A new oviraptorosaur (Dinosauria: Theropoda) from the end-Maastrichtian Hell Creek Formation of North America.

Caenagnathidae is a clade of derived, Late Cretaceous oviraptorosaurian theropods from Asia and North America. Because their remains are rare and often fragmentary, caenagnathid diversity is poorly understood. Anzu wyliei is the only caenagnathid species currently described from the late Maastrichtian Hell Creek Formation of the USA and is also among the largest and most completely preserved North American caenagnathids. Smaller, less complete caenagnathid material has long been known from the Hell Creek Formation, but it is unclear whether these are juvenile representatives of Anzu or if they represent distinct, unnamed taxa. Here, we describe a relatively small caenagnathid hindlimb from the Hell Creek Formation, and conduct osteohistological analysis to assess its maturity. Histological data and morphological differences from Anzu wyliei and other caenagnathids allow us to conclude that this specimen represents a new species of caenagnathid from the Hell Creek Formation, with a smaller adult body size than Anzu. This new taxon is also distinct from other small caenagnathid material previously described from the area, potentially indicating the coexistence of three distinct caenagnathid species in the Hell Creek Formation. These results show that caenagnathid diversity in the Hell Creek ecosystem has been underestimated.

The origin of placental mammal life histories.

After the end-Cretaceous extinction, placental mammals quickly diversified1, occupied key ecological niches2,3 and increased in size4,5, but this last was not true of other therians6. The uniquely extended gestation of placental young7 may have factored into their success and size increase8, 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 mapping9,10 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.

A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs.

Pterosaurs were the first vertebrates to evolve flight1,2 and include the largest flying animals in Earth history.3,4 While some of the last-surviving species were the size of airplanes, pterosaurs were long thought to be restricted to small body sizes (wingspans ca. <1.8-1.6 m) from their Triassic origins through the Jurassic, before increasing in size when derived long-skulled and short-tailed pterodactyloids lived alongside a diversity of birds in the Cretaceous.5 We report a new spectacularly preserved three-dimensional skeleton from the Middle Jurassic of Scotland, which we assign to a new genus and species: Dearc sgiathanach gen. et sp. nov. Its wingspan is estimated at >2.5 m, and bone histology shows it was a juvenile-subadult still actively growing when it died, making it the largest known Jurassic pterosaur represented by a well-preserved skeleton. A review of fragmentary specimens from the Middle Jurassic of England demonstrates that a diversity of pterosaurs was capable of reaching larger sizes at this time but have hitherto been concealed by a poor fossil record. Phylogenetic analysis places D. sgiathanach in a clade of basal long-tailed non-monofenestratan pterosaurs, in a subclade of larger-bodied species (Angustinaripterini) with elongate skulls convergent in some aspects with pterodactyloids.6 Far from a static prologue to the Cretaceous, the Middle Jurassic was a key interval in pterosaur evolution, in which some non-pterodactyloids diversified and experimented with larger sizes, concurrent with or perhaps earlier than the origin of birds. VIDEO ABSTRACT.

A partial oviraptorosaur skeleton suggests low caenagnathid diversity in the Late Cretaceous Nemegt Formation of Mongolia.

The Nemegt Formation of the Gobi Desert of Mongolia has produced one of the most abundant and diverse oviraptorosaur records globally. However, the caenagnathid component of this fauna remains poorly known. Two caenagnathid taxa are currently recognized from the Nemegt Formation: Elmisaurus rarus and Nomingia gobiensis. Because these taxa are known from mostly non-overlapping material, there are concerns that they could represent the same animal. A partial, weathered caenagnathid skeleton discovered adjacent to the holotype quarry of Nomingia gobiensis is referable to Elmisaurus rarus, revealing more of the morphology of the cranium, mandible, pectoral girdle, and pubis. Despite metatarsals clearly exhibiting autapomorphies of Elmisaurus rarus, overlapping elements are identical to those of Nomingia gobiensis, and add to a growing body of evidence that these taxa represent a single morphotype. In the absence of any positive evidence for two caenagnathid taxa in the Nemegt Formation, Nomingia gobiensis is best regarded as a junior synonym of Elmisaurus rarus. Low caenagnathid diversity in the Nemegt Formation may reflect broader coexistence patterns with other oviraptorosaur families, particularly oviraptorids. In contrast to North America, competition with the exceptionally diverse oviraptorids may have restricted caenagnathids to marginal roles in Late Cretaceous Asian ecosystems.

A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.

Late Cretaceous trends in Asian dinosaur diversity are poorly understood, but recent discoveries have documented a radiation of oviraptorosaur theropods in China and Mongolia. However, little work has addressed the factors that facilitated this diversification. A new oviraptorid from the Late Cretaceous of Mongolia sheds light on the evolution of the forelimb, which appears to have played a role in the radiation of oviraptorosaurs. Surprisingly, the reduced arm has only two functional digits, highlighting a previously unrecognized occurrence of digit loss in theropods. Phylogenetic analysis shows that the onset of this reduction coincides with the radiation of heyuannine oviraptorids, following dispersal from southern China into the Gobi region. This suggests expansion into a new niche in the Gobi region, which relied less on the elongate, grasping forelimbs inherited by oviraptorosaurs. Variation in forelimb length and manus morphology provides another example of niche partitioning in oviraptorosaurs, which may have made possible their incredible diversity in the latest Cretaceous of Asia.

Histology of Caenagnathid (Theropoda, Oviraptorosauria) Dentaries and Implications for Development, Ontogenetic Edentulism, and Taxonomy.

The fossil record of caenagnathid oviraptorosaurs consists mainly of their fused, complexly sculptured dentaries, but little is known about the growth and development of this diagnostic structure. Previous work has suggested that the ridges and grooves on the occlusal surface are either the vestiges of teeth and their alveoli or were adaptations to increase shearing action during mastication. In addition, the distinctiveness of the dentaries has led to their use for species-level taxonomy, without a complete understanding of their variation through ontogeny. Here, we describe additional caenagnathid mandibles from the Dinosaur Park Formation of Alberta, Canada, and perform histological analyses to assess relative ontogenetic stage and the nature of the occlusal elaborations. The results show that the mandibular symphysis is synostosed early in ontogeny and does not accurately reflect ontogenetic stage in caenagnathids. In contrast, the presence of cyclical growth marks in a large specimen shows that mandibles can be used for relative histological maturity estimation. Histological features of the ridges of bone surrounding the lingual groove indicate that they are not the vestiges of tooth-bearing tissues and that caenagnathids did not lose their teeth through ontogeny as suggested in previous work. Instead, increased secondary remodeling in these structures is consistent with their use for food processing. Unexpectedly advanced maturity in a small specimen suggests that at least three caenagnathid species of varying body sizes coexisted in the Dinosaur Park Formation. These results stress the necessity of histological analysis when assessing maturity or ontogenetic trends in fossil material. Anat Rec, 303:918-934, 2020. © 2019 Wiley Periodicals, Inc.

The first oviraptorosaur (Dinosauria: Theropoda) bonebed: evidence of gregarious behaviour in a maniraptoran theropod.

A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult material was recovered from the bonebed, but small juveniles are underrepresented. The taphonomic and sedimentological evidence suggests that the Avimimus bonebed represents a perimortem gregarious assemblage. The near absence of juveniles in the bonebed may be evidence of a transient age-segregated herd or 'flock', but the behaviour responsible for this assemblage is unclear. Regardless, the Avimimus bonebed is the first evidence of gregarious behaviour in oviraptorosaurs, and highlights a potential trend of increasing gregariousness in dinosaurs towards the end of the Mesozoic.

Reacquisition of the lower temporal bar in sexually dimorphic fossil lizards provides a rare case of convergent evolution.

Temporal fenestration has long been considered a key character to understand relationships amongst reptiles. In particular, the absence of the lower temporal bar (LTB) is considered one of the defining features of squamates (lizards and snakes). In a re-assessment of the borioteiioid lizard Polyglyphanodon sternbergi (Cretaceous, North America), we detected a heretofore unrecognized ontogenetic series, sexual dimorphism (a rare instance for Mesozoic reptiles), and a complete LTB, a feature only recently recognized for another borioteiioid, Tianyusaurus zhengi (Cretaceous, China). A new phylogenetic analysis (with updates on a quarter of the scorings for P. sternbergi) indicates not only that the LTB was reacquired in squamates, but it happened independently at least twice. An analysis of the functional significance of the LTB using proxies indicates that, unlike for T. zhengi, this structure had no apparent functional advantage in P. sternbergi, and it is better explained as the result of structural constraint release. The observed canalization against a LTB in squamates was broken at some point in the evolution of borioteiioids, whereas never re-occuring in other squamate lineages. This case of convergent evolution involves a mix of both adaptationist and structuralist causes, which is unusual for both living and extinct vertebrates.

A possible instance of sexual dimorphism in the tails of two oviraptorosaur dinosaurs.

The hypothesis that oviraptorosaurs used tail-feather displays in courtship behavior previously predicted that oviraptorosaurs would be found to display sexually dimorphic caudal osteology. MPC-D 100/1002 and MPC-D 100/1127 are two specimens of the oviraptorosaur Khaan mckennai. Although similar in absolute size and in virtually all other anatomical details, the anterior haemal spines of MPC-D 100/1002 exceed those of MPC-D 100/1127 in ventral depth and develop a hitherto unreported "spearhead" shape. This dissimilarity cannot be readily explained as pathologic and is too extreme to be reasonably attributed to the amount of individual variation expected among con-specifics. Instead, this discrepancy in haemal spine morphology may be attributable to sexual dimorphism. The haemal spine form of MPC-D 100/1002 offers greater surface area for caudal muscle insertions. On this basis, MPC-D 100/1002 is regarded as most probably male, and MPC-D 100/1127 is regarded as most probably female.