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Large trilobites in a stress-free Early Ordovician environment

Published online by Cambridge University Press:  18 May 2020

Farid Saleh*
Affiliation:
Université de Lyon, Université Claude Bernard Lyon1, École Normale Supérieure de Lyon, CNRS, UMR5276, LGL-TPE, Villeurbanne, France
Muriel Vidal
Affiliation:
Université de Brest, CNRS, IUEM Institut Universitaire Européen de la Mer, UMR 6538 Laboratoire Géosciences Océan, Place Nicolas Copernic, 29280Plouzané, France
Lukáš Laibl
Affiliation:
Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015Lausanne, Switzerland Czech Academy of Sciences, Institute of Geology, Rozvojová 269, 165 00 Prague 6, Czech Republic Institute of Geology and Palaeontology, Faculty of Science, Charles University, Albertov 6, Prague, 12843, Czech Republic
Pierre Sansjofre
Affiliation:
MNHN, Sorbonne Université, CNRS UMR 7590, Institut de Minéralogie, de Physique des Matériaux et de Cosmochimie, Paris, France
Pierre Gueriau
Affiliation:
Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015Lausanne, Switzerland
Francesc Pérez-Peris
Affiliation:
Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015Lausanne, Switzerland
Lorenzo Lustri
Affiliation:
Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015Lausanne, Switzerland
Victoire Lucas
Affiliation:
Université de Lyon, Université Claude Bernard Lyon1, École Normale Supérieure de Lyon, CNRS, UMR5276, LGL-TPE, Villeurbanne, France
Bertrand Lefebvre
Affiliation:
Université de Lyon, Université Claude Bernard Lyon1, École Normale Supérieure de Lyon, CNRS, UMR5276, LGL-TPE, Villeurbanne, France
Bernard Pittet
Affiliation:
Université de Lyon, Université Claude Bernard Lyon1, École Normale Supérieure de Lyon, CNRS, UMR5276, LGL-TPE, Villeurbanne, France
Khadija El Hariri
Affiliation:
Département des Sciences de la Terre, Faculté des Sciences et Techniques, Université Cadi-Ayyad, BP 549, 40000Marrakesh, Morocco
Allison C. Daley
Affiliation:
Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015Lausanne, Switzerland
*
Author for Correspondence: Farid Saleh, Email: [email protected]

Abstract

Understanding variations in body size is essential for deciphering the response of an organism to its surrounding environmental conditions and its ecological adaptations. In modern environments, large marine animals are mostly found in cold waters. However, numerous parameters can influence body-size variations other than temperatures, such as oxygenation, nutrient availability, predation or physical disturbances by storms. Here, we investigate trilobite size variations in the Lower Ordovician Fezouata Shale deposited in a cold-water environment. Trilobite assemblages dominated by small- to normal-sized specimens that are a few centimetres in length are found in proximal and intermediate settings, while those comprising larger taxa more than 20 cm in length are found in the most distal environment of the Fezouata Shale. Drill core material from distal settings shows that sedimentary rocks hosting large trilobites preserved in situ are extensively bioturbated with a high diversity of trace fossils, indicating that oxygen and nutrients were available in this environment. In intermediate and shallow settings, bioturbation is less extensive and shallower in depth. The rarity of storm events (minimal physical disturbance) and the lack of predators in deep environments in comparison to shallower settings would also have helped trilobites attain larger body sizes. This highly resolved spatial study investigating the effects of numerous biotic and abiotic parameters on body size has wider implications for the understanding of size fluctuations over geological time.

Type
Original Article
Copyright
© The Author(s), 2020. Published by Cambridge University Press

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References

Barry, JP and Dayton, PK (1991) Physical heterogeneity and the organization of marine communities. In Ecological Heterogeneity (eds Kolasa, J and Pickett, ST), pp. 270320. New York, NY: Springer.CrossRefGoogle Scholar
Bell, MA (2014) Patterns in palaeontology: trends of body-size evolution in the fossil record – a growing field. Palaeontology online 4, 19.Google Scholar
Bonner, JT (2006) Why Size Matters: From Bacteria to Blue Whales. Princeton: Princeton University Press, 161 p.Google Scholar
Booth, JAT, Ruhl, HA, Lovell, LL, Bailey, DM and Smith, KL (2008) Size–frequency dynamics of NE Pacific abyssal ophiuroids (Echinodermata: Ophiuroidea). Marine Biology 154(6), 933–41.CrossRefGoogle Scholar
Capéra, JC, Courtessole, R and Pillet, J (1975) Biostratigraphie de l’Ordovicien inférieur de la Montagne Noire (France méridionale) – Trémadocien inférieur. Bulletin de la Société d’Histoire Naturelle de Toulouse, 111(3–4), 337–80.Google Scholar
Capéra, JC, Courtessole, R and Pillet, J (1978) Contribution à l’étude de l’Ordovicien inférieur de la Montagne Noire. Biostratigraphie et révision des Agnostida. Annales de la Société Géologique du Nord 98, 6788.Google Scholar
Chapelle, G and Peck, LS (1999) Polar gigantism dictated by oxygen availability. Nature 399, 114–15.CrossRefGoogle Scholar
Chatterton, BDE (1980) Ontogenetic studies of Middle Ordovician trilobites from the Esbataottine Formation, Mackenzie Mountains, Canada. Palaeontographica Abteilung A, 1980, 174.Google Scholar
Chatterton, BDE and Speyer, SE (1997) Ontogeny. In Treatise on Invertebrate Paleontology, Part O, Trilobita, revised, Volume 1: Introduction (ed Kaesler, RL), pp. 173247. Boulder, Colorado: The Geological Society of America and The Univeristy of Kansas.Google Scholar
Cherel, Y and Hobson, KA (2005) Stable isotopes, beaks and predators: a new tool to study the trophic ecology of cephalopods, including giant and colossal squids. Proceedings of the Royal Society B: Biological Sciences 272(1572), 1601–07.CrossRefGoogle Scholar
Conner, W, Day, J, Baumann, R and Randall, J (1989) Influence of hurricanes on coastal ecosystems along the northern Gulf of Mexico. Wetlands Ecology and Management 1(1), 4556.CrossRefGoogle Scholar
Corbacho, J and López-Soriano, FJ (2012) A new asaphid trilobite from the Lower Ordovician (Arenig) of Morocco. Batalleria 17, 312.Google Scholar
Corbacho, J and López-Soriano, FJ (2016) Platypeltoides hammondi (Trilobita, Nileidae): A new species from the Upper Tremadoc of the Dra Valley, Morocco. Batalleria 23, 1119.Google Scholar
Corbacho, J, Lopez-Soriano, FJ, Lemke, U and Hammond, K (2017) Platypeltoides carmenae: A new Nileidae (Trilobita) from the Lower Ordovician (Tremadocian) of Guelmim area; Western Anti-Atlas, Morocco. Batalleria 25, 2029.Google Scholar
Corbacho, J, Lopez-Soriano, FJ, Lemke, U, Morrison, S and Hammond, K (2018) Diversity and distribution of the genus Platypeltoides (Nileidae) in Morocco. American Journal of Bioscience and Bioengineering 6(2), 1320.CrossRefGoogle Scholar
Courtessole, R and Pillet, J (1975) Contribution à l’étude des faunes trilobitiques de l’Ordovicien inférieur de la Montagne Noire. Les Eulominae et les Nileidae. Annales de la Société Géologique du Nord 95(4), 251–72.Google Scholar
Courtessole, R, Pillet, J and Vizcaïno, D (1981) Nouvelles données sur la biostratigraphie de l’Ordovicien inférieur de la Montagne Noire. Révision des Taihugshaniidae, de Megistaspis (Ekeraspis) et d’Asaphopsoides (Trilobites). Carcassonne: Mémoire de la Société des Etudes Scientifiques de l’Aude, 32 p.Google Scholar
Destombes, J, Hollard, H and Willefert, S (1985) Lower Palaeozoic rocks of Morocco. In Lower Palaeozoic Rocks of the World (ed Holland, C), pp. 91336. New York: Wiley. Google Scholar
Ebbestad, JOR (2016) Gastropoda, Tergomya and Paragastropoda (Mollusca) from the Lower Ordovician Fezouata Formation, Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 460, 8796.CrossRefGoogle Scholar
Fagerstrom, JA (1964) Fossil communities in paleoecology: their recognition and significance. GSA Bulletin 75(12), 1197–216.CrossRefGoogle Scholar
Fortey, R (2009) A new giant asaphid trilobite from the Lower Ordovician of Morocco. Memoirs of the Association of Australasian Palaeontologists 37, 916.Google Scholar
Fortey, RA (2010) Trilobites of the genus Dikelokephalina from Ordovician Gondwana and Avalonia. Geological Journal 46(5), 405–15.Google Scholar
Gaines, RR, Briggs, DEG, Orr, PJ and Van Roy, P (2012) Preservation of giant anomalocaridids in silica-chlorite concretions from the Early Ordovician of Morocco. PALAIOS 27(5), 317–25.CrossRefGoogle Scholar
Geiger, M, Wilson, LAB, Costeur, L, Sánchez, R and Sánchez-Villagra, MR (2013) Diversity and body size in giant caviomorphs (Rodentia) from the northern Neotropics—a study of femoral variation. Journal of Vertebrate Paleontology 33(6), 1449–56.CrossRefGoogle Scholar
Geyer, G (1993) The giant Cambrian trilobites of Morocco. Beringeria 8, 71107.Google Scholar
Guiraud, R, Bosworth, B, Thierry, J and Delplanque, A (2004) Phanerozoic geological evolution of Northern and Central Africa: An overview. Journal of African Earth Sciences 43, 83143.CrossRefGoogle Scholar
Gutiérrez-Marco, JC and García-Bellido, DC (2018) The international fossil trade from the Paleozoic of the Anti-Atlas, Morocco. In The Great Ordovician Biodiversification Event: Insights from the Tifilalt Biota, Morocco (eds Hunter, AW, Álvaro, JJ, Lefebvre, B, van Roy, P and Zamora, S). Geological Society of London, Special Publication no. 485, published online 1 January 2018, SP485.1.Google Scholar
Gutiérrez-Marco, JC, , AA, García-Bellido, DC, Rábano, I and Valério, M (2009) Giant trilobites and trilobite clusters from the Ordovician of Portugal. Geology 37(5), 443–46.Google Scholar
Harrington, HJ (1937) On some Ordovician fossils from Northern Argentina. Geological Magazine 74(3), 97124.CrossRefGoogle Scholar
Holmes, JD, Paterson, JR and Garcıa-Bellido, DC (2020) The trilobite Redlichia from the lower Cambrian Emu Bay Shale Konservat-Lagerstätte of South Australia: systematics, ontogeny and soft-part anatomy. Journal of Systematic Palaeontology 18(4), 295334.CrossRefGoogle Scholar
Horikoshi, M (1986) Biology of the deep sea. In The Seas Around Japan (eds Horikoshi, MY, Nagata, Y and Sato, T), pp. 169–92. Tokyo: Iwanami Shoten.Google Scholar
Hughes, NC and Rushton, AWA (1990) Computer-aided restoration of a late Cambrian ceratopygid trilobite from Wales, and its phylogenetic implications. Palaeontology 33(2), 429–45.Google Scholar
Hunt, J, Butler, M and Herrnkind, W (1994) Sponge mass mortality and Hurricane Andrew: catastrophe for juvenile spiny lobsters in south Florida? Bulletin of Marine Science 54(3), 1073.Google Scholar
Johnson, RG (1960) Models and methods for analysis of the mode of formation of fossil assemblages. GSA Bulletin 71(7), 1075–86.CrossRefGoogle Scholar
Klug, C, De Baets, K, Kröger, B, Bell, MA, Korn, D and Payne, JL (2015) Normal giants? Temporal and latitudinal shifts of Palaeozoic marine invertebrate gigantism and global change. Lethaia 48(2), 267–88.CrossRefGoogle Scholar
Kröger, B and Lefebvre, B (2012) Palaeogeography and palaeoecology of early Floian (Early Ordovician) cephalopods from the Upper Fezouata Formation, Anti-Atlas, Morocco. Fossil Record 15(2), 6175.CrossRefGoogle Scholar
Kröger, B, Servais, T and Zhang, Y (2009a) The origin and initial rise of pelagic cephalopods in the Ordovician. PLoS ONE 4(9), e7262.Google ScholarPubMed
Kröger, B, Zhang, Y and Isakar, M (2009b) Discosorids and oncocerids (Cephalopoda) of the Middle Ordovician Kunda and Aseri regional stages of Baltoscandia and the early evolution of these groups. Geobios 42(3), 273–92.CrossRefGoogle Scholar
Laibl, L, Fatka, O, Budil, P, Ahlberg, P, Szabad, M, Vokáč, V and Kozák, V (2015) The ontogeny of Ellipsocephalus (Trilobita) and systematic position of Ellipsocephalidae. Alcheringa: An Australasian Journal of Palaeontology 39(4), 477–87.CrossRefGoogle Scholar
Laibl, L, Fatka, O, Cronier, C and Budil, P (2014) Early ontogeny of the Cambrian trilobite Sao hirsuta from the Skryje-Týřovice Basin, Barrandian area, Czech Republic. Bulletin of Geosciences 89(2), 293309.CrossRefGoogle Scholar
Lamsdell, JC and Braddy, SJ (2009) Cope’s Rule and Romer’s theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates. Biology Letters 6(2), 265–69.CrossRefGoogle ScholarPubMed
Lebrun, P (2018) Fossiles du Maroc. Tome I, Gisements emblématiques du Paléozoïque de l’Anti-Atlas. Glavenat: les Editions du Piat, 298 p.Google Scholar
Lefebvre, B, Allaire, N, Guensburg, T, , E, Hunter, AW, Kouraïss, K, Martin, EL, , O, Nardin, E, Noailles, F, Pittet, B, Sumrall, C, , D and Zamora, S (2016) Palaeoecological aspects of the diversification of echinoderms in the Lower Ordovician of central Anti-Atlas, Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 460, 97121.CrossRefGoogle Scholar
Lefebvre, B, Guensburg, TE, Martin, ELO, Mooi, R, Nardin, E, Nohejlová, M, Saleh, F, Kouraïss, K, El Hariri, K and David, B (2019) Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes. Geobios 52, 2736.CrossRefGoogle Scholar
Luquet, G (2012) Biomineralizations: insights and prospects from crustaceans. ZooKeys 176, 103–21.CrossRefGoogle Scholar
Martí Mus, M (2016) A hyolithid with preserved soft parts from the Ordovician Fezouata Konservat-Lagerstätte of Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 460, 122–29.CrossRefGoogle Scholar
Martin, E, Lefebvre, B and Vaucher, R (2015) Taphonomy of a stylophoran-dominated assemblage in the Lower Ordovician of Zagora area (central Anti-Atlas, Morocco). In Progress in Echinoderm Palaeobiology (eds Zamora, S and Rabano, I). Madrid: Cuadernos del Museo Geominero 19, 95100.Google Scholar
Martin, ELO, Pittet, B, Gutiérrez-Marco, J-C, Vannier, J, El Hariri, K, Lerosey-Aubril, R, Masrour, M, Nowak, H, Servais, T, Vandenbroucke, TRA, Van Roy, P, Vaucher, R and Lefebvre, B (2016a) The Lower Ordovician Fezouata Konservat-Lagerstätte from Morocco: Age, environment and evolutionary perspectives. Gondwana Research 34, 274–83.CrossRefGoogle Scholar
Martin, ELO, Vidal, M, Vizcaïno, D, Vaucher, R, Sansjofre, P, Lefebvre, B and Destombes, J (2016b) Biostratigraphic and palaeoenvironmental controls on the trilobite associations from the Lower Ordovician Fezouata Shale of the central Anti-Atlas, Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 460, 142154.CrossRefGoogle Scholar
McAlister, JS and Stancyk, SE (2005) Effects of variable water motion on regeneration of Hemipholis elongata (Echinodermata, Ophiuroidea). Invertebrate Biology 122(2), 166–76.CrossRefGoogle Scholar
McCoy, VE, Young, RT and Briggs, DEG (2015a) Factors controlling exceptional preservation in concretions. Palaios 30, 272–80.CrossRefGoogle Scholar
McCoy, VE, Young, RT and Briggs, DEG (2015b) Sediment permeability and the preservation of soft-tissues in concretions: an experimental study. Palaios 30, 608–12.CrossRefGoogle Scholar
Mergl, M (2006) Tremadocian trilobites of the Prague Basin, Czech Republic. Acta Musei Nationalis Pragae, Series B – Historia Naturalis 62(1–2), 170.Google Scholar
Moran, A and Woods, A (2012) Why might they be giants? Towards an understanding of polar gigantism. Journal of Experimental Biology 215(2), 19952002.CrossRefGoogle ScholarPubMed
Nichols, G (2009) Sedimentology and Stratigraphy. Chichester: Wiley-Blackwell, 432 p.Google Scholar
Nowak, H, Pittet, B, Vaucher, R, Akodad, M, Gaines, RR and Vandenbroucke, TRA (2016) Palynomorphs of the Fezouata Shale (Lower Ordovician, Morocco): Age and environmental constraints of the Fezouata Biota. Palaeogeography, Palaeoclimatology, Palaeoecology 460, 6274.CrossRefGoogle Scholar
Park, T and Choi, DK (2009) Post-embryonic development of the Furongian (late Cambrian) trilobite Tsinania canens: implications for life mode and phylogeny. Evolution & Development 11(4), 441–55.CrossRefGoogle ScholarPubMed
Paterson, JR, García-Bellido, DC, Jago, JB, Gehling, JG, Lee, MSY and Edgecombe, GD (2016) The Emu Bay Shale Konservat-Lagerstätte: a view of Cambrian life from East Gondwana. Journal of the Geological Society 173(1), 111.CrossRefGoogle Scholar
Payne, JL, Groves, JR, Jost, AB, Nguyen, T, Moffitt, SE, Hill, TM and Skotheim, JM (2012) Late Paleozoic fusulinoidean gigantism driven by atmospheric hyperoxia. Evolution 66(9), 2929–39.CrossRefGoogle ScholarPubMed
Perillo, MM, Best, JL and Garcia, MH (2014) A new phase diagram for combined-flow bedforms. Journal of Sedimentary Research 84(4), 301–13.CrossRefGoogle Scholar
Philippart, CJM, Cadée, GC, van Raaphorst, W and Riegman, R (2000) Long-term phytoplankton-nutrient interactions in a shallow coastal sea: Algal community structure, nutrient budgets, and denitrification potential. Limnology and Oceanography 45(1), 131–44.CrossRefGoogle Scholar
Rábano, I (1990) Platypeltoides magrebiensis n. sp., Asaphina, Nileidae, del Ordovicico inferior del Anti-Atlas central, Marruecos. Boletin Geologico y Minero 101, 2147.Google Scholar
Rex, MA (1976) Biological accomodation in the deep-sea benthos: comparative evidence on the importance of predation and productivity. Deep Sea Research and Oceanographic Abstracts 23(10), 975–87.CrossRefGoogle Scholar
Rowe, GT, Clifford, CH, Smith, KL and Hamilton, PL (1975) Benthic nutrient regeneration and its coupling to primary productivity in coastal waters. Nature 255(5505), 215–17.CrossRefGoogle Scholar
Rudkin, DM, Young, GA, Elias, RJ and Dobrzanski, EP (2003) The world’s biggest trilobite— Isotelus rex new species from the upper Ordovician of northern Manitoba, Canada. Journal of Paleontology 77(1), 99112.CrossRefGoogle Scholar
Saleh, F, Candela, Y, Harper, DAT, Polechovà, M, Pittet, B and Lefebvre, B (2018) Storm-induced community dynamics in the Fezouata Biota (Lower Ordovician, Morocco). Palaios 33(12), 535–41.CrossRefGoogle Scholar
Saleh, F, Pittet, B, Perrillat, J and Lefebvre, B (2019) Orbital control on exceptional fossil preservation. Geology 47, 15.CrossRefGoogle Scholar
Sander, M and Clauss, M (2008) Sauropod gigantism. Science 322, 201–2.CrossRefGoogle ScholarPubMed
Sdzuy, K, Hammann, W and Villas, E (2001) The Upper Tremadocian fauna from Vogtendorf and the Bavarian Ordovician of the Frankenwald (Germany). Senckenbergiana Lethaea, 81(1), 207–61.CrossRefGoogle Scholar
Sigurdsen, A and Hammer, Ø (2016) Body size trends in the Ordovician to earliest Silurian of the Oslo Region. Palaeogeography, Palaeoclimatology, Palaeoecology 443, 4956.CrossRefGoogle Scholar
Speyer, SE and Brett, CE (1986) Trilobite taphonomy and Middle Devonian taphofacies. PALAIOS 1(3), 312.CrossRefGoogle Scholar
Tabb, DC and Jones, AC (1962) Effect of Hurricane Donna on the aquatic fauna of North Florida Bay. Transactions of the American Fisheries Society 91(4), 375–78.CrossRefGoogle Scholar
Taylor, AM and Goldring, R (1993) Description and analysis of bioturbation and ichnofabric. Journal of the Geological Society 150(1), 141–8.CrossRefGoogle Scholar
Teigler, DJ and Towe, KM (1975) Microstructure and composition of the trilobite exoskeleton. Fossils and Strata 4, 137–49.Google Scholar
Thoral, M (1935) Contribution à l’étude paléontologique de l’Ordovicien inférieur de la Montagne Noire et Révision sommaire de la faune cambrienne de la Montagne Noire. Montpellier: Imprimerie de la Charité, 362 p.Google Scholar
Thoral, M (1946) Cycles géologiques et formations nodulaires de la Montagne Noire. Nouvelles archives du muséum d’histoire naturelle de Lyon 1, 1141.Google Scholar
Timofeev, SF (2001) Bergmann’s principle and deep-water gigantism in marine crustaceans. Biology Bulletin 28(6), 646–50.CrossRefGoogle Scholar
Torsvik, T and Cocks, L (2011) The Palaeozoic palaeogeography of central Gondwana. In The Formation and Evolution of Africa: A Synopsis of 3.8 Ga of Earth History (eds van Hinsbergen, DJJ, Buiter, SJH, Torsvik, TH, Gaina, C and Webb, SJ), pp. 137–66. Geological Society of London, Special Publication no. 357.CrossRefGoogle Scholar
Torsvik, T and Cocks, L (2013) New global palaeogeographical reconstructions for the Early Palaeozoic and their generation. In Early Palaeozoic Biogeography and Palaeogeography (eds Harper, DAT and Servais, T), pp. 524. Geological Society of London, Memoir no. 38.Google Scholar
Twitchett, RJ (2007) The Lilliput effect in the aftermath of the end-Permian extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 252, 132–44.CrossRefGoogle Scholar
Vannier, J, Vidal, M, Marchant, R, El Hariri, K, Kouraiss, K, Pittet, B, El Albani, A, Mazurier, A. and Martin, E (2019) Collective behaviour in 480-million-year-old trilobite arthropods from Morocco. Scientific Reports 9, 14941.CrossRefGoogle ScholarPubMed
Van Roy, P and Briggs, DEG (2011) A giant Ordovician anomalocaridid. Nature 473(7348), 510–13.CrossRefGoogle ScholarPubMed
Van Roy, P, Briggs, DEG and Gaines, RR (2015a) The Fezouata fossils of Morocco; an extraordinary record of marine life in the Early Ordovician. Journal of the Geological Society 172(5), 541–49.CrossRefGoogle Scholar
Van Roy, P, Daley, AC and Briggs, DEG (2015b) Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps. Nature 522(7554), 7780.CrossRefGoogle ScholarPubMed
Van Roy, P, Orr, PJ, Botting, JP, Muir, LA, Vinther, J, Lefebvre, B, El Hariri, K and Briggs, DEG (2010) Ordovician faunas of Burgess Shale type. Nature 465(7295), 215–18.CrossRefGoogle ScholarPubMed
Vaucher, R, Martin, ELO, Hormière, H and Pittet, B (2016) A genetic link between Konzentrat- and Konservat-Lagerstätten in the Fezouata Shale (Lower Ordovician, Morocco). Palaeogeography, Palaeoclimatology, Palaeoecology 460, 2434.CrossRefGoogle Scholar
Vaucher, R, Pittet, B, Hormière, H, Martin, ELO and Lefebvre, B (2017) A wave-dominated, tide-modulated model for the Lower Ordovician of the Anti-Atlas, Morocco. Sedimentology 64(3), 777807.CrossRefGoogle Scholar
Vela, JA and Corbacho, J (2007) A new species of Lehua from Lower Ordovician of Dra Valley of Morocco. Batalleria 13, 7580.Google Scholar
Vermeij, GJ (2016) Gigantism and its implications for the history of life. PLoS ONE 11(1), e0146092.CrossRefGoogle ScholarPubMed
Vinther, J, Parry, L, Briggs, DEG and Van Roy, P (2017) Ancestral morphology of crown-group molluscs revealed by a new Ordovician stem aculiferan. Nature 542(7642), 471–74.CrossRefGoogle ScholarPubMed
Vinther, J, Van Roy, P and Briggs, DEG (2008) Machaeridians are Palaeozoic armoured annelids. Nature 451(7175), 185–88.CrossRefGoogle ScholarPubMed
Wang, B, Wang, X and Zhan, R (2003) Nutrient conditions in the Yellow Sea and the East China Sea. Estuarine, Coastal and Shelf Science 58(1), 127–36.CrossRefGoogle Scholar
Whiteley, TE, Kloc, GJ and Brett, CE (2002) Trilobites of New York: An Illustrated Guide. Ithaca, NY: Paleontological Research Institution, 20 p.Google Scholar
Zeuthen, E (1953) Oxygen uptake as related to body size in organisms. The Quarterly Review of Biology 28(1), 112.CrossRefGoogle ScholarPubMed
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