Theropod Research Articles

New evidence for the presence of carcharodontosaurid theropod dinosaurs in the Late Cretaceous of Uzbekistan

ABSTRACT An incomplete posterior dorsal vertebra ZIN PH 2646/16 from the Upper Cretaceous (Turonian) Bissekty Formation of Uzbekistan has a highly pneumatized centrum and neural arch. The centrum has a lateral pneumatic foramen that is divided by a septum. The neural arch is invaded through a lateral pneumatic foramen above the neurocentral suture and has deeply excavated prezygapophyseal-centrodiapophyseal and centrodiapophyseal fossae. The interior of the preserved portion of the neural spine is pneumatic. In the structure of the fossae on the neural arch and the presence of a double pneumatic foramen on the centrum, ZIN PH 2646/16 closely resembles the carcharodontosaurid Acrocanthosaurus from the Early Cretaceous of North America. Thus, we identify ZIN PH 2646/16 as Carcharodontosauridae indet. Carcharodontosauridae were previously known from the Bissekty Formation from Ulughbegsaurus uzbekistanensis, which is based on an incomplete maxilla.

New theropod dinosaur remains from the Upper Cretaceous of the Kem Kem Group (Eastern Morocco) clarify spinosaurid morphology

Abstract The Kem Kem Group is a lowermost lithostratigraphic unit from the Upper Cretaceous that extends along the border between Algeria and Morocco, in the northern region of Africa. This geological unit has yielded several tetrapod fossils, including a well-represented assemblage of theropod dinosaurs, after more than eight decades of research. Here, we report new occurrences of spinosaurid theropods from the spinosaurine clade in the Kem Kem Group by providing anatomical descriptions and taxonomic identifications of 11 new specimens derived from the Tafilalt region of Morocco. Among the findings, we describe a cervical vertebra of Sigilmassasaurus, in addition to several cranial, axial, and appendicular elements that can safely be attributed to Spinosaurinae. Moreover, based on a unique combination of characteristics, we also describe an isolated and partial ischium belonging to an indeterminate carcharodontosaurid. We also deliver a detailed redescription of one of the most complete snouts of a spinosaurine known to date. Therefore, the theropod dinosaurs of the Kem Kem Group show considerable diversity, but many questions, especially related to the diversity of spinosaurids and the general abundance of carnivorous dinosaurs in this region, remain unclear until new materials are discovered and complete descriptions are made.

Endocranial development in non-avian dinosaurs reveals an ontogenetic brain trajectory distinct from extant archosaurs

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form.

Isolated theropod teeth from the Upper Cretaceous of Goias State (Brazil): Northernmost occurrence of Abelisauridae from the Bauru Basin

Dinosaur fossils are becoming increasingly well-known from the Bauru Basin of Brazil, and represent some of the last occurring dinosaurs in South America before the end-Cretaceous mass extinction. However, sampling across the basin is not uniform, and comparatively little is known about the fossils from the northern part of the basin. Our fieldwork team discovered two isolated and well-preserved ziphodont theropod teeth from northern basin strata of the Maastrichtian Marília Formation of the south of the state of Goiás. We here identify them using quantitative analysis of morphometric data and comparisons to teeth previously described in the literature. Our comparisons and analyses indicate that they can be assigned to Abelisauridae. These specimens have little evident crown enamel ornamentation, a characteristic widely present in the teeth of theropods from the Southern Hemisphere. The materials described here are the northernmost Bauru Basin abelisaurids reported so far, demonstrating that these carnivorous dinosaurs were important faunal components across the extent of the basin, as they were in other regions of South America.

A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan

Abstract Recent fieldwork in the late Middle Jurassic Balabansai Formation of Kyrgyzstan has yielded a partial skeleton of a large theropod dinosaur. The material includes a few bones of the skull (postorbital, quadratojugal), dorsal and sacral vertebrae, fragments of the pectoral girdle and forelimbs, and an almost complete pelvic girdle and hindlimbs, and is here made the type of a new theropod taxon, Alpkarakush kyrgyzicus gen. et sp. nov. Alpkarakush can be diagnosed by an extremely developed orbital brow on the postorbital, a pneumatic opening leading into cavities in the neural arch from the centroprezygodiapophyseal fossa in the posterior dorsal vertebrae, an almost enclosed ventral sulcus in manual phalanx II-1, a narrow and deep intercondylar groove on the anterior side of the distal femur, and an epicondylar crest on the distal femur that is offset from the distal end. A second, fragmentary, and smaller specimen from the same site represents the same taxon. Based on long bone histology, the type of Alpkarakush represents a late subadult individual, whereas the smaller specimen is a juvenile, possibly indicating gregarious behaviour. Phylogenetic analysis places Alpkarakush in the Metriacanthosauridae, underlining the diversity and wide distribution of this clade in the Jurassic of Asia.

Iron-coated Komodo dragon teeth and the complex dental enamel of carnivorous reptiles

Komodo dragons (Varanus komodoensis) are the largest extant predatory lizards and their ziphodont (serrated, curved and blade-shaped) teeth make them valuable analogues for studying tooth structure, function and comparing with extinct ziphodont taxa, such as theropod dinosaurs. Like other ziphodont reptiles, V. komodoensis teeth possess only a thin coating of enamel that is nevertheless able to cope with the demands of their puncture–pull feeding. Using advanced chemical and structural imaging, we reveal that V. komodoensis teeth possess a unique adaptation for maintaining their cutting edges: orange, iron-enriched coatings on their tooth serrations and tips. Comparisons with other extant varanids and crocodylians revealed that iron sequestration is probably widespread in reptile enamels but it is most striking in V. komodoensis and closely related ziphodont species, suggesting a crucial role in supporting serrated teeth. Unfortunately, fossilization confounds our ability to consistently detect similar iron coatings in fossil teeth, including those of ziphodont dinosaurs. However, unlike V. komodoensis, some theropods possessed specialized enamel along their tooth serrations, resembling the wavy enamel found in herbivorous hadrosaurid dinosaurs. These discoveries illustrate unexpected and disparate specializations for maintaining ziphodont teeth in predatory reptiles.

The evolution of the air sac system in theropod dinosaurs: Evidence from the Upper Cretaceous of Madagascar.

Recent evidence suggests that the invasive air sac system evolved at least three times independently in avemetatarsalians: in pterosaurs, sauropodomorphs and theropods. Data from sauropodomorphs showed that the pneumatic architecture in vertebrae first developed in camellate-like trabeculae in the Triassic, later in camerate systems in Jurassic neosauropods, and finally camellate tissue in Cretaceous titanosaurs. This evolutionary trajectory has support from a considerable sampling of sauropodomorph taxa. However, the evolution of pneumatic bone tissues in Theropoda is less understood. We analyzed the computed tomography of Majungasaurus and Rahonavis, using densitometry rendering to differentiate the microarchitecture along the presacral axial skeleton of late Ceratosaurians and early Paravians. We also compared these results with scans of other theropod clades. Our analysis revealed an increase in pneumatic complexity in early paravians compared to the ceratosaurians. Majungasaurus presents some apneumatic neural spines, a condition also observed in Allosaurus. Majungasaurus also features some apneumatic centra despite the presence of lateral pneumatic fossae. This raises caution when evaluating PSP solely based on external morphology. We also found evidence of distinct patterns of PSP in maniraptorans. Considering that Majungasaurus, a late abelisaurid, inherited from their ceratosaurian ancestors, some apneumatic elements such as the neural spine and some centra, Rahonavis, an early paravian, took a different trajectory toward the full pneumatization of the axial skeleton. This characteristic provided paravians an advantage in gliding and flying. Also, unlike Sauropoda, pneumaticity in Theropoda apparently developed by increasing chamber volumes toward paravians. Similar studies on early Theropoda are needed to elucidate their condition and better describe the evolutionary trajectory of different groups.

Late Jurassic–Early Cretaceous dinosaur track assemblages from northwestern Hebei Province, China: Implications for paleoenvironment and paleoecology

The relationship between the Middle–Late Jurassic Yanliao Biota and the Early Cretaceous Jehol Biota has long been unresolved due to an approximately 20 Ma “vertebrate fossil gap”. However, a large number of dinosaur tracks have been reported from the Tuchengzi/Houcheng Formation in northern Hebei–western Liaoning, which occupies the stratigraphic position between the Yanliao Biota and the Jehol Biota. This paper presents new discoveries of diverse dinosaur track assemblages from the Upper Jurassic–Lower Cretaceous Houcheng Formation in the Shangyi Basin, northwestern Hebei Province, China. Based on the ichnological analyses, tracks are assigned to the sauropod Parabrontopodus, theropod Grallator, Anchisauripus, Eubrontes, and Therangospodus. The theropod tracks were likely produced by small-sized feathered theropod dinosaurs (Coelurosauria) that thrived in both the Yanliao Biota and the Jehol Biota. These findings imply that these two biotas evolved successively, without evidence of a complete turnover or extinction of biotas. Sedimentological studies of the tracksites reveal their occurrence in diverse sedimentary environments, including braided sandbar, floodplain, and deltaic plain. The coexistence of highly diverse dinosaur tracks with various preservation environments indicates that the living environment during that time was relatively comfortable. The increased diversity of dinosaur tracks in the upper part of the Houcheng Formation closely aligns with the intense volcanic activity of the Zhangjiakou Formation in northwestern Hebei Province. This suggests that volcanic activity likely served a crucial function in the proliferation of the dinosaur fauna in the Shangyi Basin and northern North China during the earliest Cretaceous.

The postcranial anatomy and osteohistology of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha) from the Late Triassic of Wales

AbstractThe earliest crocodylomorphs, known as non‐crocodyliform crocodylomorphs, first appeared during the Late Triassic. In contrast to extant crocodylians, which are all semi‐aquatic, early crocodylomorphs represent terrestrial taxa with a fully erect posture and in most cases a small body size. Their gracile skeletons suggest an active mode of life, possibly similar to contemporaneous, bipedal theropod dinosaurs. Despite this remarkable body plan, the postcranial morphology of early crocodylomorphs has rarely been documented in detail, restricting our ability to infer aspects of their functional morphology and evolution. Here, we provide a detailed description of the postcranium of Terrestrisuchus gracilis, a small‐bodied crocodylomorph from the Late Triassic of Pant‐y‐Ffynnon Quarry (southern Wales, UK), including a description of long bone tissues based on histological thin sections. Almost all elements of the postcranial skeleton have been preserved. The skeleton of Terrestrisuchus gracilis is highly gracile, even for a non‐crocodyliform crocodylomorph. Osteological correlates of the appendicular skeleton suggest that Terrestrisuchus gracilis had a digitigrade, quadrupedal posture. A quantitative analysis of limb robustness corroborates that Terrestrisuchus gracilis was a quadruped. Histological analysis suggests that all sampled specimens were skeletally immature and had fast growth at the time of death, as indicated by the lack of an external fundamental system and the predominance of fibrolamellar bone. The bone tissue is similar to that recently described for Saltoposuchus connectens and certain non‐crocodylomorph pseudosuchians, but differs from Hesperosuchus agilis and crocodyliforms, in which parallel‐fibred bone is more prevalent.

Diuqin lechiguanae gen. et sp. nov., a new unenlagiine (Theropoda: Paraves) from the Bajo de la Carpa Formation (Neuquén Group, Upper Cretaceous) of Neuquén Province, Patagonia, Argentina

BackgroundUnenlagiine paravians are among the most relevant Gondwanan theropod dinosaur clades for understanding the origin of birds, yet their fossil record remains incomplete, with most taxa being represented by fragmentary material and/or separated by lengthy temporal gaps, frustrating attempts to characterize unenlagiine evolution.Results and conclusionsHere we describe Diuqin lechiguanae gen. et sp. nov., a new unenlagiine taxon from the Upper Cretaceous (Santonian) Bajo de la Carpa Formation of the Neuquén Basin of Neuquén Province in northern Patagonia, Argentina that fills a substantial stratigraphic gap in the fossil record of these theropods. Although known only from a very incomplete postcranial skeleton, the preserved bones of Diuqin differ from corresponding elements in other unenlagiines, justifying the erection of the new taxon. Moreover, in several morphological aspects, the humerus of Diuqin appears intermediate between those of geologically older unenlagiines from the Neuquén Basin (e.g., Unenlagia spp. from the Turonian–Coniacian Portezuelo Formation) and that of the stratigraphically younger, larger-bodied Austroraptor cabazai from the Campanian–Maastrichtian Allen Formation. Consequently, the morphology of the new taxon appears to indicate a transitional stage in unenlagiine evolution. Phylogenetic analysis recovers Diuqin as a paravian with multiple plausible systematic positions, but the strongest affinity is with Unenlagiinae. The humerus of the new form exhibits subcircular punctures near its distal end that are interpreted as feeding traces most likely left by a conical-toothed crocodyliform, mammal, or theropod, the latter potentially corresponding to a megaraptorid or another unenlagiine individual. Thus, in addition to filling important morphological and temporal gaps in unenlagiine evolutionary history, the new taxon also offers information relating to the paleoecology of these theropods.

A new abelisaurid dinosaur from the end Cretaceous of Patagonia and evolutionary rates among the Ceratosauria.

Gondwanan dinosaur faunae during the 20 Myr preceding the Cretaceous-Palaeogene (K/Pg) extinction included several lineages that were absent or poorly represented in Laurasian landmasses. Among these, the South American fossil record contains diverse abelisaurids, arguably the most successful groups of carnivorous dinosaurs from Gondwana in the Cretaceous, reaching their highest diversity towards the end of this period. Here we describe Koleken inakayali gen. et sp. n., a new abelisaurid from the La Colonia Formation (Maastrichtian, Upper Cretaceous) of Patagonia. Koleken inakayali is known from several skull bones, an almost complete dorsal series, complete sacrum, several caudal vertebrae, pelvic girdle and almost complete hind limbs. The new abelisaurid shows a unique set of features in the skull and several anatomical differences from Carnotaurus sastrei (the only other abelisaurid known from the La Colonia Formation). Koleken inakayali is retrieved as a brachyrostran abelisaurid, clustered with other South American abelisaurids from the latest Cretaceous (Campanian-Maastrichtian), such as Aucasaurus, Niebla and Carnotaurus. Leveraging our phylogeny estimates, we explore rates of morphological evolution across ceratosaurian lineages, finding them to be particularly high for elaphrosaurine noasaurids and around the base of Abelisauridae, before the Early Cretaceous radiation of the latter clade. The Noasauridae and their sister clade show contrasting patterns of morphological evolution, with noasaurids undergoing an early phase of accelerated evolution of the axial and hind limb skeleton in the Jurassic, and the abelisaurids exhibiting sustained high rates of cranial evolution during the Early Cretaceous. These results provide much needed context for the evolutionary dynamics of ceratosaurian theropods, contributing to broader understanding of macroevolutionary patterns across dinosaurs.

The evolution of femoral morphology in giant non-avian theropod dinosaurs.

Theropods are obligate bipedal dinosaurs that appeared 230 million years ago and are still extant as birds. Their history is characterized by extreme variations in body mass, with gigantism evolving convergently between many lineages. However, no quantification of hindlimb functional morphology has shown if these body mass increases led to similar specializations between distinct lineages. Here we studied femoral shape variation across 41 species of theropods (n= 68 specimens) using a high-density 3D geometric morphometric approach. We demonstrated that the heaviest theropods evolved wider epiphyses and a more distally located fourth trochanter, as previously demonstrated in early archosaurs, along with an upturned femoral head and a mediodistal crest that extended proximally along the shaft. Phylogenetically informed analyses highlighted that these traits evolved convergently within six major theropod lineages, regardless of their maximum body mass. Conversely, the most gracile femora were distinct from the rest of the dataset, which we interpret as a femoral specialization to "miniaturization" evolving close to Avialae (bird lineage). Our results support a gradual evolution of known "avian" features, such as the fusion between lesser and greater trochanters and a reduction of the epiphyses' offset, independently from body mass variations, which may relate to a more "avian" type of locomotion (more knee-than hip-driven). The distinction between body mass variations and a more "avian" locomotion is represented by a decoupling in the mediodistal crest morphology, whose biomechanical nature should be studied to better understand the importance of its functional role in gigantism, miniaturization and higher parasagittal abilities.

The last ceratosaur of Asia: a new noasaurid from the Early Cretaceous Great Siberian Refugium.

The noasaurid ceratosaur Kiyacursor longipes gen. et sp. nov. is described based on a fragmentary skeleton including cervical vertebra, pectoral girdle, humerus and hind limbs from the Lower Cretaceous (Aptian) Ilek Formation at Shestakovo 1 locality in Western Siberia, Russia. This is the first ceratosaur from the Early Cretaceous of Asia, extending the stratigraphic range of Ceratosauria by 40 Myr on that continent. Kiyacursor shares unique hind limb proportions with Elaphrosaurus and Limusaurus, suggesting improved cursorial ability. These taxa show an ostrich-like specialization of the pes, with a large third metatarsal and greatly reduced second metatarsal. By contrast, all other fast running non-avian theropod dinosaurs have an arctometatarsalian pes, with the third metatarsal strongly reduced proximally. The new taxon lived in the Early Cretaceous ecosystem containing a number of other Jurassic relics, such as stem salamanders, protosuchian and shartegosuchid crocodyliforms, tritylodontid synapsids and docodontan mammaliaforms.

Morphological disparity and structural performance of the dromaeosaurid skull informs ecology and evolutionary history

Non-avialan theropod dinosaurs had diverse ecologies and varied skull morphologies. Previous studies of theropod cranial morphology mostly focused on higher-level taxa or characteristics associated with herbivory. To better understand morphological disparity and function within carnivorous theropod families, here we focus on the Dromaeosauridae, ‘raptors’ traditionally seen as agile carnivorous hunters.We applied 2D geometric morphometrics to quantify skull shape, performed mechanical advantage analysis to assess the efficiency of bite force transfer, and performed finite element analysis to examine strain distribution in the skull during biting. We find that dromaeosaurid skull morphology was less disparate than most non-avialan theropod groups. Their skulls show a continuum of form between those that are tall and short and those that are flat and long. We hypothesise that this narrower morphological disparity indicates developmental constraint on skull shape, as observed in some mammalian families. Mechanical advantage indicates that Dromaeosaurus albertensis and Deinonychus antirrhopus were adapted for relatively high bite forces, while Halszkaraptor escuilliei was adapted for high bite speed, and other dromaeosaurids for intermediate bite forces and speeds. Finite element analysis indicates regions of high strain are consistent within dromaeosaurid families but differ between them. Average strain levels do not follow any phylogenetic pattern, possibly due to ecological convergence between distantly-related taxa.Combining our new morphofunctional data with a re-evaluation of previous evidence, we find piscivorous reconstructions of Halszkaraptor escuilliei to be unlikely, and instead suggest an invertivorous diet and possible adaptations for feeding in murky water or other low-visibility conditions. We support Deinonychus antirrhopus as being adapted for taking large vertebrate prey, but we find that its skull is relatively less resistant to bite forces than other dromaeosaurids. Given the recovery of high bite force resistance for Velociraptor mongoliensis, which is believed to have regularly engaged in scavenging behaviour, we suggest that higher bite force resistance in a dromaeosaurid taxon may reflect a greater reliance on scavenging rather than fresh kills.Comparisons to the troodontid Gobivenator mongoliensis suggest that a gracile rostrum like that of Velociraptor mongoliensis is ancestral to their closest common ancestor (Deinonychosauria) and the robust rostra of Dromaeosaurus albertensis and Deinonychus antirrhopus are a derived condition. Gobivenator mongoliensis also displays a higher jaw mechanical advantage and lower resistance to bite force than the examined dromaeosaurids, but given the hypothesised ecological divergence of troodontids from dromaeosaurids it is unclear which group, if either, represents the ancestral condition. Future work extending sampling to troodontids would therefore be invaluable and provide much needed context to the origin of skull form and function in early birds. This study illustrates how skull shape and functional metrics can discern non-avialan theropod ecology at lower taxonomic levels and identify variants of carnivorous feeding.

Cranial functional specialisation for strength precedes morphological evolution in Oviraptorosauria

Oviraptorosaurians were a theropod dinosaur group that reached high diversity in the Late Cretaceous. Within oviraptorosaurians, the later diverging oviraptorids evolved distinctive crania which were extensively pneumatised, short and tall, and had a robust toothless beak, interpreted as providing a powerful bite for their herbivorous to omnivorous diet. The present study explores the ability of oviraptorid crania to resist large mechanical stresses compared with other theropods and where this adaptation originated within oviraptorosaurians. Digital 3D cranial models were constructed for the earliest diverging oviraptorosaurian, Incisivosaurus gauthieri, and three oviraptorids, Citipati osmolskae, Conchoraptor gracilis, and Khaan mckennai. Finite element analyses indicate oviraptorosaurian crania were stronger than those of other herbivorous theropods (Erlikosaurus and Ornithomimus) and were more comparable to the large, carnivorous Allosaurus. The cranial biomechanics of Incisivosaurus align with oviraptorids, indicating an early establishment of distinctive strengthened cranial biomechanics in Oviraptorosauria, even before the highly modified oviraptorid cranial morphology. Bite modelling, using estimated muscle forces, suggests oviraptorid crania may have functioned closer to structural safety limits. Low mechanical stresses around the beaks of oviraptorids suggest a convergently evolved, functionally distinct rhamphotheca, serving as a cropping/feeding tool rather than for stress reduction, when compared with other herbivorous theropods.

Prolonged faunal turnover in earliest ants revealed by North American Cretaceous amber

All ∼14,000 extant ant species descended from the same common ancestor, which lived ∼140-120 million years ago (Ma).1,2 While modern ants began to diversify in the Cretaceous, recent fossil evidence has demonstrated that older lineages concomitantly occupied the same ancient ecosystems.3 These early-diverging ant lineages, or stem ants, left no modern descendants; however, they dominated the fossil record throughout the Cretaceous until their ultimate extinction sometime around the K-Pg boundary. Even as stem ant lineages appear to be diverse and abundant throughout the Cretaceous, the extent of their longevity in the fossil record and circumstances contributing to their extinction remain unknown.3 Here we report the youngest stem ants, preserved in ∼77 Ma Cretaceous amber from North Carolina, which illustrate unexpected morphological stability and lineage persistence in this enigmatic group, rivaling the longevity of contemporary ants. Through phylogenetic reconstruction and morphometric analyses, we find evidence that total taxic turnover in ants was not accompanied by a fundamental morphological shift, in contrast to other analogous stem extinctions such as theropod dinosaurs. While stem taxa showed broad morphological variation, high-density ant morphospace remained relatively constant through the last 100 million years, detailing a parallel, but temporally staggered, evolutionary history of modern and stem ants.

A remarkable assemblage of petroglyphs and dinosaur footprints in Northeast Brazil

The Serrote do Letreiro Site, found on the northwest periphery of the Sousa Basin, Brazil, presents a remarkable convergence of paleontological and archaeological elements. It is constituted of sub-horizontal "lajeiros", or rock outcrops, intermingled with endemic Caatinga vegetation. The three prominent outcrops feature fossilized footprints of theropod, sauropod, and iguanodontian dinosaurs from the Early Cretaceous Period. Adjacent to these dinosaur tracks, indigenous petroglyphs adorn the surface. The petroglyphs, mainly characterized by circular motifs, maintain a striking resemblance to other petroglyphs found in the states of Paraíba and Rio Grande do Norte. This study primarily endeavors to delineate the site's major characteristics while concentrating on the relationship between the dinosaur footprints and the petroglyphs. It concurrently assesses the preservation status of this invaluable record, shedding light on its implications for the realms of paleontology, archaeology, and cultural heritage studies.

Reassessment of the enigmatic Late Cretaceous theropod dinosaur, Bagaraatan ostromi

Abstract The Late Cretaceous Bagaraatan ostromi, described by Osmólska in 1996, is one of the most enigmatic theropod dinosaurs. The holotype possesses a peculiar combination of features, which Osmólska suggested were indicative of a primitive position among theropods that could not be resolved further. Other researchers have pointed to affinities with either derived bird-like coelurosaurs or tyrannosaurids. Here, we reanalyse all material collected by Osmólska, which reveals it to be a chimaera of multiple theropod taxa. The femur, tibiotarsus, one pedal phalanx, and most of the bones undescribed in Osmólska’s paper are identified as Caenagnathidae indet. The mandible, cervical vertebrae, pelvis, tail, and one pedal phalanx, which we consider the holotype of B. ostromi, show tyrannosaurid affinities, which are here supported by phylogenetic analyses. We find only two potentially unique diagnostic features of the Bagaraatan holotype: double surangular foramina and a horizontal ridge on the lateral surface of the postacetabular process of the ilium. Both, however, may be ontogenetically or intraspecifically variable, and thus we conclude that the holotype of B. ostromi represents an indeterminate tyrannosaurid. The small size of the holotype and its possession of many features known to characterize juvenile Tyrannosaurus rex indicate that the skeleton belongs to a juvenile, which is one of the smallest juvenile tyrannosaurids currently known.

Quantifying the effects of exceptional fossil preservation on the global availability of phylogenetic data in deep time.

Fossil deposits with exceptional preservation ("lagerstätten") provide important details not typically preserved in the fossil record, such that they hold an outsized influence on our understanding of biodiversity and evolution. In particular, the potential bias imparted by this so-called "lagerstätten effect" remains a critical, but underexplored aspect of reconstructing evolutionary relationships. Here, we quantify the amount of phylogenetic information available in the global fossil records of 1,327 species of non-avian theropod dinosaurs, Mesozoic birds, and fossil squamates (e.g., lizards, snakes, mosasaurs), and then compare the influence of lagerstätten deposits on phylogenetic information content and taxon selection in phylogenetic analyses to other fossil-bearing deposits. We find that groups that preserve a high amount of phylogenetic information in their global fossil record (e.g., non-avian theropods) are less vulnerable to a "lagerstätten effect" that leads to disproportionate representation of fossil taxa from one geologic unit in an evolutionary tree. Additionally, for each taxonomic group, we find comparable amounts of phylogenetic information in lagerstätten deposits, even though corresponding morphological character datasets vary greatly. Finally, we unexpectedly find that ancient sand dune deposits of the Late Cretaceous Gobi Desert of Mongolia and China exert an anomalously large influence on the phylogenetic information available in the squamate fossil record, suggesting a "lagerstätten effect" can be present in units not traditionally considered lagerstätten. These results offer a phylogenetics-based lens through which to examine the effects of exceptional fossil preservation on biological patterns through time and space, and invites further quantification of evolutionary information in the rock record.

Avialan-like brain morphology in Sinovenator (Troodontidae, Theropoda)

Many modifications to the skull and brain anatomy occurred along the lineage encompassing non-avialan theropod dinosaurs and modern birds. Anatomical changes to the endocranium include an enlarged endocranial cavity, relatively larger optic lobes that imply elevated visual acuity, and proportionately smaller olfactory bulbs that suggest reduced olfactory capacity. Here, we use micro-computed tomographic (μCT) imaging to reconstruct the endocranium and its neuroanatomical features from an exceptionally well-preserved skull of Sinovenator changii (Troodontidae, Theropoda). While its overall morphology resembles the typical endocranium of other troodontids, Sinovenator also exhibits unique endocranial features that are similar to other paravian taxa and non-maniraptoran theropods. Landmark-based geometric morphometric analysis on endocranial shape of non-avialan and avialan dinosaurs points to the overall brain morphology of Sinovenator most closely resembling that of Archaeopteryx, thus indicating acquisition of avialan-grade brain morphology in troodontids and wide existence of such architecture in Maniraptora.