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Adnet-Et-Al 2020 Palaeontologi Palaeontologia Electronica palaeo-electronica.org Diversity and renewal of tropical elasmobranchs around the Middle Eocene Climatic Optimum (MECO) in North Africa: New data from the lagoonal deposits of Djebel el Kébar, Central Tunisia Sylvain Adnet, Laurent Marivaux, Henri Cappetta, Anne-Lise Charruault, El Mabrouk Essid, Suzanne Jiquel, Hayet Khayati Ammar, Bernard Marandat, Wissem Marzougui, Gilles Merzeraud, Rim Temani, Monique Vianey-Liaud, and Rodolphe Tabuce ABSTRACT Bulk sampling of an indurated glauconic sandstone horizon from Djebel el Kébar, Central Tunisia, yielded a moderately diversified assemblage of elasmobranchs (sharks and rays), dating from the late Middle Eocene (mid Bartonian). Here, 10 new taxa (nine new species and three new genera) are described among a total diversity of 33 species. These new taxa highlight the rise of mid-large predator lineages of modern nearshore seas and fill a substantial gap in the sporadic fossil record of the late Middle Eocene. Comparisons of this assemblage with other Middle-Late Eocene faunas from the Tethys seaway indicate that this Tunisian fauna shares more similarities with Late Eocene than with early Middle Eocene assemblages. This suggests that the major shift observed in tropical elasmobranch communities from the Late Eocene, with for instance the appearance of large Carcharhinids replacing Lamniforms, began during the late Middle Eocene. Such a biotic shift could be linked to the warm conditions recorded ca. 41 Ma, known as the Middle Eocene Climatic Optimum (MECO), which was characterized by a short temperature increase of all marine waters. Sylvain Adnet. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Laurent Marivaux. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] http://zoobank.org/B6B8E985-F1CF-4C10-BB00-602E5BF36C1C Adnet, Sylvain, Marivaux, Laurent, Cappetta, Henri, Charruault, Anne-Lise, Essid, El Mabrouk, Jiquel, Suzanne, Ammar, Hayet Khayati, Marandat, Bernard, Marzougui, Wissem, Merzeraud, Gilles, Temani, Rim, Vianey-Liaud, Monique, and Tabuce, Rodolphe. 2020. Diversity and renewal of tropical elasmobranchs around the Middle Eocene Climatic Optimum (MECO) in North Africa: New data from the lagoonal deposits of Djebel el Kébar, Central Tunisia. Palaeontologia Electronica, 23(2):a38. https://doi.org/10.26879/ 1085 palaeo-electronica.org/content/2020/3110-meco-elasmobranchs Copyright: August 2020 Society of Vertebrate Paleontology. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. creativecommons.org/licenses/by/4.0 ADNET ET AL.: MECO ELASMOBRANCHS Henri Cappetta. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Anne-Lise Charruault. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. anne- [email protected] El Mabrouk Essid. Office National des Mines (ONM), 24 rue 8601, 2035 La Charguia, Tunis BP: 215, 1080 Tunis, Tunisia. [email protected] Suzanne Jiquel. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Hayet Khayati Ammar. Office National des Mines (ONM), 24 rue 8601, 2035 La Charguia, Tunis BP: 215, 1080 Tunis, Tunisia. [email protected] Bernard Marandat. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Wissem Marzougui. Office National des Mines (ONM), 24 rue 8601, 2035 La Charguia, Tunis BP: 215, 1080 Tunis, Tunisia. [email protected] Gilles Merzeraud. Géosciences Montpellier (UMR 5243, CNRS/UM/Université des Antilles), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Rim Temani. Office National des Mines (ONM), 24 rue 8601, 2035 La Charguia, Tunis BP: 215, 1080 Tunis, Tunisia. [email protected] Monique Vianey-Liaud. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. monique.vianey- [email protected] Rodolphe Tabuce. Institut des Sciences de l’Evolution (UMR5554, CNRS, UM, IRD, EPHE), Université de Montpellier, Place E. Bataillon, 34095 Montpellier Cedex 05, France. [email protected] Keywords: Bartonian; faunal turnover; new species; new genera; elasmobranchs Submission: 7 April 2020. Acceptance: 17 July 2020. INTRODUCTION present on the Antarctic continent (Miller et al., 2005), while sea ice may have been present in the The Eocene was a crucial time in the Earth's Arctic Ocean (Stickley et al., 2009). Cenozoic climate evolution. Deep-ocean and high- In the tropical realm of the western Tethys latitude palaeotemperature proxies demonstrate seaway (prefiguring the paleomediterranean sea that the Eocene epoch encompasses the warmest before its Neogene closure), there is no evidence interval of the past 66 million years. The highest of major change in inshore marine depositions temperatures of the Cenozoic were reached during during the Early Eocene and early Middle Eocene. the Paleocene Eocene Thermal Maximum (PETM, Combined with a high sea-level, the warm and ~56 Ma) and the Early Eocene Climatic Optimum humid climate at this time may have favoured the (EECO, ~53-49 Ma) and were followed by long- local formation of numerous fossiliferous phos- term cooling, which continued through the Middle phate cores (dated to Early-early Middle Eocene) and Late Eocene (e.g., Zachos et al., 2001, 2008; along the Western and North African coasts (e.g., Katz et al., 2008; Bijl et al., 2010; Westerhold et al., Notholt et al., 1989) by increasing the rate of conti- 2018; Hollis et al., 2019). During this global cool- nental weathering, and thereby increasing the ing, world lands and seas were in the so-called amount of phosphate transported to the sea “doubthouse” climate state, between the “green- (Follmi, 1996; Swezey, 2009). This kind of deposit house” conditions around the EECO and the “ice- is well-known by vertebrate palaeontogists house” regime that started with the development of because it optimizes the well-preservation of fossil the Antarctic continental ice sheet in the earliest apatite (biogenic calcium triphosphate) that consti- Oligocene (e.g., Miller et al., 1987; Zachos et al., tutes the main mineral tissue of vertebrate bones 1996; DeConto and Pollard, 2003; Coxall et al., and teeth. This is particularly the case with elasmo- 2005; Miller et al., 2005; Lear et al., 2008, Liu et al., branchs, a group for which most of the main fossil- 2009). During these long “doubthouse” conditions iferous deposits are located in northern Africa, from (= Eocene), ephemeral ice sheets may have been 2 PALAEO-ELECTRONICA.ORG Morocco to Tunisia, during the early Paleogene This relatively long transition to globally cooler (Arambourg, 1952; Noubhani and Cappetta, 1997). climates was, however, interrupted by transient The quantity of phosphate in marine deposition warming events, the most prolonged and intense of (and correlatively fossil-bearing levels with well which was the “Middle Eocene Climatic Optimum” preserved elasmobranch teeth) decreases along (MECO; e.g., Bohaty and Zachos, 2003). The the inshore since the late Middle Eocene. In this MECO interrupted the cooling trend of the middle- context, most of the Late Eocene fossil-bearing late Eocene at ~ 40.6–40.0 Ma (Bohaty et al., localities with elasmobranchs are found to be 2009), during the Bartonian. First identified in sedi- related to coastal sand body complexes (Adnet et ment cores as a transient oxygen isotopic excur- al., 2007, 2010, 2011; Underwood et al., 2011). sion (~ −1.0‰; Bohaty and Zachos, 2003), the Among them, some well-exposed coastal com- MECO was recognized and more precisely dated plexes in the Western Desert of Egypt (e.g., Wadi in isotopic records from different areas. At warming al Hitan) have been sometimes interpreted to peak, the δ18O records indicate warming of ~ 4–6 reflect such global cooling and ephemeral eustatic °C of both surface and deep waters (Bohaty et al., sea level decrease (possibly related to merging of 2009; Edgar et al., 2010), with sea-surface tem- ice?) during the Late Eocene (Peters et al., 2009, peratures ranging as high as ~31 °C in the tropical 2010). However, such interpretations are always realm (Cramwinckel et al., 2018; see Figure 1). severely debated (e.g., Underwood et al., 2012; Marine biotic response to the MECO has so far Gingerich et al., 2012; King et al., 2014), and local only been studied among plankton or benthic fora- tectonic events might play an important role in minifera, from the Tethys seaway (Luciani et al., these past sedimentary cycles. 2010; Toffanin et al., 2011; Witkowski et al., 2012; Boscolo Galazo et al., 2013). These studies have FIGURE 1. Paleotemperatures (ice-free deep-ocean T°/
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