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Diverse vertebrate assemblage of the Kilmaluag Formation (Bathonian, Middle Jurassic) of Skye, Scotland

Published online by Cambridge University Press:  27 July 2020

Elsa PANCIROLI*
Affiliation:
Department of Earth Sciences, University of Oxford, South Parks Road, OxfordOX1 3AN, UK National Museum of Scotland, Chambers St, EdinburghEH1 1JF, UK School of Geosciences, University of Edinburgh, Grant Institute, Kings Buildings, EdinburghEH9 3FE, UK
Roger B. J. BENSON
Affiliation:
Department of Earth Sciences, University of Oxford, South Parks Road, OxfordOX1 3AN, UK
Stig WALSH
Affiliation:
National Museum of Scotland, Chambers St, EdinburghEH1 1JF, UK School of Geosciences, University of Edinburgh, Grant Institute, Kings Buildings, EdinburghEH9 3FE, UK
Richard J. BUTLER
Affiliation:
School of Geography, Earth and Environmental Sciences, University of Birmingham, BirminghamB15 2TT, UK
Tiago Andrade CASTRO
Affiliation:
School of Geosciences, University of Edinburgh, Grant Institute, Kings Buildings, EdinburghEH9 3FE, UK
Marc E. H. JONES
Affiliation:
Department of Cell and Developmental Biology, University College London, LondonWC1E 6BT, UK
Susan E. EVANS
Affiliation:
Department of Cell and Developmental Biology, University College London, LondonWC1E 6BT, UK
*
*Corresponding author. Email: elsa.panciroli@earth.ox.ac.uk

Abstract

The Kilmaluag Formation on the Isle of Skye, Scotland, provides one of the richest Mesozoic vertebrate fossil assemblages in the UK, and is among the richest globally for Middle Jurassic tetrapods. Since its discovery in 1971, this assemblage has predominantly yielded small-bodied tetrapods, including salamanders, choristoderes, lepidosaurs, turtles, crocodylomorphs, pterosaurs, dinosaurs, non-mammalian cynodonts and mammals, alongside abundant fish and invertebrates. It is protected as a Site of Special Scientific Interest and by Nature Conservancy Order. Unlike contemporaneous localities from England, this assemblage yields associated partial skeletons, providing unprecedented new data. We present a comprehensive updated overview of the Kilmaluag Formation, including its geology and the fossil collections made to date, with evidence of several species occurrences presented here for the first time. We place the vertebrate faunal assemblage in an international context through comparisons with relevant contemporaneous localities from the UK, Europe, Africa, Asia and the US. This wealth of material reveals the Kilmaluag Formation as a vertebrate fossil assemblage of global significance, both in terms of understanding Middle Jurassic faunal composition and the completeness of specimens, with implications for the early evolutionary histories of mammals, squamates and amphibians.

Type
Review Article
Copyright
Copyright © The Author(s) 2020. Published by Cambridge University Press on behalf of The Royal Society of Edinburgh

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References

9. References

Anderson, F. W. & Dunham, K. C. 1966. The geology of northern Skye. Memoir of the Geological Survey of Great Britain. Sheet 80 and parts of sheets 81, 90 and 91 (Scotland). Her Majesty's Stationery Office, Edinburgh, Scotland, UK.Google Scholar
Andrews, J. E. 1985. The sedimentary facies of a late Bathonian regressive episode: the Kilmaluag and Skudiburgh Formations of the Great Estuarine Group, Inner Hebrides, Scotland. Journal of the Geological Society of London 142, 1119–37.CrossRefGoogle Scholar
Anquetin, J. 2010. The anatomy of the basal turtle Eileanchelys Waldmani from the Middle Jurassic of the Isle of Skye, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101, 6796.CrossRefGoogle Scholar
Anquetin, J., Barrett, P. M., Jones, M. E. H., Moore-Fay, S. & Evans, S. E. 2009. A new stem turtle from the Middle Jurassic of Scotland: new insights into the evolution and palaeoecology of basal turtles. Proceedings of the Royal Society B 276, 879–86.CrossRefGoogle ScholarPubMed
Anquetin, J. & Claude, J. 2008. Reassessment of the oldest British turtle: Protochelys from the Middle Jurassic Stonesfield slate of Stonesfield, Oxfordshire, UK. Geodiversitas 30, 331–44.Google Scholar
Averianov, A. O., Lopatin, A. V., Skutschas, P. P., Martynovich, N. V., Leshchinskiy, S. V., Krasnolutskii, S. A. & Fayngertz, A. V. 2005. Discovery of Middle Jurassic mammals from Siberia. Acta Palaeontologica Polonica 50, 789–97.Google Scholar
Averianov, A. O., Lopatin, A. V. & Krasnolutskii, S. A. 2011. The first haramiyid (Mammalia, Allotheria) from the Jurassic of Russia. Doklady Biological Sciences 437, 103–06.CrossRefGoogle ScholarPubMed
Averianov, A. O., Martin, T., Skutschas, P. P., Danilov, I. G., Schultz, J., Schellhorn, R., Obraztsova, E., Lopatin, A., Sytchevskaya, E., Kuzmin, I. & Krasnolutskii, S. 2016. Middle Jurassic vertebrate assemblage of Berezovsk coal mine in western Siberia (Russia). Global Geology 19, 187204.Google Scholar
Averianov, A. O., Martin, T., Lopatin, A. V., Schultz, J. A., Schellhorn, R., Krasnolutskii, S., Skutschas, P. & Ivantsov, S. 2019. Haramiyidan mammals from the Middle Jurassic of Western Siberia, Russia. Part 1: Shenshouidae and Maiopatagium. Journal of Vertebrate Paleontology 39, e1669159.CrossRefGoogle Scholar
Averianov, A. O., Martin, T., Lopatin, A. V., Schultz, J. A., Schellhorn, R., Krasnolutski, S., Skutschas, P. & Ivantsov, S. 2020. Multituberculate mammals from the Middle Jurassic of western Siberia, Russia and the origin of multituberculata. Papers in Palaeontology 119.Google Scholar
Barrett, P. M. 2006. A sauropod dinosaur tooth from the Middle Jurassic of Skye, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 97, 2529.CrossRefGoogle Scholar
Barron, A. J. M., Lott, G. K. & Riding, J. B. 2012. Stratigraphic framework for the Middle Jurassic strata of Great Britain and the adjoining continental shelf: research report RR/11/06. British Geological Survey, Keyworth. 177 pp.Google Scholar
Benson, R. B. 2018. Dinosaur macroevolution and macroecology. Annual Review of Ecology, Evolution, and Systematics 49, 379408.CrossRefGoogle Scholar
Benson, R. B. J., Campione, N. E., Carrano, M. T., Mannion, P. D., Sullivan, C., Upchurch, P. & Evans, D. C. 2014. Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian lineage. PLoS Biology 12, e1001853.CrossRefGoogle Scholar
Blake, C. C. 1863. On chelonian scutes from the Stonesfield slate. The Geologist 6, 183–84.CrossRefGoogle Scholar
Buckland, W. 1824. Notice on the Megalosaurus or great fossil lizard of Stonesfield. Transactions of the Geological Society of London 2, 390–96.CrossRefGoogle Scholar
Burbrink, F. T., Grazziotin, F. G., Pyron, R. A., Cundall, D., Donnellan, S., Irish, F., Keogh, J. S., Kraus, F., Murphy, R. W., Noonan, B. & Raxworthy, C. J. 2019. Interrogating genomic-scale data for Squamata (lizards, snakes, and amphisbaenians) shows no support for key traditional morphological relationships. Systematic Biology 69, 119.Google Scholar
Butler, P. M. 1939. The teeth of the Jurassic mammals. Proceedings of the Zoological Society of London 109, 329–56.Google Scholar
Butler, P. M., Sigogneau-Russell, D. & Ensom, P. C. 2012. Possible persistence of the morganucodontans in the Lower Cretaceous Purbeck Limestone Group (Dorset, England). Cretaceous Research 33, 135–45.CrossRefGoogle Scholar
Butler, P. M. & Hooker, J. J. 2005. New teeth of allotherian mammals from the English Bathonian, including the earliest multituberculates. Acta Palaeontologica Polonica 50, 185207.Google Scholar
Butler, P. M. & Sigogneau-Russell, D. 2016. Diversity of triconodonts in the Middle Jurassic of Great Britain. Palaeontologia Polonica 67, 3565.Google Scholar
Caldwell, M. W., Nydam, R. L., Palci, A. & Apesteguía, S. 2015. The oldest known snakes from the Middle Jurassic-lower cretaceous provide insights on snake evolution. Nature Communications 6, 111.CrossRefGoogle ScholarPubMed
Chen, P.-J. & Hudson, J. D. 1991. The conchostracan fauna of the Great Estuarine Group, Middle Jurassic, Scotland. Palaeontology 34, 515–45.Google Scholar
Chure, D. J., Litwin, R., Hasiotis, S. T., Evanoff, E. & Carpenter, K. 2006. The fauna and flora of the Morrison Formation. New Mexico Museum of Natural History and Science Bulletin 36, 233–49.Google Scholar
Clark, N. D. L. 2018. Review of the dinosaur remains from the Middle Jurassic of Scotland, UK. Geosciences 8, 53.CrossRefGoogle Scholar
Clark, N. D. L., Ross, D. A. & Booth, P. 2005. Dinosaur tracks from the Kilmaluag Formation (Bathonian, Middle Jurassic) of Score Bay, Isle of Skye, Scotland, UK. Ichnos 12, 93104.CrossRefGoogle Scholar
Close, R. A., Davis, B. M., Walsh, S., Woloniewicz, A. S., Friedman, M. & Benson, R. B. J. 2016. A lower jaw of Palaeoxonodon from the Middle Jurassic of the Isle of Skye, Scotland, sheds new light on the diversity of British stem therians. Palaeontology 59, 155–69.CrossRefGoogle Scholar
Cohen, K. M., Harper, D. A. T., Gibbard, P. L. & Fan, J.-X. 2019. International chronostratigraphic chart. International Commission of Stratigraphy. https://stratigraphy.org/chart.Google Scholar
Cope, J. C. W., Dufl, K. L., Parsons, C. F., Torrens, H. S., Wimbledon, W. A. & Wright, J. 1980. A correlation of Jurassic rocks in the British Isles. Pt. 2: Middle and Upper Jurassic. Geological Society Special Report 15. 109 pp.Google Scholar
Delair, J. B. & Sarjeant, W. A. 2002. The earliest discoveries of dinosaurs: the records re-examined. Proceedings of the Geologists’ Association 113, 185–97.CrossRefGoogle Scholar
Ensom, P. C. 2007. The Purbeck Limestone Group of Dorset, southern England. Geology Today 23, 178–85.CrossRefGoogle Scholar
Evans, S. E., Barrett, P., Hilton, J., Butler, R. J., Jones, M. E. H., Liang, M.-M., Parrish, J. C., Rayfield, E. J., Sigogneau-Russell, D. & Underwood, C. J. 2006. The Middle Jurassic vertebrate assemblage of Skye, Scotland. In Barrett, P. & Evans, S. (eds) Proceedings of the ninth symposium on Mesozoic terrestrial ecosystems and biota, 3639. London: Natural History Museum.Google Scholar
Evans, S. E. 1989. New material of Cteniogenys (Reptilia: Diapsida) and a reassessment of the systematic position of the genus. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 181, 577–89.CrossRefGoogle Scholar
Evans, S. E. 1990. The skull of Cteniogenys, a choristodere (Reptilia: Archosauromorpha) from the Middle Jurassic of Oxfordshire. Zoological Journal of the Linnean Society 99, 205–37.CrossRefGoogle Scholar
Evans, S. E. 1991a. A new lizard-like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of England. Zoological Journal of the Linnean Society 103, 391412.CrossRefGoogle Scholar
Evans, S. E. 1991b. The postcranial skeleton of the choristodere Cteniogenys (Reptilia: Diapsida) from the Middle Jurassic of England. Geobios 24, 187–99.CrossRefGoogle Scholar
Evans, S. E. 1992. A sphenodontian (Reptilia: Lepidosauria) from the Middle Jurassic of England. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 1992, 449–57.CrossRefGoogle Scholar
Evans, S. E. 1994. A new anguimorph lizard from the Jurassic and Lower Cretaceous of England. Palaeontology 37, 3349.Google Scholar
Evans, S. E. 1996. Parviraptor (Squamata: Anguimorpha) and other lizards from the Morrison Formation at Fruita, Colorado. Museum of Northern Arizona Bulletin 60, 243–48.Google Scholar
Evans, S. E. 1998. Crown group lizards (Reptilia, Squamata) from the middle Jurassic of the British Isles. Palaeontographica Abteilung A-Stuttgart 250, 123–54.Google Scholar
Evans, S. E., Milner, A. R. & Mussett, F. 1988. The earliest known salamanders (Amphibia, Caudata): a record from the Middle Jurassic of England. Geobios 21, 539–52.CrossRefGoogle Scholar
Evans, S. E., Milner, A. R. & Mussett, F. 1990. A discoglossid frog from the Middle Jurassic of England. Palaeontology 33, 299311.Google Scholar
Evans, S. E. & Chure, D. C. 1998 Paramacellodid lizard skulls from the Jurassic Morrison Formation at Dinosaur national monument, Utah. Journal of Vertebrate Paleontology 18, 99114CrossRefGoogle Scholar
Evans, S. E. & Milner, A. R. 1994. Middle Jurassic microvertebrate assemblages from the British Isles. In Fraser, N. C. and Sues, H.-D. (eds) In the shadow of the dinosaurs: early Mesozoic Tetrapods, 303–21. Cambridge: Cambridge University Press.Google Scholar
Evans, S. E. & Waldman, M. 1996. Small reptiles and amphibians from the Middle Jurassic of Skye, Scotland. The continental Jurassic. Museum of Northern Arizona, Bulletin 60, 219–26.Google Scholar
Flynn, J. J., Fox, S. R., Parrish, J. M., Ranivoharimanana, L. & Wyss, A. R. 2006. Assessing diversity and paleoecology of a Middle Jurassic microvertebrate assemblage from Madagascar. New Mexico Museum of Natural History and Science Bulletin 37, 476489.Google Scholar
Foster, J. 2003a. New specimens of Eilenodon (Reptilia, Sphenodontia) from the Morrison Formation (Upper Jurassic) of Colorado and Utah. Brigham Young University Geological Studies 47, 1722.Google Scholar
Foster, J. R. 2003b. Paleoecological analysis of the vertebrate fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain region, U.S.A. New Mexico Museum of Natural History & Science Bulletin 23, 195.Google Scholar
Foster, J. R. & Heckert, A. B. 2011. Ichthyoliths and other microvertebrate remains from the Morrison Formation (Upper Jurassic) of northeastern Wyoming: a screen-washed sample indicates a significant aquatic component to the fauna. Palaeogeography, Palaeoclimatology, Palaeoecology 305, 264–79.CrossRefGoogle Scholar
Foster, J. R. & Lucas, S. G. 2006. Paleontology and geology of the Upper Jurassic Morrison Formation: bulletin 36. New Mexico Museum of Natural History and Science, 249 pp.Google Scholar
Freeman, E. F. 1976. Mammal teeth from the Forest Marble (Middle Jurassic) of Oxfordshire, England. Science (New York, N.Y.) 194, 1053–55.CrossRefGoogle ScholarPubMed
Freeman, E. F. 1979. A Middle Jurassic mammal bed from Oxfordshire. Palaeontology 22, 135–66.Google Scholar
Gao, K.-Q. & Shubin, N. H. 2012. Late Jurassic salamandroid from western Liaoning, China. Proceedings of the National Academy of Sciences 109, 5767–72.CrossRefGoogle ScholarPubMed
Gardner, J., Evans, S. E. & Sigogneau-Russell, D. 2003. Albanerpetontid amphibians from the early cretaceous of Morocco and the Middle Jurassic of England. Acta Palaeontologia Polonica 48, 301–19.Google Scholar
Gilmore, C. W. 1909. A new rhynchocephalian reptile from the Jurassic of Wyoming, with notes on the fauna of “Quarry 9”. Proceedings of the United States National Museum 37, 3542.CrossRefGoogle Scholar
Gilmore, C. W. 1928 Fossil lizards of North America. Memoirs of the National Academy of Science 22, 1169.Google Scholar
Gillham, C. 1994. A fossil turtle (Reptilia: Chelonia) from the Middle Jurassic of Oxfordshire, England. Neues Jahrbuch der Geologie und Paläontologie. Monatshefte 10, 581–96.CrossRefGoogle Scholar
Haddoumi, H., Aiméras, Y., Bodergat, A. M., Charrière, A., Mangold, C. & Benshili, K. 1998. Âges et environnements des Couches rouges d'Anoual (Jurassique moyen et Crétacé inférieur, Haut-Atlas oriental, Maroc). Comptes Rendus de l'Académie des Sciences-Series IIA-Earth and Planetary Science 327, 127–33.Google Scholar
Haddoumi, H., Allain, R., Meslouh, S., Metais, G., Monbaron, M., Pons, D., Rage, J. C., Vullo, R., Zouhri, S. & Gheerbrant, E. 2016. Guelb el Ahmar (Bathonian, Anoual Syncline, eastern Morocco): first continental flora and fauna including mammals from the Middle Jurassic of Africa. Gondwana Research 29, 290319.CrossRefGoogle Scholar
Hahn, G., Sigogneau- Russell, D. & Godefroit, P. 1991. New data on Brachyzostrodon (Mammalia; Upper Triassic). Geologica et Paleontologica 25, 237–49.Google Scholar
Harris, J. P. & Hudson, J. D. 1980. Lithostratigraphy of the Great Estuarine Group (Middle Jurassic), Inner Hebrides. Scottish Journal of Geology 16, 231–50.CrossRefGoogle Scholar
Helmdach, F. F. 1971. Stratigraphy and ostracode-fauna from the coal mine Guimarota (Upper Jurassic). Memórias dos Serviços Geológicos de Portugal, N.S. 17, 4388.Google Scholar
Hesselbo, S. P. & Coe, A. L. 2000. Jurassic sequences of the Hebrides Basin, Isle of Skye Scotland. In Graham, J. R. and Ryan, A. (eds) Field trip guidebook, 4158. Dublin: International Sedimentologists Association Meeting.Google Scholar
Ivakhnenko, M. 1978. Urodeles from the Triassic and Jurassic of Soviet Central Asia. Palaeontologicheski Zhurnal 8489. (in Russian) and Paleontology Journal 12, 362–368 (in English) [In Russian.]Google Scholar
Ji, Q., Luo, Z.-X., Yuan, C.-X. & Tabrum, A. R. 2006. A swimming mammaliaform from the Middle Jurassic and ecomorphological diversification of early mammals. Science (New York, N.Y.) 311, 1123–27.CrossRefGoogle ScholarPubMed
Jones, M. E., Anderson, C. L., Hipsley, C. A., Müller, J., Evans, S. E. & Schoch, R. R. 2013. Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards, snakes, and tuatara). BMC Evolutionary Biology 13, 208.CrossRefGoogle Scholar
Jones, M. E., Lucas, P. W., Tucker, A. S., Watson, A. P., Sertich, J. J., Foster, J. R., Williams, R., Garbe, U., Bevitt, J. J. & Salvemini, F. 2018a. Neutron scanning reveals unexpected complexity in the enamel thickness of an herbivorous Jurassic reptile. Journal of The Royal Society Interface 15, 20180039.CrossRefGoogle Scholar
Jones, M. E. H., Hill, L. E., Benson, R. B. & Evans, S. E. 2019. Three dimensional skeletons of middle Jurassic stem-group salamanders from Scotland, UK. Journal of Vertebrate Paleontology, Program and Abstracts 2018, 126.Google Scholar
Judd, J. W. 1878. The secondary rocks of Scotland. Third paper. The strata of the western coasts and islands. Quarterly Journal of the Geological Society of London 34, 660743.CrossRefGoogle Scholar
Kermack, K. A., Lee, A. J., Lees, P. M. & Mussett, F. 1987. A new docodont from the Forest Marble. Zoological Journal of the Linnean Society 89, 139.CrossRefGoogle Scholar
Kermack, K. A., Kermack, D. M., Lees, P. M. & Mills, J. R. 1998. New multituberculate-like teeth from the Middle Jurassic of England. Acta Palaeontologica Polonica 43, 581606.Google Scholar
Kielan-Jaworowska, Z. & Ensom, P. C. 1992. Multituberculate mammals from the Upper Jurassic Purbeck limestone formation of southern England. Palaeontology 35, 95126.Google Scholar
Kowallis, B. J., Christiansen, E. H., Deino, A. L., Peterson, F., Turner, C. E., Kunk, M. J. & Obradovich, J. D. 1998. The age of the Morrison Formation. Modern Geology 22, 235–60.Google Scholar
Kühne, W. G. 1956. The Liassic therapsid Oligokyphus. London: British Museum (Natural History). 149 pp.Google Scholar
Lee, M. S., Cau, A., Naish, D. & Dyke, G. J. 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science (New York, N.Y.) 345, 562–66.CrossRefGoogle Scholar
Luo, Z.-X. 2007. Transformation and diversification in early mammal evolution. Nature 450, 1011–19.CrossRefGoogle ScholarPubMed
Luo, Z.-X., Ji, Q. & Yuan, C.-X. 2007. Convergent dental adaptations in pseudo-tribosphenic and tribosphenic mammals. Nature Letters 450, 9397.CrossRefGoogle ScholarPubMed
Luo, Z.-X., Meng, Q.-J., Ji, Q., Liu, D., Zhang, Y.-G. & Neander, A. I. 2015. Evolutionary development in basal mammaliaforms as revealed by a docodontan. Science (New York, N.Y.) 347, 760–64.CrossRefGoogle ScholarPubMed
Luo, Z.-X. & Martin, T. 2007. Analysis of molar structure and phylogeny of docodontan genera. Bulletin of Carnegie Museum of Natural History 39, 2747.CrossRefGoogle Scholar
Maidment, S. C. & Muxworthy, A. 2019. A chronostratigraphic framework for the Upper Jurassic Morrison Formation, western USA. Journal of Sedimentary Research 89, 1017–38.CrossRefGoogle Scholar
Marjanović, D. & Laurin, M. 2014. An updated paleontological timetree of lissamphibians, with comments on the anatomy of Jurassic crown-group salamanders (Urodela). Historical Biology 26, 535–50.CrossRefGoogle Scholar
Marsh, O. C. 1880. Notice on Jurassic mammals representing two new orders. American Journal of Science 20, 235–39.CrossRefGoogle Scholar
Marsh, O. C. 1887 American Jurassic mammals. American Journal of Science 33, 326–48.Google Scholar
Marshall, P. 2003. Ichnofossils of the Psilonichnus ichnofacies and their paleoecological and paleoenvironmental significance in the Scottish Middle Jurassic. Ichnos 9, 95108.CrossRefGoogle Scholar
Martin, T. 2001. Mammalian fauna of the Late Jurassic Guimarota ecosystem. Publicación Electrónica de la Asociación Paleontológica Argentina 7, 1.Google Scholar
Martin, T. 2005. Postcranial anatomy of Haldanodon exspectatus (Mammalia, Docodonta) from the Late Jurassic (Kimmeridgian) of Portugal and its bearing for mammalian evolution. Zoological Journal of the Linnean Society 145, 219–48.CrossRefGoogle Scholar
Martin, T. & Krebs, B. 2000. Guimarota. A jurassic ecosystem. Munich, Germany: Verlag.Google Scholar
Martin-Silverstone, E. G., Unwin, D. M. & Barrett, P. M. 2018. A new three-dimensionally preserved monofenestratan pterosaur from the Middle Jurassic of Scotland and the complex evolutionary history of the scapula-vertebral articulation. Symposium of Vertebrate Paleontology Meeting 2019 Programme and Abstracts, Brisbane, Australia, p. 150.Google Scholar
Mateus, O. 2006. Late Jurassic dinosaurs from the Morrison Formation, the Lourinhã and Alcobaça Formations (Portugal), and the Tendaguru Beds (Tanzania): a comparison. In Foster, J. R. and Lucas, S. G. R. M. (eds) Paleontology and geology of the upper Jurassic Morrison Formation, 223–31. New Mexico Museum of Natural History & Science: Bulletin 36.Google Scholar
Mateus, O., Dinis, J. & Cunha, P. P. 2017. The Lourinhã Formation: the Upper Jurassic to lower most Cretaceous of the Lusitanian Basin, Portugal–landscapes where dinosaurs walked. Ciências da Terra/Earth Sciences Journal 19, 7597.CrossRefGoogle Scholar
McGowan, G. J. 1996. Albanerpetontid amphibians from the Jurassic (Bathonian) of Southern England. Museum of Northern Arizona Bulletin 60, 227–34.Google Scholar
Mellere, D. & Steel, R. J. 1996. Tidal sedimentation in Inner Hebrides half grabens, Scotland: the mid-Jurassic Bearreraig sandstone formation. In DeBatist, M. & Jacobs, P. (eds) Geology of siliciclastic shelf seas, 4979. London: Geological Society, Special Publications, 117.Google Scholar
Meng, Q.-J., Grossnickle, D. M., Liu, D., Zhang, Y. G., Neander, A. I., Ji, Q. & Luo, Z. X. 2017. New gliding mammaliaforms from the Jurassic. Nature 548, 291–96.CrossRefGoogle ScholarPubMed
Morton, N. 1987. Jurassic subsidence history in the Hebrides, NW Scotland. Marine and Petroleum Geology 4, 226–42.CrossRefGoogle Scholar
Morton, N. & Hudson, J. D. 1995. Field guide to the Jurassic of the Isles of Raasay and Skye, Inner Hebrides, NW Scotland. In Taylor, P. D. (ed.) Field geology of the British Jurassic, 209–80. London: Geological Society.Google Scholar
Nesov, L. A. 1988. Late Mesozoic amphibians and lizards of Soviet Middle Asia. Acta Zoologica Cracoviensia 31,475–86.Google Scholar
Nesov, L. A., Fedorov, P. V., Potapov, D. O. & Golovnyeva, L. S. 1996. The structure of the skulls of caudate amphibians collected from the Jurassic of Kirgizstan and the cretaceous of Uzbekistan. Vestnik Sankt-Petersburgskogo Universiteta, Seriya 7, Geologiya, Geografiya 1, 311. [In Russian.]Google Scholar
Panciroli, E., Benson, R. B. J. & Walsh, S. 2017a. The dentary of Wareolestes rex (Megazostrodontidae): a new specimen from Scotland and implications for morganucodontan tooth replacement. Papers in Palaeontology 3, 373–86.CrossRefGoogle Scholar
Panciroli, E., Walsh, S., Fraser, N., Brusatte, S. L. & Corfe, I. 2017b. A reassessment of the postcanine dentition and systematics of the tritylodontid Stereognathus (Cynodontia, Tritylodontidae, Mammaliamorpha), from the Middle Jurassic of the UK. Journal of Vertebrate Paleontology 37, 373–86.CrossRefGoogle Scholar
Panciroli, E., Benson, R. B. J. & Butler, R. J. 2018a. New partial dentaries of Palaeoxonodon Ooliticus (Mammalia, Amphitheriidae) from Scotland, and posterior dentary morphology in stem cladotherians. Acta Paleontologica Polonica 63, 197206.Google Scholar
Panciroli, E., Schultz, J. A. & Luo, Z.-X. 2018b. Morphology of the petrosal and stapes of Borealestes (Mammaliaformes, Docodonta) from the Middle Jurassic of Skye, Scotland. Papers in Palaeontology 5, 139–56.CrossRefGoogle Scholar
Panciroli, E., Benson, R. B. J. & Luo, Z.-X. 2019. The mandible and dentition of Borealestes Serendipitus (Docodonta) from the Middle Jurassic of Skye, Scotland. Journal of Vertebrate Paleontology 39, e1621884.CrossRefGoogle Scholar
Panciroli, E., Benson, R. B. J., Fernandez, V., Butler, R. J., Fraser, N. C., Luo, Z.-X. & Walsh, S. in press. New species of mammaliaform and the cranium of Borealestes (Mammaliformes: Docodonta) from the Middle Jurassic of the British Isles. Zoological Journal of the Linnean Society.Google Scholar
Panciroli, E. P., Benson, R. B. J. & Walsh, S. 2018c. The mammal-rich freshwater assemblage of the Middle Jurassic Kilmaluag Formation, Isle of Skye, Scotland. Abstracts of the 13th Symposium on Mesozoic Terrestrial Ecosystems, Bonn, Germany, 97.Google Scholar
Prasad, G. V. & Manhas, B. K. 2002. Triconodont mammals from the Jurassic Kota Formation of India. Geodiversitas 24, 445464.Google Scholar
Rasmussen, T. E. & Callison, G. 1981. A new herbivorous sphenodontid (Rhynchocephalia: Reptilia) from the Jurassic of Colorado. Journal of Paleontology 55, 1109–16.Google Scholar
Rauhut, O. W., Hübner, T. & Lanser, K. P. 2016. A new megalosaurid theropod dinosaur from the late Middle Jurassic (Callovian) of north-western Germany: implications for theropod evolution and faunal turnover in the Jurassic. Palaeontologia Electronica 19, 165.Google Scholar
Rees, J. & Underwood, C. J. 2005. Hybodont sharks from the Middle Jurassic of the Inner Hebrides, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 96, 351–63.CrossRefGoogle Scholar
Riding, J. B, Walton, W. & Shaw, D. 1991. Toarcian to Bathonian (Jurassic Palynology of the Inner Hebrides, northwest Scotland. Palynology 15, 115–79.CrossRefGoogle Scholar
Roelants, K., Gower, D. J., Wilkinson, M., Loader, S. P., Biju, S. D., Guillaume, K., Moriau, L. & Bossuyt, F. 2007. Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences, USA 104, 887–92.CrossRefGoogle ScholarPubMed
Rougier, G. W., Sheth, A. S., Carpenter, K., Appella-Guisafre, L. & Davis, B. M. 2015. A new species of Docodon (Mammaliaformes, Docodonta) from the Upper Jurassic Morrison Formation and a reassessment of selected craniodental characters in basal mammaliaforms. Journal of Mammalian Evolution 22, 116.CrossRefGoogle Scholar
Scheyer, T. M. & Anquetin, J. 2008. Bone histology of the Middle Jurassic turtle shell remains from Kirtlington, Oxfordshire, England. Lethaia 41, 8596.CrossRefGoogle Scholar
Schudack, M. E. 2000. Geological setting and dating of the Guimarota-beds. In Martin, T. & Krebs, B. (eds) Guimarota. A Jurassic ecosystem, 2126. Munich, Germany: Verlag.Google Scholar
Schultz, J. A., Bhullar, B. A. S. & Luo, Z.-X. 2017. Re-examination of the Jurassic mammaliaform Docodon Victor by computed tomography and occlusal functional analysis. Journal of Mammalian Evolution 26, 938.CrossRefGoogle Scholar
Seiffert, J. 1973. Contribuição para o conhecimento da Fauna da Mina de Lignito Guimarota, 3: Upper Jurassic lizards from Central Portugal. Servicox Geologicos de Portugal 22, 785.Google Scholar
Sigogneau-Russell, D. 1998. Discovery of a Late Jurassic Chinese mammal in the upper Bathonian of England. Comptes Rendus de l'Académie des Sciences-Series IIA-Earth and Planetary Science 327, 571–76.Google Scholar
Sigogneau-Russell, D. 2003a. Docodonts from the British Mesozoic. Acta Palaeontologica Polonica 48, 357–74.Google Scholar
Sigogneau-Russell, D. 2003b. Holotherian mammals from the Forest Marble (Middle Jurassic of England). Geodiversitas 25, 501–37.Google Scholar
Simpson, G. G. 1926. American terrestrial Rhynchocephalia. American Journal of Science 67, 1216.CrossRefGoogle Scholar
Simpson, G. G. 1928. A catalogue of the Mesozoic Mammalia in the Geological Department of the British Museum. London: British Museum (Natural History). 215 pp.Google Scholar
Skutschas, P. P. 2013. Mesozoic salamanders and albanerpetontids of Middle Asia, Kazakhstan, and Siberia. Palaeobiodiversity and Palaeoenvironments 93, 441–57.CrossRefGoogle Scholar
Skutschas, P. P. 2014a. Kiyatriton Leshchinskiyi Averianov et Voronkevich, 2001, a crown-group salamander from the lower Cretaceous of Western Siberia, Russia. Cretaceous Research 51, 8894.CrossRefGoogle Scholar
Skutschas, P. P. 2014b. A relict stem salamander: evidence from the early cretaceous of Siberia. Acta Palaeontologica Polonica 61,119–23.Google Scholar
Skutschas, P. P., Kolchanov, V. V., Averianov, A. O., Martin, T., Schellhorn, R., Kolosov, P. N. & Vitenko, D. D. 2018. A new relict stem salamander from the early cretaceous of Yakutia, Siberian Russia. Acta Palaeontologica Polonica 63, 519–25.CrossRefGoogle Scholar
Skutschas, P. P. & Krasnolutskii, S. A. 2011. A new genus and species of basal salamanders from the Middle Jurassic of Western Siberia, Russia. Proceedings of the Zoological Institute RAS 315, 167–75.Google Scholar
Skutschas, P. P. & Martin, T. 2011. Cranial anatomy of the stem salamander Kokartus honorarius (Amphibia: Caudata) from the Middle Jurassic of Kyrgyzstan. Zoological Journal of the Linnean Society 161, 816–38.CrossRefGoogle Scholar
Sullivan, C., Wang, Y., Hone, D. W. E., Wang, Y., Xu, X. & Shang, F. 2014. The vertebrates of the Jurassic Daohugou biota of Northeastern China. Journal of Vertebrate Paleontology 34, 243–80.CrossRefGoogle Scholar
Turner, C. E. & Peterson, F. 2004. Reconstruction of the Upper Jurassic Morrison Formation extinct ecosystem – a synthesis. Sedimentary Geology 167, 309–55.CrossRefGoogle Scholar
Wakefield, M. I. 1995. Ostracod biostratinomy at lagoonal shorelines: examples from the Great Estuarine Group, Middle Jurassic, Scotland. Proceedings of the Geologists’ Association 106, 211–18.CrossRefGoogle Scholar
Waldman, M. & Evans, S. E. 1994. Lepidosauromorph reptiles from the Middle Jurassic of Skye. Zoological Journal of the Linnean Society 112, 135–50.CrossRefGoogle Scholar
Waldman, M. & Savage, R. J. G. 1972. The first Jurassic mammal from Scotland. Journal of the Geological Society of London 128, 119–25.CrossRefGoogle Scholar
Ward, D. J. 1984. Collecting isolated microvertebrate fossils. Zoological Journal of the Linnean Society 82, 245–59.CrossRefGoogle Scholar
Weishampel, D. B., Dodson, P. & Osmolska, H. 2004. The Dinosauria. 2nd edn. Berkeley: University of California Press. 861 pp.CrossRefGoogle Scholar
Westhead, R. K. & Mather, A. E. 1996. An updated lithostratigraphy for the Purbeck Limestone Group in the Dorset type-area. Proceedings of the Geologists’ Association 107, 117–28.CrossRefGoogle Scholar
Whyte, S. & Ross, D. 2019. Jurassic Skye: dinosaurs and other fossils of the Isle of Skye. Berkshire, UK: NatureBureau. 62 pp.Google Scholar
Wills, S., Barrett, P. M. & Walker, A. 2014. New dinosaur and crocodylomorph material from the Middle Jurassic (Bathonian) Kilmaluag Formation, Skye, Scotland. Scottish Journal of Geology 50, 183–90.CrossRefGoogle Scholar
Wills, S., Bernard, E. L., Brewer, P., Underwood, C. J. & Ward, D. J. 2019. Palaeontology, stratigraphy and sedimentology of Woodeaton Quarry (Oxfordshire) and a new microvertebrate site from the White Limestone Formation (Bathonian, Jurassic). Proceedings of the Geologists’ Association 130, 170–86.CrossRefGoogle Scholar
Xu, X., Zhou, Z. H., Sullivan, C., Wang, Y. & Ren, D. 2016. An updated review of the Middle-Jurassic Yanliao Biota: chronology, taphonomy, paleontology, and paleoecology. Acta Geologica Sinica 90, 1801–40.CrossRefGoogle Scholar
Xu, X., Zhou, Z., Sullivan, C. & Wang, Y. 2017. The Yanliao Biota: a trove of exceptionally preserved Middle-Late Jurassic terrestrial life forms. In Fraser, N. C. & Sues, H.-D. (eds) Terrestrial conservation Lagerstätten, 131–67. London: Dunedin Academic Press.Google Scholar
Zhou, C. F., Wu, S., Martin, T. & Luo, Z.-X. 2013. A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations. Nature 500, 163–67.CrossRefGoogle ScholarPubMed
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