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Species-level diversity and evolution of Palaeosinopa from the Willwood Formation of the Bighorn Basin is reassessed based on substantial new material from the Bighorn, Powder River, and Wind River basins. We recognize three species of Palaeosinopa in the Willwood Formation of the Bighorn Basin: P. lutreola, P. incerta, and P. veterrima. The late Wasatchian species P. didelphoides is not present in the Bighorn Basin. The Willwood species can be differentiated based only on size. P. veterrima is the most common and wide-ranging species and is the most variable in size and morphology: the stratigraphically lowest individuals are smaller, with narrower, more crestiform lower molars; whereas the highest are larger, with wider, more bunodont teeth. Although it could be argued that these represent distinct species, we demonstrate that this morphological evolution occurred as the gradual and mosaic accumulation of features, suggesting in situ anagenetic evolution. The two smaller species are present only low in the section (biochrons Wa0–Wa4) and show no discernable evolution in size or morphology. A new skeleton of Palaeosinopa veterrima from the Willwood Formation is described, and other new postcrania are reported. The skeleton is the oldest associated skeleton of Palaeosinopa known, yet it is remarkably similar to those of younger, more derived pantolestids, the primary disparities being minor differences in proportions of the innominate, femur, and tibia, and co-ossification of the distal tibia and fibula. Either P. incerta or P. lutreola was likely the ancestral population that gave rise to the other Wasatchian Palaeosinopa. Alternatively, P. veterrima may have migrated into the Bighorn Basin from the Powder River Basin.

The Pantolestidae are an extinct family of mammals known principally from the early Paleocene to late Oligocene (from approximately 64 to 30 million years ago) of North America and Europe. Although never particularly abundant, pantolestids are relatively well represented in the Eocene and Oligocene, with several taxa known from exceptionally well-preserved skulls and postcranial material. The early evolutionary history of the group, however, similar to that of many contemporaneous mammals, remains comparatively poorly known. The current study reports on several previously undescribed pantolestids from the early late Paleocene (late middle Tiffanian, Ti4) Roche Percée local fauna, Ravenscrag Formation, of southeastern Saskatchewan, Canada. Aatotomus placochton n. gen. n. sp. resembles the enigmatic pantolestid Paleotomus in having sectorial premolars with well-developed crests and tall, sharp molar trigonids, but differs principally in possessing narrow molar talonids. Besseocetor krausei n. sp. shares numerous similarities with B. thomsoni and B. septentrionalis but differs in being considerably smaller and less robust. Palaeosinopa reclusum n. sp., the oldest species of Palaeosinopa yet discovered, reveals a unique combination of primitive and derived pantolestid features, and supports previous suggestions of a close evolutionary relationship between Palaeosinopa and Bessoecetor. The new taxa document an unusually high diversity of pantolestids in the Tiffanian of western Canada and provide important new knowledge to the evolutionary history of this group during the Paleocene.

Pantolestinae is a eutherian subfamily of mammals whose members are known from the middle early Paleocene through at least the beginning of the Oligocene of North America. They are also known from Europe, and possibly Africa. A lack of information on pantolestine skulls has prevented the use of cranial anatomy in evaluation of this group’s enigmatic higher-level phylogenetic relationships. Conversely, postcranial skeletons are well known and locomotor interpretations based on them are robust. The most complete known skull of a pantolestine, Pantolestes longicaudus (YPM 13525), is described here and compared to potential close fossil relatives and extant mammals. Semicircular canal morphology is used to test locomotor hypotheses.YPM 13525 lacks an ossified bulla. It has a mediolaterally broad basioccipital, a large entoglenoid process, and a deeply incised glaserian fissure of the squamosal, caudal and rostral tympanic processes on the petrosal, a foramen for an internal carotid artery (ICA) that entered the tympanic cavity from a posteromedial position, bony tubes enclosing the main stem and transpromontorial branch of the ICA, a large anterior carotid foramen formed within the basisphenoid, evidence of a stapedial artery ramus superior, a groove on the dorsal aspect of the basisphenoid leading to the piriform fenestra possibly for drainage of the cavernous sinus to an extracranial inferior petrosal sinus, a dorsum sellae with well-developed posterior clinoid processes, a foramen rotundum within the alisphenoid, and a sphenorbital fissure between the alisphenoid and orbitosphenoid. Overall, the morphology is not strikingly similar to any potential close relative and the phylogenetic position of Pantolestinae cannot be estimated without cladistic analysis of a character matrix that includes this new morphology and broadly samples extant and extinct eutherian taxa. Semicircular canal morphology differs from that of two likely terrestrial Paleocene mammals, Aphronorus (another pantolestid) and Eoryctes (a palaeoryctid), suggesting a different, possibly semi-aquatic, lifestyle for Pantolestes.

The Paleogene Order Taeniodonta Cope, 1876—peculiar heavy-bodied mammals, some with ever-growing cheek teeth—are grouped with the Late Cretaceous eutherian CimolestesMarsh, 1889, along with a host of other taxa in a superordinal group, the Cimolesta. Taeniodonts were thought to have arisen from Cimolestes indirectly, through Paleocene ProcerberusSloan and Van Valen, 1965. The recently described Paleocene AlveugenaEberle, 1999, until now known only from the upper dentition, has been put forth as a transitional form between cimolestids and taeniodonts on phylogenetic and biostratigraphic grounds. An older taeniodont, the Late Cretaceous SchowalteriaFox and Naylor, 2003, has since been described, complicating taeniodont origins. We describe here a lower jaw that we refer to Alveugena from the lower part of the Ludlow Member of the Fort Union Formation in North Dakota. The lower jaw comes from strata of early Early Paleocene age (Puercan 1 North American Land Mammal Age) ~8.5 m above a Cretaceous-Paleogene boundary, identified using palynological criteria. A cladistic analysis is here presented using new data on Schowalteria and Alveugena, added to that of Cimolestes, Procerberus formicarumSloan and Van Valen, 1965, P. grandisMiddleton and Dewar, 2004, and Onychodectes. This analysis revealed Alveugena as the sister taxon of the taeniodonts but with a closer relationship to Cimolestes than Procerberus, suggesting that taeniodonts evolved from a Cimolestes-like ancestor. We discuss the age relations of early taeniodonts and related taxa and propose a scenario of ancestor-descendent relations that minimizes, but does not eliminate, implied stratigraphic gaps.