Abstract Numerous early Cenozoic transient warming events (“hyperthermals”) have been recognized in the marine record, but few have been located in fossiliferous terrestrial strata. Locating warming events is critical for understanding how terrestrial animals and plants responded to past climate change. We recognize two carbon isotope excursions (CIEs) in lower Paleocene (Danian) strata of the San Juan Basin, New Mexico, United States, based on sustained decreases in carbon isotope values in bulk organic carbon. The upper excursion correlates closely with the Latest Danian Event (LDE) hyperthermal, previously known with confidence only from the marine record. We correlate the lower excursion to a smaller CIE in the marine record tied to a 405,000-year eccentricity cycle, and modest warming. Both excursions in the San Juan Basin occur in red bed intervals bounded by drab-colored rocks, suggesting transient seasonal drying. Both are also associated with marked decreases in percent total organic carbon, possibly representing decreased primary productivity. The lower excursion is associated with a strong pulse of mammalian faunal turnover. Strata above the LDE are unfossiliferous, precluding local turnover analysis, but a strong, regional pulse of turnover occurs at or near the LDE. Our results suggest warming and/or drying during these events resulted in ecological perturbations, accelerating mammalian origination and extinction rates. Additionally, the first representatives of two North American mammalian families (Arctocyonidae, Plesiadapidae) reached Europe around the time of the LDE. We hypothesize that this was in response to short-term warming, which opened a previously impassable high-latitude dispersal route connecting the continents.

It has long been debated whether non-avian dinosaurs went extinct abruptly or gradually at the end-Cretaceous (66 million years ago), because their fossil record at this time is mostly limited to northern North America. We constrain a dinosaur-rich unit to the south, the Naashoibito Member in New Mexico, to the very latest Cretaceous (~66.4 to 66.0 million years), preserving some of the last-known non-avian dinosaurs. Ecological modeling shows that North American terrestrial vertebrates maintained high diversity and endemism in the latest Cretaceous and early Paleogene, with bioprovinces shaped by temperature and geography. This counters the notion of a low-diversity cross-continental fauna and suggests that dinosaurs were diverse and partitioned into regionally distinct assemblages during the final few hundred thousand years before the end-Cretaceous asteroid impact.

The life histories of Palaeocene mammals are poorly known, but may have been central to their success in diversifying across terrestrial ecosystems after the end‐Cretaceous extinction. Among these mammalian groups, the eutherian Taeniodonta are particularly enigmatic, with few modern analogues and no living descendants, despite being one of the only lineages to apparently traverse the Cretaceous‐Palaeogene (K‐Pg) boundary. Here, we investigate the life history of an early Palaeocene taeniodont, Conoryctes comma , based on a multi‐individual, multi‐element sample. Nearly all elements sampled exhibit similar osteohistological architecture, with a small internal zone of compacted coarse cancellous bone surrounded by an internal cortex of periosteally derived fibrolamellar bone of variable thickness, and an outer cortex of lamellar bone. The well‐vascularized fibrolamellar complex in the limb bones, lacking cyclical growth marks, is indicative of overall rapid growth to near adult body size. Cyclical growth marks are present in the outer cortex after the transition to slow‐growing lamellar bone, but not in the inner cortex, suggesting sexual maturity was reached in 1 year. In some elements, an internal non‐cyclical growth mark shares histological similarities with weaning marks in living mammals and other contemporary Palaeocene mammals, and occurred at the body size predicted for this transition in therian mammals. The unusual presence of compacted coarse cancellous bone near the midshafts of multiple limb bones may be related to cortical thickening, and is similar to the arrangement described in some fossorial mammals, supporting previous assertions of this lifestyle in Conoryctes . Altogether, these palaeohistological signals suggest a life history in C. comma similar to living eutherians, despite uncertainty about whether it is within crown Placentalia or a close outgroup. Thus, our data are consistent with an early origin of placental‐like reproductive strategies in their eutherian ancestors, although this attribute was likely shared more broadly among Mesozoic mammal lineages prior to the end‐Cretaceous extinction.

Mixodectids are poorly understood placental mammals from the Paleocene of western North America that have variably been considered close relatives of euarchontan mammals (primates, dermopterans, and scandentians) with hypothesized relationships to colugos, extinct plagiomenids, and/or microsyopid plesiadapiforms. Here we describe the most complete dentally associated skeleton yet recovered for a mixodectid, specifically Mixodectes pungens from the early Paleocene of the San Juan Basin, New Mexico. A partial skull with all the teeth erupted and associated axial skeleton, forelimbs, and hind limbs, with epiphyses fused, indicate that it was a mature adult. Results from cladistic analyses incorporating new data robustly support primatomorphan (Primates + Dermoptera) affinities of Mixodectidae, but relationships within Euarchonta are less clear, with Mixodectes recovered as a stem primatomorphan, stem dermopteran, or stem primate. Analyses of postcrania suggest that M. pungens was a relatively large (~ 1.3 kg), claw-climbing arborealist capable of frequent clinging on large diameter vertical supports. With teeth suggesting an omnivorous diet that included leaves, M. pungens occupied a unique ecological niche in the early Paleocene of North America that differed from contemporary, arboreal plesiadapiforms that were smaller and more frugivorous. Euarchontans were thus a more diverse radiation in the early Cenozoic than previously appreciated.

Ever since the discovery of Macrauchenia patachonica by Charles Darwin in 1834, the affinities of litopterns-a group of extinct South American Native Ungulates (SANUs)-have been elusive. In particular, the interfamilial relationships and timing of the familial diversification within the order Litopterna have not been addressed with adequate taxon and character sampling, and modern phylogenetic methods. We address these issues using a new morphological matrix of 703 dental and mandibular characters, scored for the earliest litopterns alongside early SANUs of other orders (Astrapotheria, Notoungulata, Pyrotheria, and Xenungulata). We also included members of families that have been often included within Litopterna, such as Didolodontidae, Protolipternidae, Indaleciidae, and Notonychopidae, the last two sometimes grouped in the order Notopterna. We conducted maximum parsimony and undated and tip-dated Bayesian phylogenetic analyses. Our results indicate that (1) didolodontids, protolipternids, and kollpaniines should be considered early SANUs, but not particularly closely related to any order, (2) Indaleciidae and Notonychopidae usually form a monophyletic group (Notopterna), and (3) Litopterna is a monophyletic unit composed of four families [Adianthidae, Anisolambdidae (subfamilies Anisolambdinae and Sparnotheriodontinae), Macraucheniidae, Proterotheriidae], and tentatively the notoptern families Indaleciidae and Notonychopidae with a probable origin similar to 64.0 Mya in the Early Palaeocene.

Following the end-Cretaceous mass extinction, mammals underwent an increase in body size, taxonomic diversity and ecological specialization throughout the Paleocene, exemplifying their adaptability. One especially enigmatic group is the "Triisodontidae", medium- to large-sized ungulate-like placentals from the Paleocene which are best known from their teeth that exhibit adaptations towards carnivory. The "triisodontids" were the first large carnivorous mammals and pre-date, and may have given rise to, Mesonychia, a group of more specialized placental carnivores. The "triisodontids" have been well-described from dental material, although very little is known about their postcrania. Here, we describe the postcrania of Triisodon crassicuspis-the most completely represented species of the genus to date-from a specimen (NMMNH P-72096) recovered from basal Torrejonian strata of the Nacimiento Formation in the San Juan Basin, New Mexico. Anatomical comparisons reveal that the forelimb long bones of Tri. crassicuspis are robust relative to its size, more so than other "triisodontids". Attachment sites on the ulna are evidence of well-developed muscles involved in powerful extension and flexion of the manus. In Tri. crassicuspis, the range of pronation-supination was limited as evident from the humeroradial morphology. Qualitative functional assessment of osteological features of the forelimb of Tri. crassicuspis is suggestive of terrestrial locomotion with at least moderate digging ability. Re-analyses of the dentition confirmed that Tri. crassicuspis had specializations for carnivory, and provide a body mass estimate of ca. 32-44 kg based on dental proxies. In summary, Tri. crassicuspis was a relatively large and powerful terrestrial animal, and one of the first known placentals to fill a largely carnivorous niche.

Conoryctes comma is a member of the enigmatic group Taeniodonta, Paleogene mammals that have been found only in North America. Taeniodonts were part of the first wave of placental mammal diversification after the end-Cretaceous extinction. The lack of postcranial elements has limited the understanding of the anatomy and locomotion of Conoryctes, and how it compared to other taeniodonts. We here describe the postcranial anatomy and functional morphology of Conoryctes, based largely on nine new specimens found in the San Juan Basin, New Mexico, USA. The specimens include elements of the axial column, such as the axis, sacrum, and ribs; the humerus, ulna, radius, and part of the manus; the innominate, femur, tibia, and part of the pes, including the tarsals. Conoryctes was a medium-sized mammal, with a robust humerus, radius, and femur, and with anatomical similarities to other conoryctid taeniodonts and Onychodectes. The tarsal elements of Conoryctes show characteristics of the “leptictimorph astragalocalcaneal morphology” as seen in other Paleogene mammals, such as Escavadodon, Palaeanodon, and Procerberus. Anatomical features of the forelimb and hindlimb of Conoryctes indicate that it was a scratch-digging animal with powerful forearm muscles and well-stabilized digits, features that may have helped it adapt to the subtropical forests of the San Juan Basin, approximately 63 million years ago. This corroborates the previous hypothesis that digging adaptations are seen in all members of Taeniodonta for which the postcranial elements are known, and that digging ability was present in the common ancestor of the clade and potentially central to their radiation after the environmental destruction of the end-Cretaceous extinction.

Caenagnathid oviraptorosaurian dinosaurs, long considered enigmas, have now become relatively easy to recognize in the fossil record of Asia and North America. This has revealed their presence in several formations spanning the Late Cretaceous, providing new insights including showing that they were more widespread in the southern parts of North America than previously recognized. Here we add to this record by describing a nearly complete right tibia from the upper Campanian De-na-zin Member of the Kirtland Formation of New Mexico, which represents the first caenagnathid material from these beds. The bone is nearly identical in morphology and size to its counterpart in the Mongolian Elmisaurus rarus, reinforcing a suite of previously-identified features that allow for straightforward recognition of caenagnathid tibiae. The growing body of caenagnathid material continues to blur boundaries between features once considered distinct, raising doubts that some taxa established from sparse material, including Ojoraptorsaurus boerei, are actually valid.

After successfully diversifying during the Paleocene, the descendants of the first wave of mammals that survived the end-Cretaceous mass extinction waned throughout the Eocene. Competition with modern crown clades and intense climate fluctuations may have been part of the factors leading to the extinction of these archaic groups. Why these taxa went extinct has rarely been studied from the perspective of the nervous system. Here, we describe the first virtual endocasts for the archaic order Tillodontia. Three species from the middle Eocene of North America were analyzed: Trogosus hillsii, Trogosus grangeri, and Trogosus castoridens. We made morphological comparisons with the plaster endocast of another tillodont, Tillodon fodiens, as well as groups potentially related to Tillodontia: Pantodonta, Arctocyonidae, and Cimolesta. Trogosus shows very little inter-specific variation with the only potential difference being related to the fusion of the optic canal and sphenorbital fissure. Many ancestral features are displayed by Trogosus, including an exposed midbrain, small neocortex, orbitotemporal canal ventral to rhinal fissure, and a broad circular fissure. Potential characteristics that could unite Tillodontia with Pantodonta, and Arctocyonidae are the posterior position of cranial nerve V3 exit in relation to the cerebrum and the low degree of development of the subarcuate fossa. The presence of large olfactory bulbs and a relatively small neocortex are consistent with a terrestrial lifestyle. A relatively small neocortex may have put Trogosus at risk when competing with artiodactyls for potentially similar resources and avoiding predation from archaic carnivorans, both of which are known to have had larger relative brain and neocortex sizes in the Eocene. These factors may have possibly exacerbated the extinction of Tillodontia, which showed highly specialized morphologies despite the increase in climate fluctuations throughout the Eocene, before disappearing during the middle Eocene.

A new, exceptionally well-preserved articulated skull roof and braincase of Melvius chauliodous, from the Kirtland Formation of New Mexico, reveals new features that provide detailed osteological information for Melvius, thus allowing for a reassessment of characters that defines the genus, its two recognized species M. chauliodous and M. thomasi, and its phylogenetic relationships within Vidalamiinae. The new features include the presence of dermopterotic rays (‘ribs’) and parietal ray bundles, previously unknown for both species of Melvius. The presence of elongated dermopterotic rays, a feature previously considered to be distinctive for the genus Calamopleurus, and tribe Calamopleurini, is now documented in Melvius, but these rays differ in their point of origination at the posteromedial corners of both dermopterotics. Bundles of short fan-shaped rays emanating from below the posterolateral corners of the parietals are also documented in both species of Melvius. Melvius was previously placed in the tribe Vidalamiini (Subfamily Vidalamiinae), but because it shares some unique features with members of the tribe Calamopleurini and the genus Calamopleurus, its placement within Vidalamiini, and the recognition of these two distinct clades (tribes) within Vidalamiinae, are not supported in our phylogenetic analysis. Previous characters distinguishing Melvius species, particularly differences in postinfraorbital ornamentation between M. chauliodous and M. thomasi, are considered dubious. However, other features may serve to determine interspecific variation or differences. For the present, we retain both species until more definitive material of M. thomasi is described.

After the end-Cretaceous extinction, placental mammals quickly diversified(1), occupied key ecological niches(2,3) and increased in size(4,5), but this last was not true of other therians(6). The uniquely extended gestation of placental young(7) may have factored into their success and size increase(8), but reproduction style in early placentals remains unknown. Here we present the earliest record of a placental life history using palaeohistology and geochemistry, in a 62 million-year-old pantodont, the clade including the first mammals to achieve truly large body sizes. We extend the application of dental trace element mapping(9,10) by 60 million years, identifying chemical markers of birth and weaning, and calibrate these to a daily record of growth in the dentition. A long gestation (approximately 7 months), rapid dental development and short suckling interval (approximately 30-75 days) show that Pantolambda bathmodon was highly precocial, unlike non-placental mammals and known Mesozoic precursors. These results demonstrate that P. bathmodon reproduced like a placental and lived at a fast pace for its body size. Assuming that P. bathmodon reflects close placental relatives, our findings suggest that the ability to produce well-developed, precocial young was established early in placental evolution, and that larger neonate sizes were a possible mechanism for rapid size increase in early placentals.

It has long been debated why groups such as non-avian dinosaurs became extinct whereas mammals and other lineages survived the Cretaceous/Paleogene mass extinction 66 million years ago. We used Markov networks, ecological niche partitioning, and Earth System models to reconstruct North American food webs and simulate ecospace occupancy before and after the extinction event. We find a shift in latest Cretaceous dinosaur faunas, as medium-sized species counterbalanced a loss of megaherbivores, but dinosaur niches were otherwise stable and static, potentially contributing to their demise. Smaller vertebrates, including mammals, followed a consistent trajectory of increasing trophic impact and relaxation of niche limits beginning in the latest Cretaceous and continuing after the mass extinction. Mammals did not simply proliferate after the extinction event; rather, their earlier ecological diversification might have helped them survive.

Mammals are the most encephalized vertebrates, with the largest brains relative to body size. Placental mammals have particularly enlarged brains, with expanded neocortices for sensory integration, the origins of which are unclear. We used computed tomography scans of newly discovered Paleocene fossils to show that contrary to the convention that mammal brains have steadily enlarged over time, early placentals initially decreased their relative brain sizes because body mass increased at a faster rate. Later in the Eocene, multiple crown lineages independently acquired highly encephalized brains through marked growth in sensory regions. We argue that the placental radiation initially emphasized increases in body size as extinction survivors filled vacant niches. Brains eventually became larger as ecosystems saturated and competition intensified.

The Late Cretaceous dinosaur Tyrannosaurus rex was recently split into three species based on the premise that variation in the T. rex hypodigm is exceptional, indicating cryptic species and “robust” and “gracile” morphs. The morphs are based on proportional ratios throughout the skeleton. The species are claimed to be stratigraphically separate, with an early robust species followed by robust and gracile descendants. There are problems with the hypothesis: the taxon diagnoses are based on two features that overlap between the species; several skulls cannot be identified based on the diagnoses; proportional comparisons between Tyrannosaurus and other theropods are based on incomparable samples; the tooth data are problematic; the stratigraphic framework divides the Hell Creek Formation into thirds, without the stratigraphic position of each specimen, or independent age control showing the subdivisions are coeval over the entire geographic area; previous work found variation in T. rex , but it cannot be parsed into discrete categories. We tested for “gracile” and “robust” morphs by analyzing the femoral and tooth ratios that were published in the multiple species study using agglomerative hierarchical clustering. The results found that each set of ratios are explained by one cluster, showing that dimorphism is not supported. We tested for exceptional variation of the femoral ratio of Tyrannosaurus ; we calculated the mean intraspecific robusticity for 112 species of living birds and 4 nonavian theropods. The results showed that the absolute variation in Tyrannosaurus is unexceptional and it does not indicate cryptic diversity. We conclude that “ T. regina ” and “ T. imperator ” are subjective junior synonyms of T. rex .

Mammals exhibit vast ecological diversity, including a panoply of locomotor behaviours. The foundations of this diversity were established in the Mesozoic, but it was only after the end-Cretaceous mass extinction that mammals began to increase in body size, diversify into many new species and establish the extant orders. Little is known about the palaeobiology of the mammals that diversified immediately after the extinction during the Palaeocene, which are often perceived as ‘archaic’ precursors to extant orders. Here, we investigate the locomotor ecology of Palaeocene mammals using multivariate and disparity analyses. We show that tarsal measurements can be used to infer locomotor mode in extant mammals, and then demonstrate that Palaeocene mammals occupy distinctive regions of tarsal morphospace relative to Cretaceous and extant therian mammals, that is distinguished by their morphological robustness. We find that many Palaeocene species exhibit tarsal morphologies most comparable with morphologies of extant ground-dwelling mammals. Disparity analyses indicate that Palaeocene mammals attained similar morphospace diversity to the extant sample. Our results show that mammals underwent a post-extinction adaptive radiation in tarsal morphology relating to locomotor behaviour by combining a basic eutherian bauplan with anatomical specializations to attain considerable ecomorphological diversity.

Tyrannosaurid theropods topped the terrestrial food chain in North America and Asia during the latest Cretaceous. Most tyrannosaurids, exemplified by Tyrannosaurus rex, had deep snouts, thick teeth, and large jaw muscles that could generate high bite forces. They coexisted in Asia with a morphologically divergent group of long-snouted relatives, called alioramins. Qianzhousaurus sinensis, from the Maastrichtian of Ganzhou, China, is the largest alioramin yet discovered, but has only been briefly described. Here we present a detailed osteological description of the holotype cranium and mandible of Qianzhousaurus. We identify several new autapomorphic features of the genus, and new synapomorphies that unite alioramins (Qianzhousaurus, Alioramus altai, Alioramus remotus) as a clade, including a laterally projecting rugosity on the jugal. We clarify that the elongate skull of alioramins involves lengthening of the anterior palate but not the premaxilla, and is reflected by lengthening of the posterior bones of the lower jaw, even though the posterior cranium (orbit and lateral temporal fenestra) are proportionally similar to deep-skulled tyrannosaurids. We show that much of the variation among the alioramin species is consistent with growth trends in other tyrannosaurids, and that A. altai, A. remotus, and Qianzhousaurus represent different ontogenetic stages of progressive maturity, across which the signature nasal rugosites of alioramins became less prominent. We predict that the holotype skull of Qianzhousaurus represents the adult level of maturity for alioramins, and propose that the skull morphology of Qianzhousaurus indicates a much weaker bite than deep-skulled tyrannosaurids, suggestive of differences in prey choice and feeding style.

We describe the tympanic anatomy of the petrosal of Deltatherium fundaminis, an enigmatic Paleocene mammal based on cranial specimens recovered from New Mexico, U.S.A. Although the ear region of Deltatherium has previously been described, there has not been a comprehensive, well-illustrated contribution using current anatomical terminology. The dental and cranial anatomy of Deltatherium is a chimera, with morphological similarities to both 'condylarth' and 'cimolestan' taxa. As such, the phylogenetic relationships of this taxon have remained elusive since its discovery, and it has variably been associated with Arctocyonidae, Pantodonta and Tillodontia. The petrosal of Deltatherium is anteriorly bordered by an open space comprising a contiguous carotid opening and pyriform fenestra. The promontorium features both a small rostral tympanic process and small epitympanic wing but lacks well-marked sulci. A large ventral facing external aperture of the canaliculus cochleae is present and bordered posteriorly by a well-developed caudal tympanic process. The hiatus Fallopii opens on the ventral surface of the petrosal. The tegmen tympani is mediolaterally broad and anteriorly expanded, and its anterior margin is perforated by a foramen for the ramus superior of the stapedial artery. The tympanohyal is small but approximates the caudal tympanic process to nearly enclose the stylomastoid notch. The mastoid is widely exposed on the basicranium and bears an enlarged mastoid process, separate from the paraoccipital process. These new observations provide novel anatomical data corroborating previous hypotheses regarding the plesiomorphic eutherian condition but also reveal subtle differences among Paleocene eutherians that have the potential to help inform the phylogeny of Deltatherium.

The lower Paleocene Ojo Alamo Sandstone and Nacimiento Formation from the San Juan Basin (SJB) in northwestern New Mexico preserve arguably the best early Paleocene mammalian record in North America and is the type location for the Puercan (Pu) and Torrejonian (To) North American Land Mammal ages (NALMA). However, the lack of precise depositional age constraints for the Ojo Alamo Sandstone and lower Nacimiento Formation has hindered our understanding of the timing and pacing of mammalian community change in the SJB following the Cretaceous-Paleogene mass extinction. Here we produced a high-resolution age model for the Ojo Alamo Sandstone and lower Nacimiento Formation combining magnetostratigraphy and 40Ar/39Ar geochronology spanning the first ~3.5 Myr of the Paleocene. Mean sediment accumulation rates during C29n were relatively low (<50 m/Myr) and equalized from basin center to basin margin indicating an accommodation minimum; sediment accumulation rates approximately double (> 90 m/Myr) during C28r and are highest in the basin center and lowest on basin margin indicating high accommodation and an increase in basin subsidence near the C29n/C28r boundary (~64.96 Ma). Puercan fossil localities were restricted to C29n, Torrejonian 1 localities to C28n, and lower Torrejonian 2 localities to C27r. Our revised age model for the SJB suggests that the first appearance of To1 mammals may have been diachronous across North America, with the Torrejonian 1 mammals first appearing in the north (Montana and North Dakota) during C29n, then in middle latitudes (Utah) in C28r, and lastly in southern North America (New Mexico) in C28n.

ABSTRACTTyrannosaurus rex and other tyrannosaurid dinosaurs were apex predators during the latest Cretaceous, which combined giant size and advanced neurosensory systems. Computed tomography (CT) data have shown that tyrannosaurids had a trademark system of a large brain, large olfactory bulbs, elongate cochlear ducts, and expansive endocranial sinuses surrounding the brain and sense organs. Older, smaller tyrannosauroid relatives of tyrannosaurids developed some, but not all, of these features, raising the hypothesis that tyrannosaurid‐style brains evolved before the enlarged tyrannosaurid‐style sinuses, which might have developed only with large body size. This has been difficult to test, however, because little is known about the brains and sinuses of the first large‐bodied tyrannosauroids, which evolved prior to Tyrannosauridae. We here present the first CT data for one of these species, Bistahieversor sealeyi from New Mexico. Bistahieversor had a nearly identical brain and sinus system as tyrannosaurids like Tyrannosaurus, including a large brain, large olfactory bulbs, reduced cerebral hemispheres, and optic lobes, a small tab‐like flocculus, long and straight cochlear ducts, and voluminous sinuses that include a supraocciptal recess, subcondyar sinus, and an anterior tympanic recess that exits the braincase via a prootic fossa. When characters are plotted onto tyrannosauroid phylogeny, there is a two‐stage sequence in which features of the tyrannosaurid‐style brain evolved first (in smaller, nontyrannosaurid species like Timurlengia), followed by features of the tyrannosaurid‐style sinuses (in the first large‐bodied nontyrannosaurid tyrannosauroids like Bistahieversor). This suggests that the signature tyrannosaurid sinus system evolved in concert with large size, whereas the brain did not. Anat Rec, 303:1043–1059, 2020. © 2020 American Association for Anatomy