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The Late Triassic fauna of the Lossiemouth Sandstone Formation (LSF) from the Elgin area, Scotland, has been pivotal in expanding our understanding of Triassic terrestrial tetrapods. Frustratingly, due to their odd preservation, interpretations of the Elgin Triassic specimens have relied on destructive moulding techniques, which only provide incomplete, and potentially distorted, information. Here, we show that micro-computed tomography (μCT) could revitalise the study of this important assemblage. We describe a long-neglected specimen that was originally identified as a pseudosuchian archosaur, Ornithosuchus woodwardi. μCT scans revealed dozens of bones belonging to at least two taxa: a small-bodied pseudosuchian and a specimen of the procolophonid Leptopleuron lacertinum. The pseudosuchian skeleton possesses a combination of characters that are unique to the clade Erpetosuchidae. As a basis for investigating the phylogenetic relationships of this new specimen, we reviewed the anatomy, taxonomy and systematics of other erpetosuchid specimens from the LSF (all previously referred to Erpetosuchus). Unfortunately, due to the differing representation of the skeleton in the available Erpetosuchus specimens, we cannot determine whether the erpetosuchid specimen we describe here belongs to Erpetosuchus granti (to which we show it is closely related) or if it represents a distinct new taxon. Nevertheless, our results shed light on rarely preserved details of erpetosuchid anatomy. Finally, the unanticipated new information extracted from both previously studied and neglected specimens suggests that fossil remains may be much more widely distributed in the Elgin quarries than previously recognised, and that the richness of the LSF might have been underestimated.
Dinosaur body fossil material is rare in Scotland, previously known almost exclusively from the Great Estuarine Group on the Isle of Skye. We report the first unequivocal dinosaur fossil from the Isle of Eigg, belonging to a Bathonian (Middle Jurassic) taxon of uncertain affinity. The limb bone NMS G.2020.10.1 is incomplete, but through a combination of anatomical comparison and osteohistology, we determine it most likely represents a stegosaur fibula. The overall proportions and cross-sectional geometry are similar to the fibulae of thyreophorans. Examination of the bone microstructure reveals a high degree of remodelling and randomly distributed longitudinal canals in the remaining primary cortical bone. This contrasts with the histological signal expected of theropod or sauropod limb bones, but is consistent with previous studies of thyreophorans, specifically stegosaurs. Previous dinosaur material from Skye and broadly contemporaneous sites in England belongs to this group, including Loricatosaurus and Sarcolestes and a number of indeterminate stegosaur specimens. Theropods such as Megalosaurus and sauropods such as Cetiosaurus are also known from these localities. Although we find strong evidence for a stegosaur affinity, diagnostic features are not observed on NMS G.2020.10.1, preventing us from referring it to any known genera. The presence of this large-bodied stegosaur on Eigg adds a significant new datapoint for dinosaur distribution in the Middle Jurassic of Scotland.
Taxonomic diversity and morphological disparity are different measures of biodiversity that together can describe large-scale evolutionary patterns. Diversity measures are often corrected by extending lineages back in time or adding additional taxa necessitated by a phylogeny, but disparity analyses focus on observed taxa only. This is problematic because some morphologies required by phylogeny are not included, some of which may help fill poorly sampled time bins. Moreover the taxic nature of disparity analyses makes it difficult to compare disparity measures with phylogenetically corrected diversity or morphological evolutionary rate curves. We present a general method for using phylogeny to correct measures of disparity, by including reconstructed ancestors in the disparity analysis. We apply this method to discrete character data sets focusing on Triassic archosaurs, Cenozoic carnivoramorph mammals, and Cretaceous-Cenozoic euarchontogliran mammals. Phylogenetic corrections do not simply mirror the taxic disparity patterns, but affect the three analyses in heterogeneous ways. Adding reconstructed ancestors can inflate morphospace, and the amount and direction of expansion differs depending on the taxonomic group in question. In some cases phylogenetic corrections give a temporal disparity curve indistinguishable from the taxic trend, but in other cases disparity is elevated in earlier time intervals relative to later bins, due to the extension of unsampled morphologies further back in time. The phylogenetic disparity curve for archosaurs differs little from the taxic curve, supporting a previously documented pattern of decoupled disparity and rates of morphological change in dinosaurs and their early contemporaries. Although phylogenetic corrections should not be used blindly, they are helpful when studying clades with major unsampled gaps in their fossil records.
The Middle Jurassic is a poorly sampled time interval for non-pelagic neosuchian crocodyliforms, which obscures our understanding of the origin and early evolution of major clades. Here we report a lower jaw from the Middle Jurassic (Bathonian) Duntulm Formation of the Isle of Skye, Scotland, UK, which consists of an isolated and incomplete left dentary and part of the splenial. Morphologically, the Skye specimen closely resembles the Cretaceous neosuchians Pachycheilosuchus and Pietraroiasuchus, in having a proportionally short mandibular symphysis, shallow dentary alveoli and inferred weakly heterodont dentition. It differs from other crocodyliforms in that the Meckelian canal is dorsoventrally expanded posterior to the mandibular symphysis and drastically constricted at the 7th alveolus. The new specimen, together with the presence of Theriosuchus sp. from the Valtos Formation and indeterminate neosuchians from the Kilmaluag Formation, indicates the presence of a previously unrecognised, diverse crocodyliform fauna in the Middle Jurassic of Skye, and Europe more generally. Small-bodied neosuchians were present, and ecologically and taxonomically diverse, in nearshore environments in the Middle Jurassic of the UK.
Fossils of Mesozoic terrestrial vertebrates from Lithuania and the wider East Baltic region of Europe have previously been unknown. We here report the first Mesozoic terrestrial vertebrate fossils from Lithuania: two premaxillary specimens and three teeth that belong to Phytosauria, a common clade of semiaquatic Triassic archosauriforms. These specimens represent an uncrested phytosaur, similar to several species within the genera Paleorhinus, Parasuchus, Rutiodon and Nicrosaurus. Because phytosaurs are currently only known from the Upper Triassic, their discovery in northwestern Lithuania (the Šaltiškiai clay-pit) suggests that at least part of the Triassic succession in this region is Late Triassic in age, and is not solely Early Triassic as has been previously considered. The new specimens are among the most northerly occurrences of phytosaurs in the Late Triassic, as Lithuania was approximately 7–10° further north than classic phytosaur-bearing localities in nearby Germany and Poland, and as much as 40° further north than the best-sampled phytosaur localities in North America. The far northerly occurrence of the Lithuanian fossils prompts a review of phytosaur biogeography and distribution, which suggests that these predators were widely distributed in the Triassic monsoonal belt but rarer in more arid regions.
The rise of archosaurs during the Triassic and Early Jurassic has been treated as a classic example of an evolutionary radiation in the fossil record. This paper reviews published studies and provides new data on archosaur lineage origination, diversity and lineage evolution, morphological disparity, rates of morphological character change, and faunal abundance during the Triassic–Early Jurassic. The fundamental archosaur lineages originated early in the Triassic, in concert with the highest rates of character change. Disparity and diversity peaked later, during the Norian, but the most significant increase in disparity occurred before maximum diversity. Archosaurs were rare components of Early–Middle Triassic faunas, but were more abundant in the Late Triassic and pre-eminent globally by the Early Jurassic. The archosaur radiation was a drawn-out event and major components such as diversity and abundance were discordant from each other. Crurotarsans (crocodile-line archosaurs) were more disparate, diverse, and abundant than avemetatarsalians (bird-line archosaurs, including dinosaurs) during the Late Triassic, but these roles were reversed in the Early Jurassic. There is no strong evidence that dinosaurs outcompeted or gradually eclipsed crurotarsans during the Late Triassic. Instead, crurotarsan diversity decreased precipitously by the end-Triassic extinction, which helped usher in the age of dinosaurian dominance.
The Middle Jurassic was a critical time in the evolution of theropod dinosaurs, highlighted by the origination and radiation of the large-bodied and morphologically diverse Tetanurae. Middle Jurassic tetanurans are rare but have been described from Europe, South America and China. In particular, China has yielded a number of potential basal tetanurans, but these have received little detailed treatment in the literature. Here we redescribe the postcranial skeleton of one of the most complete Chinese Middle Jurassic theropods, Monolophosaurus. Several features confirm the tetanuran affinities of Monolophosaurus, but the possession of ‘primitive’ traits such as a double-faceted pubic peduncle of the ilium and a hood-like supracetabular crest suggest a basal position within Tetanurae. This conflicts with most published cladistic analyses that place Monolophosaurus in a more derived position within Allosauroidea. We review the Middle Jurassic record of Chinese theropods and compare Monolophosaurus to other Middle Jurassic theropods globally. These comparisons suggest that Monolophosaurus and Chuandongocoelurus formed an endemic theropod clade limited to the Middle Jurassic of Asia. Other Middle Jurassic Chinese theropods deserve further study.
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