Skip to main content Accessibility help

Diverse and durophagous: Early Carboniferous chondrichthyans from the Scottish Borders

  • Kelly R. Richards (a1), Janet E. Sherwin (a2), Timothy R. Smithson (a1), Rebecca F. Bennion (a1), Sarah J. Davies (a2), John E. A. Marshall (a3) and Jennifer A. Clack (a1)...


Chondrichthyan teeth from a new locality in the Scottish Borders supply additional evidence of Early Carboniferous chondrichthyans in the UK. The interbedded dolostones and siltstones of the Ballagan Formation exposed along Whitrope Burn are interpreted as representing a restricted lagoonal environment that received significant amounts of land-derived sediment. This site is palynologically dated to the latest Tournaisian–early Viséan. The diverse dental fauna documented here is dominated by large crushing holocephalan toothplates, with very few, small non-crushing chondrichthyan teeth. Two new taxa are named and described. Our samples are consistent with worldwide evidence that chondrichthyan crushing faunas are common following the Hangenberg extinction event. The lagoonal habitat represented by Whitrope Burn may represent a temporary refugium that was host to a near-relict fauna dominated by large holocephalan chondrichthyans with crushing dentitions. Many of these had already become scarce in other localities by the Viséan and become extinct later in the Carboniferous. This fauna provides evidence of early endemism or niche separation within European chondrichthyan faunas at this time. This evidence points to a complex picture in which the diversity of durophagous chondrichthyans is controlled by narrow spatial shifts in niche availability over time.


Corresponding author

*Current address: University of Oxford Museum of Natural History, Parks Road, Oxford, OX3 1PW, UK.
**Current addresses: (1) Geology Research Unit, Université de Liège, 14 Allée du 6 Août, 4000 Liège, Belgium. (2) Directorate of Earth and History of Life, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, Belgium.


Hide All
Agassiz, L. 1838. Recherches sur les Poisons Fossiles. Volume 3. Neuchâtel, Switzerland: Petitpierre.
Agassiz, L. 1843. Recherches sur les Poisons Fossiles. Volume 5. Neuchâtel, Switzerland: Petitpierre.
Behan, C., Walkden, G. & Cuny, G. 2012. A Carboniferous chondrichthyan assemblage from residues within a Triassic Karst system at Cromhall Quarry. Palaeontology 55, 12451263.
Bennett, C. E., Siveter, D. J., Davies, S. J., Williams, M., Wilkinson, I. P., Browne, M. & Miller, C. G. 2012. Ostracods from freshwater and brackish environments of the Carboniferous of the Midland Valley of Scotland: the early colonization of terrestrial water bodies. Geological Magazine 149, 366396.
Berg, L. S. 1940. Sistema ryboobraznykh i ryb, nyne zhivushchikh i iskopaemykh. [Classification of fishes, both recent and fossil]. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute, USSR Academy of Sciences] 5, 1–517.
Bonaparte, C. L. 1831. Saggio di una distribuzione metodica degli animali vertebrati. Giornale Arcadico di Scienze 49, 177.
Bonaparte, C. L. 1838. Selachorum tabula analytica. Nuovi Annali delle Scienze Naturali (Bologna) 1, 195214.
Brandon, A., Riley, N. J., Wilson, A. A. & Ellison, R. A. 1995. Three new early Namurian (E1c–E2a) marine bands in central and northern England, UK, and their bearing on correlations with the Askrigg Block. Proceedings of the Yorkshire Geological Society 50, 333335.
Bromley, R. G. 1996. Trace fossil: biology, taphonomy and applications. London: Chapman & Hall. 378 pp.
Capetta, H., Duffin, C. J. & Zidek, J. 1993. Chondrichthyes. In Benton, M. J. (ed.) The Fossil Record, Vol 2, 593609. London: Chapman & Hall. xviii+846 pp.
Compagno, L. J. V. 1977. Phyletic Relationships of Living Sharks and Rays. American Zoologist. 17, 303322.
Davis, J. W. 1883. On the fossil fishes of the Carboniferous limestone series of Great Britain. Scientific Transactions of the Royal Dublin Society 1, 327600.
Dean, B. 1909. Studies on fossil fishes (sharks, chimaeroids, and arthrodires). Memoirs of the American Museum of Natural History 9, 211287.
Duffin, C. J. & Ginter, M. 2006. Comments on the Selachian Genus “Cladodus” Agassiz, 1843. Journal of Vertebrate Paleontology 26, 253266.
Duffin, C. J. & Ward, D. 1983. Neoselachian sharks' teeth from the Lower Carboniferous of Britain and the Lower Permian of the USA. Palaeontology 26, 93110.
Duncan, M. 2003. Early Carboniferous chondrichthyan Thrinacodus from Ireland, and a reconstruction of jaw apparatus. Acta Palaeontological Polonica 48, 113122.
Duncan, M. 2004. Chondrichthyan genus Lissodus from the Lower Carboniferous of Ireland. Acta Palaeontologica Polonica 49, 417428.
Duncan, M. 2006. Various chondrichthyan microfossil faunas from the Lower Mississippian (Carboniferous) of Ireland. Irish Journal of Earth Sciences 24, 5169.
Edwards, W. & Stubblefield, C. J. 1947. Marine bands and other faunal marker-horizons in relation to the sedimentary cycles of the middle coal measures of Nottinghamshire and Derbyshire. Quarterly Journal of the Geological Society 103, 209256.
Finarelli, J. A. & Coates, M. I. 2014. Chondrenchelys problematica (Traquair, 1888) redescribed: a Lower Carboniferous, eel-like holocephalan from Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 105, 3559.
Garvey, J. & Turner, S. 2006. Vertebrate remains from the presumed earliest Carboniferous of the Mansfield Basin, Victoria. Alcheringa 30, 4362.
Ginter, M. 1999. Famennian-Tournaisian chondrichthyan microremains from the eastern Thuringian Slate Mountains. Abhandlungen und Berichte für Naturkunde, Magdeburg 21, 2547.
Ginter, M., Hairapetian, V. & Klug, S. 2002. Famennian chondrichthyans from the shelves of North Gondwana. Acta Geologica Polonica 52, 169215.
Ginter, M., Hampe, O. & Duffin, C. J. 2010. Chondrichthyes. Paleozoic Elasmobranchii: teeth. In Schultze, H-P. (ed.) Handbook of Paleoichthyology, Volume 3D. München: Verlag Dr. Friedrich Pfeil. 512 pp.
Ginter, M., Duffin, C. J., Dean, M. T. & Korn, D. 2015. Late Viséan pelagic chondrichthyans from northern Europe. Acta Palaeontologica Polonica 60, 899922.
Ginter, M. & Sun, Y. 2007. Chondrichthyan remains from the Lower Carboniferous of Muhua, southern China. Acta Palaeontologica Polonica 52, 705727.
Glikman, L. S. 1964. Akuly paleogena I ih stratigrafičeskoe značenie [Sharks of the Paleogene and their stratigraphic significance]. Moscow: Doklady Akademii Nauk SSSR [Academy of Sciences of the USSR].
Gunnell, F. H. 1933. Conodonts and fish remains from the Cherokee, Kansas City, and Wabaunsee Groups of Missouri and Kansas. Journal of Paleontology 7, 261297.
Habibi, T. & Ginter, M. 2011. Early Carboniferous chondrichthyans from the Mobarak Formation, central Alborz Mountains, Iran. Acta Geologica Polonica 61, 2734.
Hairapetian, V. & Ginter, M. 2009. Fammenian chondrichthyan remains from the Chariseh section, Central Iran. Acta Geologica Polonica 59, 173200.
Hay, O. P. 1899. On some changes in the names, generic and specific, of certain fossil fishes. American Naturalist 33, 783792.
Hay, O. P. 1902. Bibliography and catalogue of the fossil vertebrata of North America. Bulletin of the United States Geological Survey 179, 1–868.
Huxley, T. H. 1880. On the application of the laws of evolution to the arrangement of the Vertebrata, and more particularly of the Mammalia. Proceedings of the Zoological Society of London 43, 649662.
Ivanov, A. O. 1996. The Early Carboniferous chondrichthyans of the South Urals, Russia. Geological Society, London, Special Publications 107, 417425.
Ivanov, A. 2000. Permian elasmobranchs (Chondrichthyes) of Russia. Ichthyolith Issues, Special Publication 6, 3942.
Ivanov, A. 2005. Early Permian Chondrichthyans of the Middle and South Urals. Revista Brasileira de Paleontologia 8, 127138.
Jaekel, O. 1925. Das Mundskelett der Wirbeltiere. Gegenbaurs Morphologisches Jahrbuch 55, 402409.
Jerve, A., Johanson, Z., Ahlberg, P. & Boisvert, C. 2014. Embryonic development of fin spine in Callorhincus milii (Holocephali); implications for chondrichthyan fin spine evolution. Evolution & Development 16, 339353.
Kriwet, J., Kiessling, W. & Klug, S. 2009. Diversification trajectories and evolution life-history traits in early sharks and batoids. Proceedings of the Royal Society, London, Series B 276, 945951.
Lebedev, O. A. 1996. Fish assemblages in the Tournaisian–Viséan environments of the East European Platform. Geological Society, London, Special Publications 107, 387415.
Lund, R. 1982. Harpagofututor volsellorhinus New genus and species (Chondrichthyes, Chondrenchelyiformes) from the Namurian Bear Gulch Limestone, Chondrenchelys problematica Traquair (Viséan), and their sexual dimorphism. Journal of Paleontology 56, 938958.
Lund, R. & Grogan, E. D. 1997. Relationships of the Chimaeriformes and the basal radiation of the Chondrichthyes. Reviews in Fish Biology and Fisheries 7, 65123.
Maisey, J. G. 1975. The interrelationships of the phalacanthous selachians. Neues Jahrbuch für Geologie und Paläontologie 9, 553567.
Maisey, J. G. 1977. The fossil selachian fishes Palaeospinax Egerton, 1872 and Nemacanthus Agassiz, 1837. Zoological Journal of the Linnean Society 60, 259273.
Maisey, J. G. 1978. Growth and form of finspines in hybodont sharks. Palaeontology 21, 657666.
Manski, C. F. & Lucas, S. G. 2013. Romer's Gap Revisited: Continental Assemblages and Ichno-assemblages from the basal Carboniferous of Blue Beach, Nova Scotia, Canada. The Carboniferous–Permian Transition. Bulletin of the New Mexico Museum of Natural History and Science 60, 244273.
Mapes, R. H. & Hansen, M. C. 1984. Pennsylvanian shark-cephalopod predation: a case study. Lethaia 17, 175183.
McCoy, F. A. 1855. A Systematic description of the British Palaeozoic fossils in the Geological Museum of the University of Cambridge. 1–661.
McKenzie, M. A. & Bamber, E. W. 1979. An occurrence of Lower Carboniferous fish remains from Alberta, Canada. Canadian Journal of Earth Sciences 16, 16281631.
Morris, J. & Roberts, G. E. 1862. On the Carboniferous limestone of Oreton and Farlow, Clee Hills, Shropshire. Quarterly Journal of the Geological Society of London 18, 94106.
Newberry, J. S. & Worthen, A. H. 1866. Descriptions of new species of vertebrates, mainly from the Sub-Carboniferous Limestone and Coal Measures of Illinois. Geological Survey of Illinois 2, 9–134.
Newberry, J. S. & Worthen, A. H. 1870. Geology and Paleontology. Descriptions of fossil vertebrates. Geological Survey of Illinois 4, 343374.
Obruchev, D. V. 1953. Lzuchenie edestid y raboty A. P. Karpinskogo [Studies on edestids and the works of A. P. Karpinski]. Trudy Paleontologièeskog Instituta Akademii Nauk SSSR [Proceedings of the Paleontological Institute, USSR Academy of Sciences] 45, 185.
Owen, R. 1867. On the dental characters of genera and species, chiefly of fishes from the Low Main Seam and shales of coal, Northumberland. Transactions of the Odontological Society of Great Britain 5, 329334.
Patterson, C. 1965. The phylogeny of chimaeroids. Philosophical Transactions of the Royal Society, London, Series B 249, 101219.
Patterson, C. 1992. Interpretation of the toothplates of chimaeroid fishes. Zoological Journal of the Linnean Society 106, 3361.
Pemberton, S. G., Spila, M., Pulham, A. J., Saunders, T., MacEachern, J. A., Robbins, D. & Sinclair, I. K. 2001. Ichnology and sedimentology of shallow to marginal marine systems: Ben Nevis & Avalon reservoirs, Jeanne D'Arc Baisin. Geological Association of Canada, Short Course Notes 15. St Johns, Newfoundland: Geological Association of Canada. 343 pp.
Richards, K. R. 2016. Carboniferous chondrichthyans from the Derbyshire limestones. Doctoral Thesis, University of Cambridge. 326 pp.
Roelofs, B., Barham, M., Mory, A. J. & Trinajstic, K. 2016. Late Devonian and Early Carboniferous chondrichthyans from the Fairfield Group, Canning Basin, Western Australia. Palaeontologia Electronica 19, 128.
Salamon, M. A., Gorzelak, P., Niedźwiedzki, R., Trzęsiok, D. & Baumiller, T. K. 2014. Trends in shell fragmentation as evidence of mid-Paleozoic changes in marine predation. Paleobiology 40, 1423.
Sallan, L. C., Kammer, T. W., Ausich, W. I. & Cook, L. A. 2011. Persistent predator–prey dynamics revealed by mass extinction. Proceedings of the National Academy of Sciences USA 108, 83358338.
Sallan, L. C. & Coates, M. I. 2010. End-Devonian extinction and a bottleneck in the early evolution of modern jawed vertebrates. Proceedings of the National Academy of Sciences USA 107, 10131–35.
Sallan, L. C. & Coates, M. I. 2013. Styracopterid (Actinopterygii) ontogeny and the multiple origins of post-Hangenberg deep-bodied fishes. Zoological Journal of the Linnean Society 169, 156199.
Schram, F. R. 1979. British Carboniferous Malacostraca. Fieldiana Geology 40, 1–129.
Smithson, T. R., Richards, K. R. & Clack, J. A. 2015. Lungfish diversity in Romer's Gap: reaction to the end-Devonian extinction. Palaeontology 59, 2944.
Stahl, B. J. 1999. Chondrichthyes III. Holocephali, In Schultze, H-P. (ed.) Handbook of Paleoichthyology, Volume 3. München: Verlag Dr. Friedrich Pfeil.
St John, O. H. & Worthen, A. H. 1875. Descriptions of fossil fishes. Geological Survey of Illinois 6, 245488.
St John, O. H. & Worthen, A. H. 1883. Descriptions of fossil fishes; a partial revision of the Cochliodonts and Psammodonts. Geological Survey of Illinois 7, 55264.
Stoddart, W. W. 1875. The Geology of the Bristol coal-field. Proceedings of the Bristol Naturalist's Society 1, 115140.
Stone, P., Millward, D., Young, B., Merritt, J. W., Clarke, S. M., McCormac, M. & Lawrence, D. J. D. 2010. British Regional Geology: Northern England (Fifth edition). Keyworth, Nottingham, UK: British Geological Survey.
Turnau, E., Avchimovitch, V.I., Byvsheva, T. V., Carson, B., Clayton, G. & Owens, B. 1997. The first appearance in Europe of Lycospora pusilla (Ibrahim) Somers and its relationship to the Tournaisian/Viséan boundary. Prace Państwowego Instytutu Geologicznego 157, 289293.
Turner, S. 1982. Middle Palaeozoic elasmobranch remains from Australia. Journal of Vertebrate Paleontology 2, 117131.
Turner, S. 1991. Palaeozoic vertebrate microfossils of Australia. In Vickers-Rich, P., Monaghan, J. N., Baird, R. F. & Rich, T. H. (eds) Vertebrate palaeontology of Australasia, 429464. Melbourne: Pioneer Design Studios. xvi+1437 pp.
Turner, S. 1993. Palaeozoic microvertebrates biostratigraphy of Eastern Gondwana. In Long, J. A. (ed.) Palaeozoic vertebrate biostratigraphy and biogeography, 174207. London: Belhaven Press. 384 pp.
Underwood, C. J., Mitchell, S. F. & Veltkamp, K. J. 1999. Shark and ray teeth from the Hauterivian (Lower Cretaceous) of northeast England. Palaeontology 42, 287302.
Waters, C. N., Waters, R. A., Jonas, N. S., Cleal, C. J. & Davies, J. R. 2011. Bristol, Mendips and Forest of Dean. In Waters, C. N., Somerville, I. D., Jones, N. S., Cleal, C. J., Collinson, J. D., Waters, R. A., Besly, B. M., Dean, M. T., Stephenson, M. H., Davies, J. R., Freshney, E. C., Jackson, D. I., Mitchell, W. I., Powell, J. H., Barclay, W. J., Browne, M. A. E., Leveridge, B. E., Long, S. L. & McLean, D. (eds) A revised correlation of Carboniferous rocks in the British Isles. Geological Society of London, Special Report 26, 3743. London & Bath: The Geological Society. 186 pp.
Whitenack, L. B., Simkins, D. C. & Motta, P. J. 2011. Biology meets engineering: the structural mechanics of fossil and extant shark teeth. Journal of Morphology 272, 169179.
Whitenack, L. B. & Motta, P. J. 2010. Performance of shark teeth during puncture and draw: implications for the mechanics of cutting. Biological Journal of the Linnean Society 100, 271286.
Williams, M., Leng, M. L., Stephenson, M. H., Andrews, J. E., Wilkinson, I. P., Siveter, D. J., Horne, D. J. & Vannier, J. M. C. 2006. Evidence that Early Carboniferous ostracods colonised coastal flood plain brackish water environments. Palaeogeography, Palaeoclimatology, Palaeoecology 230, 299318.
Williams, M. E. 1990. Feeding behavior in Cleveland Shale fishes. In Boucot, A. J. (ed.) Evolutionary Paleobiology of Behavior and Coevolution, 273287. Amsterdam: Elsevier Science. 725 pp.
Wood, S. P. & Rolfe, W. D. I. 1985. Introduction to the palaeontology of the Dinantian of Foulden, Berwickshire, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 76, 16.
Woodward, A. S. 1889. Catalogue of the fossil fishes in the British Museum (Natural History) Part 1. London: British Museum (Natural History).
Zangerl, R. 1981. Chondrichthyes I. Paleozoic Elasmobranchii. In Schultze, H-P. (ed.) Handbook of Paleoichthyology, Vol. 3A. Stuttgart: Gustav Fischer. 115 pp.


Diverse and durophagous: Early Carboniferous chondrichthyans from the Scottish Borders

  • Kelly R. Richards (a1), Janet E. Sherwin (a2), Timothy R. Smithson (a1), Rebecca F. Bennion (a1), Sarah J. Davies (a2), John E. A. Marshall (a3) and Jennifer A. Clack (a1)...


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed