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Large Trilobites in a Stress-Free Early Ordovician Environment

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Large Trilobites in a Stress-Free Early Ordovician Environment Large trilobites in a stress-free Early Ordovician environment Farid Saleh, Muriel Vidal, Lukáš Laibl, Pierre Sansjofre, Pierre Guériau, Francesc Perez Peris, Lorenzo Lustri, Victoire Lucas, Bertrand Lefebvre, Bernard Pittet, et al. To cite this version: Farid Saleh, Muriel Vidal, Lukáš Laibl, Pierre Sansjofre, Pierre Guériau, et al.. Large trilobites in a stress-free Early Ordovician environment. Geological Magazine, Cambridge University Press (CUP), In press, pp.1-10. 10.1017/S0016756820000448. hal-03005087 HAL Id: hal-03005087 https://hal.archives-ouvertes.fr/hal-03005087 Submitted on 13 Nov 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Geological Magazine Large trilobites in a stress-free Early Ordovician environment Journal: Geological Magazine Manuscript ID GEO-19-2360.R2 Manuscript Type: Original Article Date Submitted by the 01-Apr-2020 Author: Complete List of Authors: Saleh, Farid; Université Claude Bernard Lyon 1, Sciences de la Terre Vidal, Muriel; Brest University Laibl, Lukáš; University of Lausanne ForSansjofre, Peer Pierre; Brest Review University Gueriau, Pierre; University of Lausanne, Institute of Earth Sciences Perez peris, Francesc; University of Lausanne Lustri, Lorenzo; University of Lausanne Lucas, Victoire; Université Claude Bernard Lyon 1 Lefebvre, Bertrand; Université Lyon 1, UMR CNRS 5276 LGLTPE PITTET, Bernard; Université Claude Bernard Lyon 1, LGL-TPE El Hariri, Khadija; Cadi Ayyad University Daley, Allison; University of Lausanne, Institute of Earth Sciences Keywords: Arthropod, Paleozoic, Body-size, Fezouata Shale 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 few cm in length are found in proximal and intermediate settings, while those comprising larger taxa more than 20cm in length Abstract: 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 have also 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. Cambridge University Press Page 1 of 38 Geological Magazine 1 2 3 1 Original Articles 4 5 6 2 Large trilobites in a stress-free Early Ordovician environment 7 8 3 FARID SALEH1*, MURIEL VIDAL2, LUKÁŠ LAIBL3,4,5, PIERRE SANSJOFRE2, PIERRE 9 10 4 GUERIAU3, FRANCESC PEREZ PERIS3, LORENZO LUSTRI3, VICTOIRE LUCAS2, 11 12 5 BERTRAND LEFEBVRE1, BERNARD PITTET1, KHADIJA EL HARIRI6 and ALLISON 13 14 3 15 6 C. DALEY 16 17 7 1Université de Lyon, Université Claude Bernard Lyon1, École Normale Supérieure de Lyon, 18 19 8 CNRS, UMR5276, LGL-TPE, Villeurbanne, France 20 21 For Peer Review 2 22 9 Univ. Brest, CNRS, IUEM Institut Universitaire Européen de la Mer, UMR 6538 Laboratoire 23 24 10 Géosciences Océan, Place Nicolas Copernic, 29280 Plouzané, France 25 26 11 3Institute of Earth Sciences, University of Lausanne, Géopolis, CH-1015 Lausanne, 27 28 12 Switzerland 29 30 31 13 4The Czech Academy of Sciences, Institute of Geology, Rozvojová 269, 165 00 Prague 6, 32 33 14 Czech Republic 34 35 15 5Institute of Geology and Palaeontology, Faculty of Science, Charles University, Albertov 6, 36 37 38 16 Prague, 12843, Czech Republic 39 40 17 6Département des Sciences de la Terre, Faculté des Sciences et Techniques, Université Cadi- 41 42 18 Ayyad, BP 549, 40000 Marrakesh, Morocco 43 44 * 45 19 [email protected] 46 47 20 Short running title: Large Ordovician trilobites 48 49 50 51 52 53 54 55 56 57 58 59 60 1 Cambridge University Press Geological Magazine Page 2 of 38 1 2 3 22 Abstract 4 5 6 23 Understanding variations in body-size is essential for deciphering the response of an organism 7 8 24 to its surrounding environmental conditions and its ecological adaptations. In modern 9 10 25 environments, large marine animals are mostly found in cold waters. However, numerous 11 12 26 parameters can influence body size variations other than temperatures, such as oxygenation, 13 14 15 27 nutrient availability, predation, or physical disturbances by storms. Here, we investigate 16 17 28 trilobite size variations in the Lower Ordovician Fezouata Shale deposited in a cold water 18 19 29 environment. Trilobite assemblages dominated by small- to normal-sized specimens that are 20 21 For Peer Review 22 30 few cm in length are found in proximal and intermediate settings, while those comprising 23 24 31 larger taxa more than 20cm in length are found in the most distal environment of the Fezouata 25 26 32 Shale. Drill core material from distal settings shows that sedimentary rocks hosting large 27 28 33 trilobites preserved in-situ are extensively bioturbated with a high diversity of trace fossils, 29 30 31 34 indicating that oxygen and nutrients were available in this environment. In intermediate and 32 33 35 shallow settings, bioturbation is less extensive and shallower in depth. The rarity of storm 34 35 36 events (minimal physical disturbance) and the lack of predators in deep environments in 36 37 38 37 comparison to shallower settings would have also helped trilobites attain larger body sizes. 39 40 38 This highly resolved spatial study investigating the effects of numerous biotic and abiotic 41 42 39 parameters on body size has wider implications for the understanding of size fluctuations over 43 44 45 40 geological time. 46 47 41 Keywords: Arthropod, Body-size, Paleozoic, Fezouata Shale 48 49 50 51 52 53 54 55 56 57 58 59 60 2 Cambridge University Press Page 3 of 38 Geological Magazine 1 2 3 43 1. Introduction 4 5 6 44 Considered one of the most important aspects of animal biology (Bonner, 2006), body size 7 8 9 45 results from numerous biotic and abiotic factors (Bell, 2014). Vertebrate size variations over 10 11 46 geological time have received considerable attention (Sander & Clauss, 2008; Geiger et al., 12 13 47 2013). Comparatively, marine invertebrates have been less studied (Lamsdell & Braddy, 14 15 16 48 2009; Klug et al., 2015; Sigurdsen & Hammer, 2016). For instance, it is well agreed that low 17 18 49 temperatures can be responsible of the large sizes of modern marine invertebrates (i.e. 19 20 50 Bergmann's rule; Timofeev, 2001; Moran & Woods, 2012). Nevertheless, if this was the sole 21 For Peer Review 22 51 parameter controlling body size, all taxa at high latitudes should be larger than genera found 23 24 25 52 at lower latitudes. This is rarely the case because size variations occur locally in a specific 26 27 53 paleoenvironment, owing to changes in water depth, oxygenation, predation, nutrient 28 29 54 availability or even physical disturbances caused by storm events (Saleh et al., 2018). 30 31 32 55 During the Ordovician, Morocco was part of the Gondwana margins, at high latitudes, close 33 34 56 to the South Pole. The Fezouata Shale was deposited near the Zagora region in Morocco, 35 36 57 under cold waters at the transition between two major evolutionary events: the Cambrian 37 38 39 58 Explosion and the Great Ordovician Biodiversification Event (Martin et al., 2016a). In this 40 41 59 formation, two sedimentary intervals have yielded thousands of exceptionally preserved 42 43 60 fossils belonging to different groups such as arthropods, echinoderms, mollusks, and sponges 44 45 61 (Vinther et al., 2008, 2017; Van Roy et al., 2010, 2015a; Martí Mus, 2016; Lefebvre et al., 46 47 48 62 2019). A striking feature of this formation is extreme body size fluctuations at both taxon and 49 50 63 assemblage scales between localities and even between different levels of the same locality 51 52 64 (for further details see Ebbestad, 2016; Lefebvre et al., 2016; Martin, 2016; Saleh et al., 53 54 55 65 2018). Trilobites occur in all sites from the Fezouata Shale and show a large body-size range 56 57 66 in this formation. Abundant and spectacular specimens of very large trilobites were found at 58 59 67 Ouled Slimane near the Tanskhit bridge (Rábano, 1990; Fortey, 2009; Lebrun, 2018). In this 60 3 Cambridge University Press Geological Magazine Page 4 of 38 1 2 3 68 study, the sedimentological and taphonomic contexts of levels with large trilobites from the 4 5 6 69 Fezouata Shale are elucidated, in order to contribute to the understanding of body-size 7 8 70 fluctuations in the geological record (see also Lamsdell & Braddy, 2009; Klug et al., 2015; 9 10 71 Sigurdsen & Hammer, 2016).
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