Michael J. Ryan’s research while affiliated with Carleton University and other places

The herbivorous dinosaur clade Ceratopsia flourished in the northern hemisphere during the Late Jurassic to Late Cretaceous. Previous palaeobiogeographic studies have suggested that their initial diversification occurred in Asia, with early‐branching neoceratopsians extending their geographical range to North America sometime during the Barremian to Albian. However, the specific timing and mode of their dispersal from Asia to North America remains unknown. Here we describe a new, early‐branching neoceratopsian, Sasayamagnomus saegusai gen. et sp. nov., from the Albian Ohyamashimo Formation in southwestern Japan, representing the easternmost fossil record of ceratopsians in Asia. Sasayamagnomus exhibits three diagnostic features in the jugal, squamosal and coracoid, respectively, and also has a unique combination of characters in the lacrimal. Our phylogenetic analysis indicates that Sasayamagnomus forms a clade with Aquilops americanus , one of the earliest neoceratopsians from North America, followed by the sister taxon Auroraceratops rugosus from China. The present time‐calibrated phylogenetic tree indicates that the immigration of neoceratopsians from Asia to North America occurred during the latest Aptian or early Albian, refining the previously suggested timeframe. This aligns with fragmentary neoceratopsian fossil records from the Lower Cretaceous of North America and the initial formation of the Bering land bridge. Furthermore, the simultaneous occurrence of global warming (which enabled the development of extensive forests in the Arctic region) and the emergence of the Bering land bridge during the Aptian–Albian, probably played a crucial role in facilitating the immigration of neoceratopsians from Asia to North America.

CMN 8917 is a small, partial skull of a duck-billed dinosaur from the upper Campanian Dinosaur Park Formation in what is now Dinosaur Provincial Park, Alberta. It represents one of the few nestling-sized juvenile hadrosaurines known to date. Support for this phylogenetic placement includes a narial vestibule not enclosed within the premaxillary dorsal and lateral processes, the presence of an anterodorsal maxillary process, and a maxillary dorsal process that is longer anteroposteriorly than dorsoventrally. The skull also possesses tooth traits traditionally associated with lambeosaurines, such as secondary ridges on some maxillary and dentary tooth crowns, and denticulation on some maxillary tooth crowns. The occurrence of these features in a juvenile hadrosaurine suggests that they were modified during ontogeny, calling into question their taxonomic utility for identifying juvenile specimens. The dentary teeth of CMN 8917 are similar to those of many adult hadrosaurids in that they possess a concave occlusal surface with steeper lingual and shallower buccal wear zones. This differs from the occlusal surface morphology present in some other juvenile hadrosaurids, which suggests interspecific differences in dental battery development—possibly reflective of dietary differences—occurred during early ontogeny in some taxa.

In this study, we use an exceptional skeleton of the pachycephalosaur Stegoceras validum (UALVP 2) to inform a comprehensive appendicular muscle reconstruction of the animal, with the goal of better understanding the functional morphology of the pachycephalosaur postcranial skeleton. We find that S. validum possessed a conservative forelimb musculature, particularly in comparison to early saurischian bipeds. By contrast, the pelvic and hind limb musculature are more derived, reflecting peculiarities of the underlying skeletal anatomy. The iliotibialis, ischiocaudalis, and caudofemoralis muscles have enlarged attachment sites and the caudofemoralis has greater leverage owing to the distal displacement of the fourth trochanter along the femur. These larger muscles, in combination with the wide pelvis and stout hind limbs, produced a stronger, more stable pelvic structure that would have proved advantageous during hypothesized intraspecific head-butting contests. The pelvis may have been further stabilized by enlarged sacroiliac ligaments, which stemmed from the unique medial iliac flange of the pachycephalosaurs. Although the pubis of UALVP 2 is not preserved, the pubes of other pachycephalosaurs are highly reduced. The puboischiofemoralis musculature was likely also reduced accordingly, and compensated for by the aforementioned improved pelvic musculature.

The early centrosaurine ceratopsid, Wendiceratops pinhornensis, was discovered in Alberta, Canada in a medium density monodominant bonebed from the Oldman Formation (mid-Campanian, ~79 Ma). The bonebed contains abundant, well-preserved, adult-sized and some juvenile-sized postcranial material, allowing for the first description of a number of elements of the postcrania of this basal centrosaurine ceratopsid. The postcranial elements described are generally consistent with postcrania described for more derived centrosaurine taxa. However the rectangular-shaped distal terminus of the ischium previously considered to be an apomorphy of Wendiceratops. is shown to also be present in Medusaceratops, and thus may be a synapomorphy of basal centrsaurines. The bonebed represents a lag deposit within a mudstone-bearing overbank facies and contains individuals from multiple age classes. It contains over 95% ceratopsid remains, with all identifiable elements referable to Wendiceratops. The elements are completely disarticulated, but have undergone little weathering or abrasion (both Stage 0), although the ends of long bones and processes capped by cartilage in life frequently exhibit evidence of wet rot and breakage by hydrological reworking after decomposition. The taphonomy of the bonebed is consist with other monodominant centrosaurine bonebeds that have been interpreted as mass death assemblages preserving evidence of gregarious (herding) behavior. At approximately 79 million years old, the Wendiceratops bonebed is approximately two million years older than other ceratopsid bonebeds indicating that this bonebed is the oldest documented evidence of herding behavior in a ceratopsid.

A monodominant Gryposaurus sp. bonebed in the lower unit of the Campanian Oldman Formation of southern Alberta is the oldest hadrosauroid bonebed documented in the province and the first described from the formation. The sedimentology of the locality and the taphonomy of the hadrosaurid material indicates that the bonebed represents an assemblage of juvenile-sized individuals that were probably transported only a short distance from where they died to where they were finally deposited and preserved in a fine-grained mudstone within an overbank sequence. Histological examination of six limb elements confirms that all individuals are juveniles, with two age classes (<1 and <2 years of age at the time of death) that likely died in the same event. Bone microstructure data indicate that Gryposaurus experienced rapid growth over the 2-year life spans documented, equivalent to other Late Cretaceous hadrosaurids in North America. The parautochthonous nature of the bonebed, and the lack of small neonate (newborn) material and almost complete lack of large adult material, suggests that the bonebed represents a segregated group of juveniles. This group of immature individuals may have been an autonomous unit that had separated itself from a larger social grouping, possibly in an effort to increase their survivability.

The pre-Campanian trionychid fossil record in North America is composed of highly fragmentary specimens, which are often not identifiable beyond Pan-Trionychidae. Here, we describe a new species of plastomenid soft-shelled turtle based on a partial shell (ROM 56647) from the Santonian Milk River Formation of southern Alberta, estimated at approximately 84 Ma. This represents the oldest relatively complete shell of a trionychid from North America. Plastomenidae, a pan-trionychid clade known only from the fossil record, is classically characterized by the complete suturing of its posterior plastral bones along the midline, a crescent-shaped entoplastron, and enlarged costals VIII. ROM 56647 has a unique combination of plastomenid characters (i.e., mid-line contact of hypoplastra and xiphiplastra, anteroposteriorly long eighth costal) and apomorphies (an emarginate nuchal, enlarged tubercles on the carapace, a wide pygal notch with a straight anterior edge, and a fused hyo-hypoplastron) that allows us to identify it as a new taxon, Jimemys glaebosus gen. et sp. nov. Phylogenetic analysis places J. glaebosus within Plastomenidae as the sister taxon to a clade containing Plastomenus, Helopanoplia, and Hutchemys. This phylogenetic position implies that Aspideretoides foveatus Leidy, 1856, Atoposemys, and Gilmoremys, all of which are more basal within Plastomenidae, had ghost lineages extending at least to the Santonian. As the oldest pan-trionychid that is diagnostic to the species level in North America, J. glaebosus provides new insights into both the early evolution of trionychids in North America and the biodiversity of southern Alberta during a poorly sampled time in the Late Cretaceous of North America.

Goniopholididae is a group of basal neosuchian crocodyliforms closely related to Paralligatoridae and Eusuchia that lived during the Jurassic and Early Cretaceous. Goniopholidids have the long, flat snout and secondary palate of modern crocodylians, the acquisition of which is regarded as a key feature in the early evolution of crocodylian body plan and their aquatic adaptation. Here, we report a new species, Amphicotylus milesi , with the description from the best-preserved specimen to date of Goniopholididae from Wyoming, USA. Its posterior extension of the nasopharyngeal passage (pterygoid secondary palate) and the shortening and dorsal deflection of the ceratobranchial suggest that basal neosuchians could raise their gular valve to separate oral and pharyngeal cavities as in modern crocodylians. The anatomy of Amphicotylus milesi sheds light on the acquisition of this new respiratory system in the crocodyliform evolution and their early aquatic adaptation, leading to modern crocodylians.

Hadrosaurids are a diverse and widely distributed group of ornithischian dinosaurs that are particularly well represented in the upper Campanian Dinosaur Park Formation of the Belly River Group of Alberta. However, the origin of this hadrosaurid diversity in Alberta is poorly understood, as the lower Campanian terrestrial deposits of the underlying Oldman and Foremost formations of the group have produced comparatively few body fossils. Here we provide the first description of a partially articulated hadrosaurid and hadrosaurid material from a bonebed from the Foremost Formation and refer it to the brachylophosaurin Probrachylophosaurus sp. indet. The material represents the oldest occurrence of Brachylophosaurini in Alberta and the oldest known hadrosaurid diagnostic to the genus level from Canada. In Alberta, Hadrosaurinae display a distinct pattern of replacement with the tribes Brachylophosaurini and Kritosaurini being confined to the lower to middle Campanian strata (below the marine Bearpaw Formation) and replaced above the Bearpaw Formation by members of Saurolophini (Prosaurolophus, Saurolophus) and Edmontosaurini (Edmontosaurus), with the latter clade persisting to the end of the Maastrichtian. Although the worldwide stratigraphic distribution of the Hadrosaurinae is complex, this pattern generally holds true for northern Laramidian hadrosaurine tribes, suggesting that their pattern of evolution and replacement may be driven by some common underlying factor such as an environmental response to fluctuations in the margins of the Western Interior Seaway due to sea level change.

A new articulated postcranial specimen of an indeterminate ankylosaurid dinosaur from the Upper Cretaceous (middle-upper Campanian) Baruungoyot Formation from Hermiin Tsav, southern Gobi Desert, Mongolia includes twelve dorsal vertebrae, ribs, pectoral girdles, forelimbs, pelvic girdles, hind limbs, and free osteoderms. The new specimen shows that Asian ankylosaurids evolved rigid bodies with a decreased number of pedal phalanges. It also implies that there were at least two forms of flank armor within Ankylosauridae, one with spine-like osteoderms and the other with keeled rhomboidal osteoderms. Unique anatomical features related to digging are present in Ankylosauridae, such as dorsoventrally flattened and fusiform body shapes, extensively fused series of vertebrae, anteroposteriorly broadened dorsal ribs, a robust humerus with a well-developed deltopectoral crest, a short robust ulna with a well-developed olecranon process, a trowel-like manus, and decreased numbers of pedal phalanges. Although not fossorial, ankylosaurids were likely able to dig the substrate, taking advantage of it for self-defence and survival.