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The evolution of quadrupedality from bipedal ancestors is an exceptionally rare transition in tetrapod evolution, but it has occurred several times within the herbivorous dinosaur clade Ornithischia. Stegosauria, Ankylosauria, and Ceratopsidae are all uncontroversially quadrupedal, while basal ornithischians and basal ornithopods are uncontroversially bipedal. However, stance in iguanodontian ornithopods, including the hadrosaurs, and in non-ceratopsid ceratopsians is debated because robust osteological correlates of quadrupedality have not been identified. We examine a suite of characteristics that have been previously proposed as osteological correlates for bipedality or quadrupedality in dinosaurs. These include both discrete anatomical features, which we assess as correlates for quadrupedality using character optimization onto a composite cladogram, and proportional ratios, which we assess as correlates by reconstructing nodal ancestral states using squared-change parsimony, followed by optimization. We also examine the correlation of these features with body size. An anterolateral process on the proximal ulna, hoof-shaped manual unguals, a transversely broadened ilium, a reduced fourth trochanter and a femur longer than the tibia are found to be robust correlates of quadrupedality in ornithischian dinosaurs. Along the ceratopsid “stem” lineage, quadrupedal characters were acquired in a stepwise fashion, with forelimb characters developing prior to changes in the hind limb. In contrast, iguanodontid ornithopods display a mosaic of character states, indicating varying degrees of facultative quadrupedality that probably arose for a variety of different reasons. Hadrosaurs are found to possess all character states associated with quadrupedality and were probably predominantly quadrupedal. In general, quadrupedal ornithischians do not appear to have been constrained by their bipedal ancestry to a particular order of character acquisition.
Two partial skeletons from Montana represent the northernmost occurrences of Stegosauria within North America. One of these specimens represents the northernmost dinosaur fossil ever recovered from the Morrison Formation. Consisting of fragmentary cranial and postcranial remains, these specimens are contributing to our knowledge of the record and distribution of dinosaurs within the Morrison Formation from Montana. While the stegosaurs of the Morrison Formation consist of Alcovasaurus, Hesperosaurus, and Stegosaurus, the only positively identified stegosaur from Montana thus far is Hesperosaurus. Unfortunately, neither of these new specimens exhibit diagnostic autapomorphies. Nonetheless, these specimens are important data points due to their geographic significance, and some aspects of their morphologies are striking. In one specimen, the teeth express a high degree of wear usually unobserved within this clade—potentially illuminating the progression of the chewing motion in derived stegosaurs. Other morphologies, though not histologically examined in this analysis, have the potential to be important indicators for maturational inferences. In suite with other specimens from the northern extent of the formation, these specimens contribute to the ongoing discussion that body size may be latitudinally significant for stegosaurs—an intriguing geographical hypothesis which further emphasizes that size is not an undeviating proxy for maturity in dinosaurs.
The earliest definitive ornithischian dinosaurs are from the Early Jurassic and are rare components of early dinosaur faunas. The Lower Lufeng Formation (Hettangian–Sinemurian) of Yunnan Province, China, has yielded a diverse Early Jurassic terrestrial vertebrate fauna. This includes several incomplete specimens have been referred to Ornithischia, including the type specimen of the thyreophoran “Tatisaurus” and other generically indeterminate material. The highly fragmentary Lufeng ornithischian Bienosaurus lufengensis was described briefly in 2001 and identified as an ankylosaurian dinosaur. Recent studies have cast doubt on this hypothesis, however, and given that the referral of Bienosaurus to Ankylosauria would result in an extensive ghost-lineage extending between it and the first definitive eurypodans (ankylosaurs + stegosaurs) in the Middle Jurassic, the holotype specimen is re-examined and re-described. We identify Bienosaurus as a probable thyreophoran dinosaur, although the fragmentary nature of the material and the absence of autapomorphies means that the specimen should be regarded as a nomen dubium.
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