasses diversified and spread, becoming the dominant flora similar to what we see today.

The Cretaceous (along with the Mesozoic) ended with the Cretaceous-Paleogene (K-Pg) extinction event, a large mass extinction in which many groups, including non-avian dinosaurs, pterosaurs, and large marine reptiles, died out. This event, likely triggered by an asteroid impact, is marked by the abrupt K-Pg boundary, a distinct geologic layer separating the Mesozoic and Cenozoic Eras. In its aftermath, mammals and avian dinosaurs rapidly diversified, becoming the dominant land animals of the Cenozoic Era.

Calvarius is an iguanodontian. Iguanodontia was an important branch of the ornithopoda, a group of herbivorous dinosaurs that would become one of the most successful clades of the Cretaceous. These dinosaurs first appeared in the Late Jurassic as small bipedal grazers, but diversified through the Early Cretaceous, gradually increasing into larger more robust forms. While the more derived hadrosaurs of the Late Cretaceous became highly specialized for chewing tough plant material, basal iguanodontians retained a mix of primitive and advanced traits. They were among the first large-bodied ornithopods to spread widely across the Northern Hemisphere, with fossils found in Europe, North Africa, and Asia.

Early iguanodontians possessed powerful hind limbs for efficient bipedal movement but could also walk on all fours when foraging. One of their most distinctive features was their unique hand structure, which included a stiff, spike-like thumb that may have been used for defense or breaking apart tough vegetation. Although they lacked the fully developed dental batteries of later hadrosaurs, their teeth were already adapted for efficient plant processing, giving them an evolutionary advantage as herbivores. These adaptations laid the groundwork for the eventual dominance of hadrosaurs, which would expand into even more diverse habitats and continue to thrive until the end of the Cretaceous.