Skip to main content Accessibility help

Stylonurine eurypterids from the Strud locality (Upper Devonian, Belgium): new insights into the ecology of freshwater sea scorpions

  • James C Lamsdell (a1), Linda Lagebro (a2), Gregory D Edgecombe (a3), Graham E Budd (a2) and Pierre Gueriau (a4)...


The Upper Famennian (Upper Devonian) Strud locality has yielded very abundant and diversified flora as well as vertebrate and arthropod faunas. The arthropod fauna, mostly recovered from fine shales deposited in a calm, confined floodplain habitat including temporary pools, has delivered a putative insect and various crustaceans including eumalacostracans and notostracan, spinicaudatan and anostracan branchiopods. Here we present the Strud eurypterids, consisting of semi-articulated juvenile specimens assigned to Hardieopteridae recovered from the pool and floodplain deposits, as well as larger isolated fragments of potential adults recovered from stratigraphically lower, coarser dark sandy layers indicative of a higher-energy fluvial environment. The Strud fossils strongly suggest that, as proposed for some Carboniferous eurypterids, juvenile freshwater eurypterids inhabited sheltered nursery pools and migrated to higher-energy river systems as they matured.


Corresponding author


Hide All
Andrews, HE, Brower, JC, Gould, SJ and Reyment, RA (1974) Growth and variation in Eurypterus remipes Dekay. Bulletin of the Geological Institution of the University of Uppsala 4, 81114.
Botton, ML, Tankersley, RA and Loveland, RE (2010) Developmental ecology of the American horseshoe crab Limulus polyphemus. Current Zoology 56, 550–62.
Braddy, SJ (2001) Eurypterid palaeoecology: palaeobiological, ichnological and comparative evidence for a ‘mass-moult–mate’ hypothesis. Palaeogeography, Palaeoclimatology, Palaeoecology 172, 115–32.
Brower, JC and Veinus, J (1974) The statistical zap versus the shotgun approach. Mathematical Geology 6, 311–32.
Brower, JC and Veinus, J (1978) Multivariate analysis of allometry using point coordinates. Journal of Paleontology 52, 1037–53.
Burmeister, H (1843) Die Organisation der Trilobiten, aus ihren lebenden Verwandten entwickelt, nebst einer systematischen Uebersicht aller zeither beschriebenen Arten. Berlin: Reimer, 147 pp. + 6 pls.
Clément, G, Ahlberg, PE, Blieck, A, Blom, H, Clack, JA, Poty, E, Thorez, J and Janvier, P (2004) Devonian tetrapod from Western Europe. Nature 427, 412–13.
Clément, G and Boisvert, CA (2006) Lohest’s true and false ‘Devonian amphibians’: evidence for the rhynchodipterid lungfish Soederberghia in the Famennian of Belgium. Journal of Vertebrate Paleontology 26, 276–83.
Cuggy, MJ (1994) Ontogenetic variation in Silurian eurypterids from Ontario and New York State. Canadian Journal of Earth Sciences 31, 728–32.
Denayer, J, Prestianni, C, Gueriau, P, Olive, S and Clément, G (2016) Stratigraphy and depositional environments of the Late Famennian (Late Devonian) of Southern Belgium and characterization of the Strud locality. Geological Magazine 153, 112–27.
Diener, C (1924) Fossilium catalogus I: animalia, Pars 25: Eurypterida. Berlin: Junk, 28 pp.
Dunlop, JA and Lamsdell, JC (2017) Segmentation and tagmosis in Chelicerata. Arthropod Structure & Development 46, 395418.
Ehlers, GM (1935) A new eurypterid from the Upper Devonian of Pennsylvania. Contribution from the Museum of Paleontology, University of Michigan 4, 291–5.
Fraipont, J (1889) Euryptérides nouveaux du Dévonien supérieur de Belgique (Psammites du Condroz). Annales de la Société géologique de Belgique 17, 5362.
Garrouste, R, Clément, G, Nel, P, Engel, MS, Grandcolas, P, D’Haese, C, Lagebro, L, Denayer, J, Gueriau, P, Lafaite, P, Olive, S, Prestianni, C and Nel, A (2012) A complete insect from the Late Devonian. Nature 488, 82–5.
Garrouste, R, Clément, G, Nel, P, Engel, MS, Grandcolas, P, D’Haese, C, Lagebro, L, Denayer, J, Gueriau, P, Lafaite, P, Olive, S, Prestianni, C and Nel, A (2013) Garrouste etal. reply. Nature 494, E4–5.
Gueriau, P, Charbonnier, S and Clément, G (2014 a) First decapod crustaceans in a Late Devonian continental ecosystem. Palaeontology 57, 1203–13.
Gueriau, P, Charbonnier, S and Clément, G (2014 b) Angustidontid crustaceans from the Late Devonian of Strud (Namur Province, Belgium): insights into the origin of Decapoda. Neues Jahrbuch für Geologie und Paläontologie 273, 327–37.
Gueriau, P, Rabet, N, Clément, G, Lagebro, L, Vannier, J, Briggs, DEG, Charbonnier, S, Olive, S and Béthoux, O (2016) A 365-million-year-old freshwater community reveals morphological and ecological stasis in branchiopod crustaceans. Current Biology 26, 383–90.
Gueriau, P, Rabet, N and Du Tien Hat, E (2018) The Strud crustacean fauna (Late Devonian, Belgium): updated review and palaeoecology of an early continental ecosystem. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 107, 7990.
Hall, J (1884) Description of a new species of Stylonurus from the Catskill Group. In 36th Annual Report of the New York State Museum of Natural History, pp. 76–7. Albany, NY: University of the State of New York.
Heymons, R (1901) Die Entwicklungsgeschichte der Scolopender. Zoologica 13, 1244.
Hörnschemeyer, T, Haug, J, Béthoux, O, Beutel, RG, Charbonnier, S, Hegna, TA, Koch, M, Rust, J, Wedmann, S, Bradler, S and Willmann, R (2013) Is Strudiella a Devonian insect? Nature 494, E3–4.
Jeram, AJ and Selden, PA (1994) Eurypterids from the Viséan of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 84, 301–8.
Kjellesvig-Waering, EN (1966) A revision of the families and genera of the Stylonuracea (Eurypterida). Fieldiana Geology 12, 85106.
Lagebro, L, Gueriau, P, Hegna, TA, Rabet, N and Budd, GE (2015) The oldest notostracan (Upper Devonian Strud locality, Belgium). Palaeontology 58, 497509.
Lamsdell, JC (2011) The eurypterid Stoermeropterus conicus from the Lower Silurian Pentland Hills, Scotland. Monograph of the Palaeontographical Society, London 165, 184, pls. 1–15.
Lamsdell, JC (2013) Redescription of Drepanopterus pentlandicus Laurie 1892, the earliest known mycteropoid (Chelicerata: Eurypterida) from the early Silurian (Llandovery) of the Pentland Hills, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 103, 77103.
Lamsdell, JC (2016) Horseshoe crab phylogeny and independent colonizations of fresh water: ecological invasion as a driver for morphological innovation. Palaeontology 59, 181–94.
Lamsdell, JC and Braddy, SJ (2010) Cope’s Rule and Romer’s theory: patterns of diversity and gigantism in eurypterids and Paleozoic vertebrates. Biology Letters 6, 265–9.
Lamsdell, JC, Braddy, SJ, Loeffler, EJ and Dineley, DL (2010 a) Early Devonian stylonurine eurypterids from Arctic Canada. Canadian Journal of Earth Sciences 47, 1405–15.
Lamsdell, JC, Braddy, SJ and Tetlie, OE (2009) Redescription of Drepanopterus abonensis (Chelicerata: Eurypterida: Stylonurina) from the Late Devonian of Portishead, UK. Palaeontology 52, 1113–39.
Lamsdell, JC, Braddy, SJ and Tetlie, OE (2010 b) The systematics and phylogeny of the Stylonurina (Arthropoda: Chelicerata: Eurypterida). Journal of Systematic Palaeontology 8, 4961.
Lamsdell, JC and Briggs, DEG (2017) The first diploaspidid (Chelicerata: Chasmataspidida) from North America (Silurian, Bertie Group, New York State) is the oldest species of Diploaspis. Geological Magazine 154, 175–80.
Lamsdell, JC, Briggs, DEG, Liu, HP, Witzke, BJ and McKay, RM (2015) The oldest described eurypterid: a giant Middle Ordovician (Darriwilian) megalograptid from the Winneshiek Lagerstätte of Iowa. BMC Evolutionary Biology, 15, 169. doi: 10.1186/s12862-015-0443-9.
Lamsdell, JC, Congreve, CR, Hopkins, MJ, Krug, AZ and Patzkowsky, ME (2017) Phylogenetic paleoecology: tree-thinking and ecology in deep time. Trends in Ecology and Evolution 32, 452–63.
Lamsdell, JC, Hoşgör, İ and Selden, PA (2013) A new Ordovician eurypterid (Arthropoda: Chelicerata) from southeast Turkey: evidence for a cryptic Ordovician record of Eurypterida. Gondwana Research 23, 354–66.
Lamsdell, JC and Selden, PA (2013) Babes in the wood – a unique window into sea scorpion ontogeny. BMC Evolutionary Biology 13, 98.
Lamsdell, JC and Selden, PA (2017) From success to persistence: identifying an evolutionary regime shift in the diverse Paleozoic aquatic arthropod group Eurypterida, driven by the Devonian biotic crisis. Evolution 71, 95110.
Laurie, M (1892) On some eurypterid remains from the Upper Silurian rocks of the Pentland Hills. Transactions of the Royal Society of Edinburgh: Earth Sciences 37, 151–61.
Lehman, JP (1959) Les Dipneustes du Dévonien supérieur du Groenland. Meddelelser om Grønland 160, 158.
Olive, S (2015) Devonian antiarch placoderms from Belgium revisited. Acta Palaeontologica Polonica 60, 711–31.
Olive, S, Ahlberg, PE, Pernègre, VN, Poty, É, Steurbaut, É and Clément, G (2016 a) New discoveries of tetrapods (ichthyostegid-like and whatcheeriid-like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium). Palaeontology 59, 827–40.
Olive, S, Clément, G, Daeschler, EB and Dupret, V (2015 a) Characterization of the placoderm (Gnathostomata) assemblage from the tetrapod-bearing locality of Strud (Belgium, upper Famennian). Palaeontology 58, 9811002.
Olive, S, Clément, G, Daeschler, EB and Dupret, V (2016 b) Placoderm assemblage from the tetrapod-bearing locality of Strud (Belgium, Upper Famennian) provides evidence for a fish nursery. PLoS ONE 11, e0161540. doi: 10.1371/journal.pone.0161540.
Olive, S, Clément, G, Denayer, J, Derycke, C, Dupret, V, Gerienne, P, Gueriau, P, Marion, JM, Mottequin, B and Prestianni, C (2015 b) Flora and fauna from a new Famennian (Upper Devonian) locality at Becco, eastern Belgium. Geologica Belgica 18, 92101.
Plax, DP, Lamsdell, JC, Vrazo, MB and Barbikov, DV (2018) A new genus of eurypterid (Chelicerata, Eurypterida) from the Upper Devonian salt deposits of Belarus. Journal of Paleontology 92, 838–49.
Ponomarenko, AG (1985) King crabs and eurypterids from the Permian and Mesozoic of the USSR. Paleontological Journal 3, 100–4.
Prestianni, C, Steel, M, Thorez, J and Gerrienne, P (2007) Strud: old quarry, new discoveries. Preliminary report. In Recent Advances in Palynology (eds Steemans, P and Javaux, E), pp. 43–7. Carnets de Géologie, Memoir.
Selden, PA (1981) Functional morphology of the prosoma of Baltoeurypterus tetragonophthalmus (Fischer) (Chelicerata: Eurypterida). Transactions of the Royal Society of Edinburgh, Earth Sciences 72, 948.
Siebold, CT von (1848) Lehrbuch der vergleichenden Anatomie der Wirbellosen Thiere. Erster Theil. In Lehrbuch der verglichenden Anatomie (eds Siebold, CT von and Stannius, H). Berlin: Verlag von Veit & Comp, 679 pp.
Størmer, L (1934) Merostomata from the Downtonian Sandstone of Ringerike, Norway. Skrifter utgitt av Det Norske Vidensakps-Akademi i Oslo I, Matem.-Naturvid. Klasse 10, 1125.
Størmer, L (1951) A new eurypterid from the Ordovician of Montgomeryshire, Wales. Geological Magazine 88, 409–22.
Størmer, L (1955) Merostomata. In Treatise on Invertebrate Paleontology, Pt. P, Arthropoda 2 (ed. Moore, RC), pp. 316. Lawrence, Kansas: Geological Society of America and University of Kansas Press.
Størmer, L and Waterston, CD (1968) IV. Cyrtoctenus gen. nov., a large late Palaeozoic arthropod with pectinate appendages. Transactions of the Royal Society of Edinburgh 68, 63104.
Tetlie, OE (2008) Hallipterus excelsior, a stylonurid (Chelicerata: Eurypterida) from the Late Devonian Catskill Delta complex, and its phylogenetic position in the Hardieopteridae. Bulletin of the Peabody Museum of Natural History 49, 1930.
Tetlie, OE, Anderson, LI and Poschmann, M (2007) Kiaeropterus (Eurypterida; Stylonurina) recognized from the Silurian of the Pentland Hills. Scottish Journal of Geology 43, 17.
Tetlie, OE, Brandt, DS and Briggs, DEG (2008) Ecdysis in sea scorpions (Chelicerata: Eurypterida). Palaeogeography, Palaeoclimatology, Palaeoecology 265, 182–94.
Tetlie, OE and Cuggy, MB (2007) Phylogeny of the basal swimming eurypterids (Chelicerata; Eurypterida; Eurypterina). Journal of Systematic Palaeontology 5, 345–56.
Tetlie, OE and Van Roy, P (2006) A reappraisal of Eurypterus dumonti Stainier, 1917 and its position within the Adelophthalmidae Tollerton, 1989. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre 76, 7990.
Tollerton, VP Jr (1989) Morphology, taxonomy, and classification of the order Eurypterida Burmeister, 1843. Journal of Paleontology 63, 642–57.
Waterston, CD (1979) Problems of functional morphology and classification in stylonuroid eurypterids (Chelicerata, Merostomata), with observations on the Scottish Silurian Stylonuroidea. Transactions of the Royal Society of Edinburgh, Earth Sciences 70, 251322.
Whyte, MA (2005) A gigantic fossil arthropod trackway. Nature 438, 576.


Stylonurine eurypterids from the Strud locality (Upper Devonian, Belgium): new insights into the ecology of freshwater sea scorpions

  • James C Lamsdell (a1), Linda Lagebro (a2), Gregory D Edgecombe (a3), Graham E Budd (a2) and Pierre Gueriau (a4)...


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