Predator-prey dynamics are an important evolutionary driver of escalating predation mode and efficiency, and commensurate responses of prey [1-3]. Among these strategies, camouflage is important for visual concealment, with countershading the most universally observed [4-6]. Extant terrestrial herbivores free of significant predation pressure, due to large size or isolation, do not exhibit countershading. Modern predator-prey dynamics may not be directly applicable to those of the Mesozoic due to the dominance of very large, visually oriented theropod dinosaurs . Despite thyreophoran dinosaurs' possessing extensive dermal armor, some of the most extreme examples of anti-predator structures [8, 9], little direct evidence of predation on these and other dinosaur megaherbivores has been documented. Here we describe a new, exquisitely three-dimensionally preserved nodosaurid ankylosaur, Borealopelta markmitchelli gen. et sp. nov., from the Early Cretaceous of Alberta, which preserves integumentary structures as organic layers, including continuous fields of epidermal scales and intact horn sheaths capping the body armor. We identify melanin in the organic residues through mass spectroscopic analyses and observe lighter pigmentation of the large parascapular spines, consistent with display, and a pattern of countershading across the body. With an estimated body mass exceeding 1,300 kg, B. markmitchelli was much larger than modern terrestrial mammals that either are countershaded or experience significant predation pressure as adults. Presence of countershading suggests predation pressure strong enough to select for concealment in this megaherbivore despite possession of massive dorsal and lateral armor, illustrating a significant dichotomy between Mesozoic predator-prey dynamics and those of modern terrestrial systems.
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.
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.
Few ankylosaurs are known from more than a single specimen, but the ankylosaurid Euoplocephalus tutus (from the Late Cretaceous of Alberta, Canada and Montana, USA) is represented by dozens of skulls and partial skeletons, and is therefore an important taxon for understanding intraspecific variation in ankylosaurs. Euoplocephalus is unusual compared to other dinosaurs from the Late Cretaceous of Alberta because it is recognized from the Dinosaur Park, Horseshoe Canyon, and Two Medicine formations. A comprehensive review of material attributed to Euoplocephalus finds support for the resurrection of its purported synonyms Anodontosaurus lambei and Scolosaurus cutleri, and the previously resurrected Dyoplosaurus acutosquameus. Anodontosaurus is found primarily in the Horseshoe Canyon Formation of Alberta and is characterized by ornamentation posterior to the orbits and on the first cervical half ring, and wide, triangular knob osteoderms. Euoplocephalus is primarily found in Megaherbivore Assemblage Zone 1 in the Dinosaur Park Formation of Alberta and is characterized by the absence of ornamentation posterior to the orbits and on the first cervical half ring, and keeled medial osteoderms on the first cervical half ring. Scolosaurus is found primarily in the Two Medicine Formation of Montana (although the holotype is from Dinosaur Provincial Park), and is characterized by long, back-swept squamosal horns, ornamentation posterior to the orbit, and low medial osteoderms on the first cervical half ring; Oohkotokia horneri is morphologically indistinguishable from Scolosaurus cutleri. Dyoplosaurus was previously differentiated from Euoplocephalus sensu lato by the morphology of the pelvis and pes, and these features also differentiate Dyoplosaurus from Anodontosaurus and Scolosaurus; a narrow tail club knob is probably also characteristic for Dyoplosaurus.
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.
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.
Ankylosaurs are one of the least explored clades of dinosaurs regarding endocranial anatomy, with few available descriptions of braincase anatomy and even less information on brain and inner ear morphologies. The main goal of this study is to provide a detailed description of the braincase and internal structures of the Early Cretaceous nodosaurid Pawpawsaurus campbelli, based on recently made CT scans.
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids- the Struthiosaurinae.
The ankylosaurs are one of the major dinosaur groups and are characterized by unique body armor. Previous studies on other dinosaur taxa have revealed growth patterns, life history and evolutionary mechanisms based on their long bone histology. However, to date nothing is known about long bone histology in the Ankylosauria. This study is the first description of ankylosaurian long bone histology based on several limb elements, which were sampled from different individuals from the Ankylosauridae and Nodosauridae. The histology is compared to that of other dinosaur groups, including other Thyreophora and Sauropodomorpha. Ankylosaur long bone histology is characterized by a fibrolamellar bone architecture. The bone matrix type in ankylosaurs is closest to that of Stegosaurus. A distinctive mixture of woven and parallel-fibered bone together with overall poor vascularization indicates slow growth rates compared to other dinosaurian taxa. Another peculiar characteristic of ankylosaur bone histology is the extensive remodeling in derived North American taxa. In contrast to other taxa, ankylosaurs substitute large amounts of their primary tissue early in ontogeny. This anomaly may be linked to the late ossification of the ankylosaurian body armor. Metabolically driven remodeling processes must have liberated calcium to ossify the protective osteodermal structures in juveniles to subadult stages, which led to further remodeling due to increased mechanical loading. Abundant structural fibers observed in the primary bone and even in remodeled bone may have improved the mechanical properties of the Haversian bone.
A new ankylosaurid (Ankylosauria: Dinosauria), Ziapelta sanjuanensis, gen. et sp. nov., is based on a complete skull, an incomplete first cervical half ring, a possible fragment of the second cervical half ring, and additional fragmentary osteoderms. The holotype specimen is from the Upper Cretaceous (Upper Campanian, Kirtlandian Land-Vertebrate Age) Kirtland Formation (De-na-zin Member) at Hunter Wash, San Juan Basin, in northwestern New Mexico, USA. Diagnostic characters of Ziapelta include: a large, prominent triangular median nasal caputegulum; a mixture of flat and bulbous frontonasal caputegulae; ventrolaterally oriented squamosal horns with a sharp, prominent dorsal keel; and the ventral surface of basicranium with three prominent anteroposteriorly oriented fossae. A phylogenetic analysis suggests that Ziapelta is not closely related to the other ankylosaurid from the De-na-zin Member, Nodocephalosaurus, but allies it to the northern North American ankylosaurids Ankylosaurus, Anodontosaurus, Euoplocephalus, Dyoplosaurus, and Scolosaurus.