Nicholas Fraser
National Museums Scotland, Department of Natural Sciences, Faculty Member
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Archosauromorph reptiles underwent rapid lineage diversification, increases in morphological and body size disparity, and expansion into new adaptive landscapes. Several of the primary early archosauromorph clades (e.g. rhynchosaurs) are... more
Archosauromorph reptiles underwent rapid lineage diversification, increases in morphological and body size disparity, and expansion into new adaptive landscapes. Several of the primary early archosauromorph clades (e.g. rhynchosaurs) are easy to differentiate from others because of their characteristic body types, whereas the more lizard‐like and carnivorous forms with long necks (e.g. tanystropheids) were historically all relegated to the groups Protorosauria or Prolacertiformes. However, it is now clear that these groups are polyphyletic and that a lizard‐like, carnivorous form is plesiomorphic for Archosauromorpha, and multiple subclades started with that body plan. Among these early forms is Malerisaurus from the Upper Triassic of India (M. robinsonae) and the Upper Triassic of south‐western USA (M. langstoni). In this paper, we critically re‐evaluate the genus. We find both species of Malerisaurus as valid, and identify Malerisaurus as an early diverging, but late‐surviving, carnivorous member of Azendohsauridae within Allokotosauria. Our histological analysis and assessment of ontogenetic changes of limb bones of small and large individuals demonstrate that the skeletons of the small forms grew slowly and became more robust through ontogeny, and that the larger recovered bones are at or near the maximum size of the taxon. Malerisaurus and Malerisaurus‐like taxa were common members of the Otischalkian–Adamanian (late Carnian to mid‐Norian) faunal assemblages from Upper Triassic strata of the south western USA, but they are absent from the younger Revueltian holochronozone. Specimens from western North America show that Allokotosauria had a near‐Pangaean distribution for much of the Middle Triassic to Late Triassic.
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The historical clade “Protorosauria” represents an important group of archosauromorph reptiles that had a wide geographic distribution between the Late Permian and Late Triassic. “Protorosaurs” are characterized by their long necks, which... more
The historical clade “Protorosauria” represents an important group of archosauromorph reptiles that had a wide geographic distribution between the Late Permian and Late Triassic. “Protorosaurs” are characterized by their long necks, which are epitomized in the genusTanystropheusand inDinocephalosaurus orientalis. Recent phylogenetic analyses have indicated that “Protorosauria” is a polyphyletic clade, but the exact relationships of the various “protorosaur” taxa within the archosauromorph lineage is currently uncertain. Several taxa, although represented by relatively complete material, have previously not been assessed phylogenetically. We present a new phylogenetic hypothesis that comprises a wide range of archosauromorphs, including the most exhaustive sample of “protorosaurs” to date and several “protorosaur” taxa from the eastern Tethys margin that have not been included in any previous analysis. The polyphyly of “Protorosauria” is confirmed and therefore we suggest the usage o...
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We describe a new small-bodied coelophysoid theropod dinosaur, Pendraig milnerae gen. et sp. nov, from the Late Triassic fissure fill deposits of Pant-y-ffynnon in southern Wales. The species is represented by the holotype, consisting of... more
We describe a new small-bodied coelophysoid theropod dinosaur, Pendraig milnerae gen. et sp. nov, from the Late Triassic fissure fill deposits of Pant-y-ffynnon in southern Wales. The species is represented by the holotype, consisting of an articulated pelvic girdle, sacrum and posterior dorsal vertebrae, and an associated left femur, and by two referred specimens, comprising an isolated dorsal vertebra and a partial left ischium. Our phylogenetic analysis recovers P. milnerae as a non-coelophysid coelophysoid theropod, representing the first-named unambiguous theropod from the Triassic of the UK. Recently, it has been suggested that Pant-y-ffynnon and other nearby Late Triassic to Early Jurassic fissure fill faunas might have been subjected to insular dwarfism. To test this hypothesis for P. milnerae , we performed an ancestral state reconstruction analysis of body size in early neotheropods. Although our results indicate that a reduced body size is autapomorphic for P. milnerae , ...
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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... more
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.
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The Middle and early Late Triassic of southern China is well known for a remarkable diversity of marine vertebrates, particularly reptiles, including an abundance of intriguing new forms (e.g., Jiang et al., 2005; Hu et al., 2011; Li et... more
The Middle and early Late Triassic of southern China is well known for a remarkable diversity of marine vertebrates, particularly reptiles, including an abundance of intriguing new forms (e.g., Jiang et al., 2005; Hu et al., 2011; Li et al., 2016). Here we describe a new diapsid from Yunnan Province. It possesses an elongate neck that exhibits a remarkable similarity to that of many Protorosauria, yet in other respects the skull and postcranium are much less derived. The new taxon is part of the so-called Panxian-Luoping Fauna and the deposits correspond to the Upper Member of the Guanling Formation, comprising thin to medium bedded, gray to dark-gray laminated marly limestone and limestone, with several layers of bentonite intercalated in the fossil level at Panxian (Wan, 2002; Motani et al., 2008; Jiang et al., 2009). Their age is Pelsonian (middle Anisian, Middle Triassic) as is indicated by the conodont Nicoraella kockeli Zone (Sun et al., 2006; Zhang et al., 2009). A recent U-P...
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Docodonta are one of the earliest diverging groups of mammaliaforms, and their morphology provides key information on the transition between non-mammalian cynodonts and Mammalia. We describe the partial skulls of two docodontans... more
Docodonta are one of the earliest diverging groups of mammaliaforms, and their morphology provides key information on the transition between non-mammalian cynodonts and Mammalia. We describe the partial skulls of two docodontans Borealestes serendipitus and Borealestes cuillinensis sp. nov. from the Kilmaluag Formation (Middle Jurassic: Bathonian), Isle of Skye, Scotland. We visualize their cranial anatomy using laboratory and synchrotron X-ray micro-CT. The skulls belong to two partial skeletons, currently comprising the most complete Mesozoic mammal fossils reported from the British Isles. The associated upper and lower dentitions show that the lower dentition of Borealestes is not diagnostic to species level. We establish, B. cuillinensis, based on upper molar characters, and re-identify upper molars previously assigned to ‘Borealestes’ mussettae as belonging to B. cuillinensis. ‘Borealestes’ mussettae, based on distinctive lower molars, is found to be morphologically and phyloge...
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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,... more
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 anat...
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Newly discovered fossils of the Middle Triassic reptile Atopodentatus unicus call for a radical reassessment of its feeding behavior. The skull displays a pronounced hammerhead shape that was hitherto unknown. The long, straight anterior... more
Newly discovered fossils of the Middle Triassic reptile Atopodentatus unicus call for a radical reassessment of its feeding behavior. The skull displays a pronounced hammerhead shape that was hitherto unknown. The long, straight anterior edges of both upper and lower jaws were lined with batteries of chisel-shaped teeth, whereas the remaining parts of the jaw rami supported densely packed needle-shaped teeth forming a mesh. The evidence indicates a novel feeding mechanism wherein the chisel-shaped teeth were used to scrape algae off the substrate, and the plant matter that was loosened was filtered from the water column through the more posteriorly positioned tooth mesh. This is the oldest record of herbivory within marine reptiles.
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We describe a new species of the aetosaur Coahomasuchus, C. chathamensis, based on an incomplete, but largely articulated, anterior portion of a skeleton recovered from a quarry in the Upper Triassic Pekin Formation of Chatham County,... more
We describe a new species of the aetosaur Coahomasuchus, C. chathamensis, based on an incomplete, but largely articulated, anterior portion of a skeleton recovered from a quarry in the Upper Triassic Pekin Formation of Chatham County, North Carolina. This is only the second documented occurrence of Coahomasuchus, with the other being the holotype of C. kahleorum Heckert and Lucas, 1999 from the Upper Triassic Colorado City Formation of Texas. Although much of the specimen is the same size as the holotype of C. kahleorum, the dorsal paramedian osteoderms of the North Carolina taxon are considerably (~1.3×) wider than homologous counterparts in C. kahleorum, and the ventral thoracic osteoderms are also rectangular (~1.5× wider than long), rather than square, presumably to accommodate the wider body. This is a rare instance where two articulated specimens of closely related aetosaur species are available for direct comparison of homologous osteoderms. Isolated osteoderms with similar o...
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THE end of the Triassic period is pivotal in the evolution of modern ecosystems1. Despite this, the Triassic remains one of the poorest known periods in the evolutionary history of the terrestrial arthropods. Here we report on... more
THE end of the Triassic period is pivotal in the evolution of modern ecosystems1. Despite this, the Triassic remains one of the poorest known periods in the evolutionary history of the terrestrial arthropods. Here we report on fossiliferous shales preserving a nearly complete marginal lacustrine community from the Virginia-North Carolina border that sheds considerable light on this critical interval. Three species of insect were previously described from this locality, but the full extent and significance of its diversity have only now been discovered: we report here the oldest definitive records for three orders of insect and numerous families and super families. Furthermore, in addition to new taxa, the flora is shown to contain an unusual diversity of forms, some of which have only been previously reported either from Europe or the Southern Hemisphere. The abundance of complete insects and the preservation of soft-part anatomy on some of the vertebrates elevates the site to one of the most significant Lagerstätte in the world.
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Research Interests: Paleoecology and Mesozoic
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Contrary to the fast radiation of most metazoans after the end-Permian mass extinction, it is believed that early marine reptiles evolved slowly during the same time interval. However, emerging discoveries of Early Triassic marine... more
Contrary to the fast radiation of most metazoans after the end-Permian mass extinction, it is believed that early marine reptiles evolved slowly during the same time interval. However, emerging discoveries of Early Triassic marine reptiles are questioning this traditional view. Here we present an aberrant basal ichthyosauriform with a hitherto unknown body design that suggests a fast radiation of early marine reptiles. The new species is larger than coeval marine reptiles and has an extremely small head and a long tail without a fluke. Its heavily-built body bears flattened and overlapping gastral elements reminiscent of hupehsuchians. A phylogenetic analysis places the new species at the base of ichthyosauriforms, as the sister taxon of Cartorhynchus with which it shares a short snout with rostrally extended nasals. It now appears that ichthyosauriforms evolved rapidly within the first one million years of their evolution, in the Spathian (Early Triassic), and their true diversity ...
The Triassic period is generally viewed as the beginning of the Age of Dinosaurs. For paleontologists, however, it also marks the rise of the world&amp;amp;amp;#x27;s first modern land ecosystems. Over the past three decades,... more
The Triassic period is generally viewed as the beginning of the Age of Dinosaurs. For paleontologists, however, it also marks the rise of the world&amp;amp;amp;#x27;s first modern land ecosystems. Over the past three decades, extensive, worldwide fieldwork has led to the discovery of many new species of Triassic animals and plants, suggesting that faunal and floral changes already began in the Middle Triassic and were more protracted than previously thought. The Late Triassic is a pivotal time in the evolution of life on land, with many of the major groups ...
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ABSTRACT A new species of Macrocnemus is described on the basis of two incomplete specimens from the Lower Ladinian Prosanto Formation of southeastern Switzerland. The new form can be distinguished by its gracile limb elements and having... more
ABSTRACT A new species of Macrocnemus is described on the basis of two incomplete specimens from the Lower Ladinian Prosanto Formation of southeastern Switzerland. The new form can be distinguished by its gracile limb elements and having a noticeably longer tibia than either Macrocnemus bassanii or Macrocnemus fuyuanensis. One of the new specimens exhibits soft part preservation in the region of the pelvic girdle. The hind part of this individual is preserved fully articulated and allows a complete count of the caudal vertebrae to be made for the first time with 52 caudals. It is postulated that this specimen was predated upon and that the anterior part of the animal was consumed by a large predator such as a nothosaurid reptile or the actinopterygian fish Saurichthys.
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This book is the first attempt to collate all the information known to date on the small vertebrates, e.g. mammals, crocodiles, turtles, lizards, frogs, salamanders, etc., and features contributions by experts with international... more
This book is the first attempt to collate all the information known to date on the small vertebrates, e.g. mammals, crocodiles, turtles, lizards, frogs, salamanders, etc., and features contributions by experts with international reputations in their fields. There are chapters on the taxonomy and phylogeny of the key vertebrate groups followed by a section dealing with the most significant fossiliferous assemblages worldwide. The final section looks at how faunal turnover at this time is measured and examines the possibility of mass extinctions.
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... RIEPPEL Olivier ; FRASER Nicholas C. ; NOSOTTI Stefania ; Résumé / Abstract. This paper presents a preliminary analysis of protorosaur interrelationships based on the combination of the data published by Benton & Allen (1997),... more
... RIEPPEL Olivier ; FRASER Nicholas C. ; NOSOTTI Stefania ; Résumé / Abstract. This paper presents a preliminary analysis of protorosaur interrelationships based on the combination of the data published by Benton & Allen (1997), Jalil (1997), and Dilkes (1998). ...
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Atoposaurids were a clade of semiaquatic crocodyliforms known from the Late Jurassic to the latest Cretaceous. Tentative remains from Europe, Morocco, and Madagascar may extend their range into the Middle Jurassic. Here we report the... more
Atoposaurids were a clade of semiaquatic crocodyliforms known from the Late Jurassic to the latest Cretaceous. Tentative remains from Europe, Morocco, and Madagascar may extend their range into the Middle Jurassic. Here we report the first unambiguous Middle Jurassic (late Bajocian-Bathonian) atoposaurid: an anterior dentary from the Isle of Skye, Scotland, UK. A comprehensive review of atoposaurid specimens demonstrates that this dentary can be referred to T heriosuchus based on several derived characters, and differs from the five previously recognized species within this genus. Despite several diagnostic features, we conservatively refer it to T heriosuchus sp., pending the discovery of more complete material. As the oldest known definitively diagnostic atoposaurid, this discovery indicates that the oldest members of this group were small-bodied, had heterodont dentition, and were most likely widespread components of European faunas. Our review of mandibular and dental features in atoposaurids not only allows us to present a revised diagnosis of T heriosuchus, but also reveals a great amount of variability within this genus, and indicates that there are currently five valid species that can be differentiated by unique combinations of dental characteristics. This variability can be included in future broad-scale cladistics analyses of atoposaurids and closely related crocodyliforms, which promise to help untangle the complicated taxonomy and evolutionary history of Atoposauridae. © 2015 The Authors. Zoological Journal of the Linnean Society published by John Wiley &amp;amp;amp; Sons Ltd on behalf of The Linnean Society of London.
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... New vertebrate material from the Otter Sandstone Formation of Devon includes a cleithrolepidid fish, temnospondyl amphibians, a procolophonid reptile, and a tooth which may be attributed to a tanystropheid reptile. ... 7в) which is... more
... New vertebrate material from the Otter Sandstone Formation of Devon includes a cleithrolepidid fish, temnospondyl amphibians, a procolophonid reptile, and a tooth which may be attributed to a tanystropheid reptile. ... 7в) which is characteristically procolophonid. ...