SciCombinator

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Concept: Ornithischia

170

A fragmentary cervico-pectoral lateral spine and partial humerus of an ankylosaur from the Early Cretaceous (early Valanginian) of Gronau in Westfalen, northwestern Germany, are described. The spine shows closest morphological similarities to the characteristic cervical and pectoral spines of Hylaeosaurus armatus from the late Valanginian of England. An extensive comparison of distal humeri among thyreophoran dinosaurs supports systematic differences in the morphology of the distal condyli between Ankylosauria and Stegosauria and a referral of the Gronau specimen to the former. The humerus fragment indicates a rather small individual, probably in the size range of H. armatus, and both specimens are determined herein as ?Hylaeosaurus sp.. A short overview of other purported ankylosaur material from the Berriasian-Valanginian of northwest Germany shows that, aside from the material described herein, only tracks can be attributed to this clade with confidence at present.

Concepts: Cretaceous, Dinosaur, Iguanodon, Ankylosauria, Thyreophora, Ornithischia, Early Cretaceous, Hylaeosaurus

94

For 130 years, dinosaurs have been divided into two distinct clades-Ornithischia and Saurischia. Here we present a hypothesis for the phylogenetic relationships of the major dinosaurian groups that challenges the current consensus concerning early dinosaur evolution and highlights problematic aspects of current cladistic definitions. Our study has found a sister-group relationship between Ornithischia and Theropoda (united in the new clade Ornithoscelida), with Sauropodomorpha and Herrerasauridae (as the redefined Saurischia) forming its monophyletic outgroup. This new tree topology requires redefinition and rediagnosis of Dinosauria and the subsidiary dinosaurian clades. In addition, it forces re-evaluations of early dinosaur cladogenesis and character evolution, suggests that hypercarnivory was acquired independently in herrerasaurids and theropods, and offers an explanation for many of the anatomical features previously regarded as notable convergences between theropods and early ornithischians.

Concepts: Bird, Phylogenetics, Dinosaur, Sauropodomorpha, Saurischia, Theropoda, Coelurosauria, Ornithischia

77

Minmi is the only known genus of ankylosaurian dinosaur from Australia. Seven specimens are known, all from the Lower Cretaceous of Queensland. Only two of these have been described in any detail: the holotype specimen Minmi paravertebra from the Bungil Formation near Roma, and a near complete skeleton from the Allaru Mudstone on Marathon Station near Richmond, preliminarily referred to a possible new species of Minmi. The Marathon specimen represents one of the world’s most complete ankylosaurian skeletons and the best-preserved dinosaurian fossil from eastern Gondwana. Moreover, among ankylosaurians, its skull is one of only a few in which the majority of sutures have not been obliterated by dermal ossifications or surface remodelling. Recent preparation of the Marathon specimen has revealed new details of the palate and narial regions, permitting a comprehensive description and thus providing new insights cranial osteology of a basal ankylosaurian. The skull has also undergone computed tomography, digital segmentation and 3D computer visualisation enabling the reconstruction of its nasal cavity and endocranium. The airways of the Marathon specimen are more complicated than non-ankylosaurian dinosaurs but less so than derived ankylosaurians. The cranial (brain) endocast is superficially similar to those of other ankylosaurians but is strongly divergent in many important respects. The inner ear is extremely large and unlike that of any dinosaur yet known. Based on a high number of diagnostic differences between the skull of the Marathon specimen and other ankylosaurians, we consider it prudent to assign this specimen to a new genus and species of ankylosaurian. Kunbarrasaurus ieversi gen. et sp. nov. represents the second genus of ankylosaurian from Australia and is characterised by an unusual melange of both primitive and derived characters, shedding new light on the evolution of the ankylosaurian skull.

Concepts: Mammal, Cretaceous, Facial nerve, Dinosaur, Ankylosauridae, Ankylosauria, Thyreophora, Ornithischia

45

Although the evolution and function of “exaggerated” bony projections in ornithischian dinosaurs has been subject to significant debate recently, our understanding of the structure and morphology of their epidermal keratinized coverings is greatly limited. The holotype of Borealopelta, a new nodosaurid ankylosaur, preserves osteoderms and extensive epidermal structures (dark organic residues), in anatomic position across the entire precaudal length. Contrasting previous specimens, organic epiosteodermal scales, often in the form of horn-like (keratinous) sheaths, cap and exaggerate nearly all osteoderms, allowing for morphometric and allometric analyses of both the bony osteoderms and their horny sheaths. A total of 172 osteoderms were quantified, with osteoderm spine length and height being positively allometric with respect to basal length and width. Despite tight correlations between the different measures amongst all other osteoderms, the large parascapular spines represent consistent outliers. Thickness and relative contribution of the keratinized epiosteodermal scales/sheaths varies greatly by region, ranging from 2% to 6% for posterior thoracics, to ∼25% (1.3×) for the parascapular spines-similar to horn sheaths in some bovid analogues. Relative to the bony cores, the horny portions of the spines are strongly positively allometric (slope = 2.3, CI = 1.8-2.8). Strong allometric scaling, species-specific morphology, and significant keratinous extension of the cervicoscapular spines is consistent with elaboration under socio-sexual selection. This marks the first allometric analysis of ornithischian soft tissues.

Concepts: Biology, Anatomy, Height, Length, Allometry, Dinosaur, Ankylosauria, Ornithischia

41

We describe a new species of an ornithischian dinosaur, Isaberrysaura mollensis gen. et sp. nov. The specimen, consisting in an almost complete skull and incomplete postcranium was collected from the marine-deltaic deposits of the Los Molles Formation (Toarcian-Bajocian), being the first reported dinosaur for this unit, one of the oldest from Neuquén Basin, and the first neornithischian dinosaur known from the Jurassic of South America. Despite showing a general stegosaurian appearance, the extensive phylogenetic analysis carried out depicts Isaberrysaura mollensis gen. et sp. nov. as a basal ornithopod, suggesting that both Thyreophora and neornithischians could have achieved significant convergent features. The specimen was preserved articulated and with some of its gut content place in the middle-posterior part of the thoracic cavity. Such stomach content was identified as seeds, most of them belonging to the Cycadales group. This finding reveals a possible and unexpected role of this ornithischian species as seed-dispersal agent.

Concepts: Biology, South America, Dinosaur, Edmontosaurus, Jurassic, Ankylosauria, Thyreophora, Ornithischia

36

Middle Jurassic to Early Cretaceous deposits from northeastern China have yielded varied theropod dinosaurs bearing feathers. Filamentous integumentary structures have also been described in ornithischian dinosaurs, but whether these filaments can be regarded as part of the evolutionary lineage toward feathers remains controversial. Here we describe a new basal neornithischian dinosaur from the Jurassic of Siberia with small scales around the distal hindlimb, larger imbricated scales around the tail, monofilaments around the head and the thorax, and more complex featherlike structures around the humerus, the femur, and the tibia. The discovery of these branched integumentary structures outside theropods suggests that featherlike structures coexisted with scales and were potentially widespread among the entire dinosaur clade; feathers may thus have been present in the earliest dinosaurs.

Concepts: Bird, Reptile, Dinosaur, Archaeopteryx, Theropoda, Coelurosauria, Tyrannosaurus, Ornithischia

34

Crocodyliforms serve as important taphonomic agents, accumulating and modifying vertebrate remains. Previous discussions of Mesozoic crocodyliform feeding in terrestrial and riverine ecosystems have often focused on larger taxa and their interactions with equally large dinosaurian prey. However, recent evidence suggests that the impact of smaller crocodyliforms on their environments should not be discounted. Here we present direct evidence of feeding by a small crocodyliform on juvenile specimens of a ‘hypsilophodontid’ dinosaur from the Upper Cretaceous (Campanian) Kaiparowits Formation of southern Utah. Diagnostic crocodyliform bite marks present on a left scapula and a right femur, as well as a partial probable crocodyliform tooth crown (ovoid in cross-section) preserved within a puncture on the right femur, comprise the bulk of the feeding evidence. Computed tomography scans of the femoral puncture reveal impact damage to the surrounding bone and that the distal tip of the embedded tooth was missing prior to the biting event. This is only the second reported incidence of a fossil crocodyliform tooth being found embedded directly into prey bone. These bite marks provide insight into the trophic interactions of the ecosystem preserved in the Kaiparowits Formation. The high diversity of crocodyliforms within this formation may have led to accentuated niche partitioning, which seems to have included juvenile dinosaurian prey.

Concepts: Ecology, Cretaceous, Fossil, Dinosaur, Cretaceous–Tertiary extinction event, Plesiosaur, Ornithischia, Crocodylomorpha

21

The partial skeleton of a leptoceratopsid dinosaur, Ischioceratops zhuchengensis gen. et sp. nov., was excavated from the bone-beds of the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China. This fossil represents the second leptoceratopsid dinosaur specimen recovered from the Kugou locality, a highly productive site in Zhucheng. The ischium of the new taxon is morphologically unique among known Dinosauria, flaring gradually to form an obturator process in its middle portion and resembling the shaft of a recurve bow. An elliptical fenestra perforates the obturator process, and the distal end of the shaft forms an axehead-shaped expansion. The discovery of Ischioceratops increases the known taxonomic diversity and morphological disparity of the Leptoceratopsidae.

Concepts: Cretaceous, Shandong, Paleontology, Liaoning, Dinosaur, Ornithischia, Late Cretaceous, Ceratopsia

16

Several skulls of the ornithischian dinosaur Lesothosaurus diagnosticus (Lower Jurassic, southern Africa) are known, but all are either incomplete, deformed, or incompletely prepared. This has hampered attempts to provide a comprehensive description of skull osteology in this crucial early dinosaurian taxon. Using visualization software, computed tomographic scans of the Lesothosaurus syntypes were digitally segmented to remove matrix, and identify and separate individual cranial and mandibular bones, revealing new anatomical details such as sutural morphology and the presence of several previously undescribed elements. Together with visual inspection of exposed skull bones, these CT data enable a complete description of skull anatomy in this taxon. Comparisons with our new data suggest that two specimens previously identified as Lesothosaurus sp. (MNHN LES 17 and MNHN LES 18) probably represent additional individuals of Lesothosaurus diagnosticus.

Concepts: Bone, Skull, Anatomical terms of location, Dinosaur, Jurassic, Diapsid, Ornithischia, Heterodontosauridae

14

The systematic relationships of taxa traditionally referred to as ‘basal ornithopods’ or ‘hypsilophodontids’ remain poorly resolved since it was discovered that these taxa are not a monophyletic group, but rather a paraphyletic set of neornithischian taxa. Thus, even as the known diversity of these taxa has dramatically increased over the past two decades, our knowledge of their placement relative to each other and the major ornithischian subclades remained incomplete. This study employs the largest phylogenetic dataset yet compiled to assess basal ornithischian relationships (255 characters for 65 species level terminal taxa). The resulting strict consensus tree is the most well-resolved, stratigraphically consistent hypothesis of basal ornithischian relationships yet hypothesized. The only non-iguanodontian ornithopod (=basal ornithopod) recovered in this analysis is Hypsilophodon foxii. The majority of former ‘hypsilophodontid’ taxa are recovered within a single clade (Parksosauridae) that is situated as the sister-taxon to Cerapoda. The Parksosauridae is divided between two subclades, the Orodrominae and the Thescelosaurinae. This study does not recover a clade consisting of the Asian taxa Changchunsaurus, Haya, and Jeholosaurus (=Jeholosauridae). Rather, the former two taxa are recovered as basal members of Thescelosaurinae, while the latter taxon is recovered in a clade with Yueosaurus near the base of Neornithischia.The endemic South American clade Elasmaria is recovered within the Thescelosaurinae as the sister taxon to Thescelosaurus. This study supports the origination of Dinosauria and the early diversification of Ornithischia within Gondwana. Neornithischia first arose in Africa by the Early Jurassic before dispersing to Asia before the late Middle Jurassic, where much of the diversification among non-cerapodan neornithischians occurred. Under the simplest scenario the Parksosauridae originated in North America, with at least two later dispersals to Asia and one to South America. However, when ghost lineages are considered, an alternate dispersal hypothesis has thescelosaurines dispersing from Asia into South America (via North America) during the Early Cretaceous, then back into North America in the latest Cretaceous. The latter hypothesis may explain the dominance of orodromine taxa prior to the Maastrichtian in North America and the sudden appearance and wide distribution of thescelosaurines in North America beginning in the early Maastrichtian. While the diversity of parksosaurids has greatly increased over the last fifteen years, a ghost lineage of over 40 myr is present between the base of Parksosauridae and Cerapoda, indicating that much of the early history and diversity of this clade is yet to be discovered. This new phylogenetic hypothesis provides a comprehensive framework for testing further hypotheses regarding evolutionary patterns and processes within Ornithischia.

Concepts: Phylogenetics, Cladistics, North America, South America, Cretaceous, Dinosaur, Ornithischia, Hypsilophodont