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YCRES4077_proof ■ 15 February 2019 ■ 1/26 Cretaceous Research xxx (xxxx) xxx 55 Contents lists available at ScienceDirect 56 57 Cretaceous Research 58 59 60 journal homepage: www.elsevier.com/locate/CretRes 61 62 63 64 65 1 Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz, 66 2 67 3 1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of 68 4 69 Q7 northeastern Italy 5 70 6 a, * a, f a b 71 7 Q6 Jacopo Amalfitano , Luca Giusberti , Eliana Fornaciari , Fabio Marco Dalla Vecchia , c d e 72 8 Valeria Luciani , Jürgen Kriwet , Giorgio Carnevale 73 9 a Department of Geosciences, University of Padova, Via Gradenigo 6, I-35131 Padova, Italy 74 10 b Institut Catala de Paleontologia Miquel Crusafont (ICP), Carrer de L'Escola Industrial 23, E-08201 Sabadell, Spain 75 11 c Department of Physics and Earth Sciences, University of Ferrara, Scientific and Technological Pole, Via G. Saragat 1, I-44100 Ferrara, Italy 76 12 d University of Vienna, Department of Palaeontology, Geozentrum, Althanstrasse 14, 1090 Vienna, Austria e 77 13 Department of Earth Sciences, University of Torino, Via Valperga Caluso 35, I-10125 Torino, Italy f Institute of Geosciences and Georesources, CNR, Via Gradenigo 6, 35131 Padova, Italy 78 14 79 15 80 16 article info abstract 81 17 82 18 Article history: Cretoxyrhina mantelli was a large pelagic lamniform shark geographically widespread during the Late 83 19 Received 18 September 2018 Cretaceous, and well known because of several nearly complete skeletons from the Western Interior 84 20 Received in revised form Seaway of North America. Here we report 15 partial skeletons belonging to lamniform sharks from the 85 28 December 2018 21 ‘lastame’ lithozone of the Upper Cretaceous Scaglia Rossa Formation of the Lessini Mountains (north- 86 Accepted in revised form 2 February 2019 22 eastern Italy). Seven partial but articulated skeletons include tooth sets that allow a confident attribution 87 Available online xxx 23 to Cretoxyrhina mantelli based on dental morphologies. We review the taxonomic history of C. mantelli, 88 24 evidencing that the taxon was erected by Agassiz (1835) and tracing back four of the original syntypes. Keywords: 89 25 Lamniform sharks Based on calcareous plankton biostratigraphy, the rock in which the Italian skeletal remains are 90 26 Taxonomy embedded is constrained to the middle-upper Turonian. Total length estimates of the specimens suggest 91 that the sample includes the largest specimen of Cretoxyrhina mantelli (615e650 cm estimated total 27 Upper Cretaceous 92 28 Paleobiology length) known to date. The placoid scale morphology indicates that C. mantelli most likely was a fast Taphonomy 93 29 swimmer with a similar ecology as the extant white shark, Carcharodon carcharias. The associated skeletal elements of the specimens represent large chondrichthyan deadfalls and the cadavers decayed 94 30 on the seafloor where they remained exposed for several months, as indicated by bioerosional traces, 95 31 some of which are interpreted as a product of bone-eating worm activities and other bioerosional traces 96 32 with Gastrochaenolites-like structure. The Cretoxyrhina mantelli remains described herein provide new 97 33 information about the ʻlastameʼ vertebrate assemblage, which seemingly was strongly dominated by 98 34 chondrichthyans, especially lamniform sharks. 99 35 © 2019 Elsevier Ltd. All rights reserved. 100 36 101 37 102 38 103 39 104 40 105 Q1 1. Introduction reported from many localities all around the world (Africa, Asia, 41 106 Europe, North and South America; Cappetta, 2012). Some nearly 42 107 The ʻginsuʼ shark, Cretoxyrhina mantelli (Agassiz), is probably complete skeletons were discovered in the Smoky Hill Chalk 43 108 one of the best known and ecologically relevant top predators of Member of the Niobrara Chalk in the Western Interior Seaway of 44 109 the Late Cretaceous seas. Isolated teeth of this shark have been North America (Bourdon, Everhart, 2011; Shimada, 1993a,b, 45 110 1994aec, 1997aee, 2008; Shimada et al., 2006). These skeletons 46 111 indicate that Cretoxyrhina mantelli had a general morphology close 47 112 to that of the white shark (Carcharodon carcharias Linnaeus, 1758) 48 113 * Corresponding author. (Shimada, 1997b; Shimada et al., 2006), being similar or even larger 49 E-mail addresses: jacopo.amalfi[email protected] (J. Amalfitano), luca. 114 in size. 50 [email protected] (L. Giusberti), [email protected] (E. Fornaciari), fabio. 115 Isolated teeth are so far the most common remains of C. mantelli 51 [email protected] (F.M. Dalla Vecchia), [email protected] (V. Luciani), juergen. 116 [email protected] (J. Kriwet), [email protected] (G. Carnevale). reported from Italy (e.g., Bassani, 1876; D'Erasmo, 1922), although 52 117 53 https://doi.org/10.1016/j.cretres.2019.02.003 118 54 0195-6671/© 2019 Elsevier Ltd. All rights reserved. 119 Please cite this article as: Amalfitano, J et al., Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz,1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy, Cretaceous Research, https://doi.org/10.1016/j.cretres.2019.02.003 YCRES4077_proof ■ 15 February 2019 ■ 2/26 2 J. Amalfitano et al. / Cretaceous Research xxx (xxxx) xxx 1 some partial skeletons were found in the Scaglia Rossa Formation of The ʻlastameʼ (ʻlithozoneʼ 2 of the Scaglia Rossa Formation of 66 2 Lessini Mountains (Verona Province, Veneto Region) during the Lozar and Grosso, 1997) crops out in the surroundings of S. Anna 67 3 1970s (Amalfitano et al., 2017b,c; Cigala Fulgosi et al., 1980; Dalla d'Alfaedo village (Verona Province) (Fig. 1), on Loffa Mount (west- 68 4 Vecchia et al., 2005; Ginevra et al., 2000). The majority of these ern Lessini Mountains). It consists of a 7-8 m-thick package of 69 5 skeletal remains consist of teeth associated with calcified vertebral limestone that is subdivided into 72 flaser-nodular whitish to 70 6 centra and fragments of calcified cranial cartilage; in some cases, reddish layers, rich in planktonic foraminifera and calcareous 71 7 they include also placoid scales. nannofossils (Amalfitano et al., 2017b). Such layers are laterally 72 8 The goal of this paper is to provide an overview of all the skeletal continuous with a thickness ranging from 4 to 35 cm and are 73 9 remains of C. mantelli discovered in northeastern Italy, and to separated by thin, dark red and shaly layers (marls and clays; 74 10 discuss their paleobiological, stratigraphic and taphonomical im- Amalfitano et al., 2017b). According to Lozar, Grosso (1997), the 75 11 plications. The historical record of the genus Cretoxyrhina in ʻlastameʼ spans from the lower Turonian to the lower Santonian, 76 12 northeastern Italy is also discussed and we propose a solution to while other authors assigned a Turonian-Coniacian age to it (e.g., 77 13 the nomenclatural conundrum of the exact date of the original Cigala Fulgosi et al., 1980). A stratigraphic revision of the whole 78 14 erection of Cretoxyrhina mantelli. lithozone, however, is currently in progress. The macrofossils of the 79 15 ʻlastameʼ include several remains of large marine vertebrates 80 fi 16 2. Geological background (chondrichthyans, bony shes, marine turtles, and mosasaurs; 81 17 Amadori et al., 2019; Amalfitano et al., 2017a-c; Capellini, 1884; 82 fi 18 All of the articulated skeletons of Cretoxyrhina mantelli discov- Chesi, Del no, 2007; Dalla Vecchia et al., 2005; Cigala Fulgosi et al., 83 19 84 ered in northeastern Italy come from the Scaglia Rossa Formation, 1980; Palci et al., 2013). ʻ ʼ 20 which crops out and is extensively quarried in several localities of The Pietra di Castellavazzo crops out in the surroundings of 85 21 86 the western Lessini Mountains (Verona Province; Massari and Castellavazzo village (near Longarone, Belluno Province) in the Piave ʻ ʼ e 22 Savazzi, 1981; Amalfitano et al., 2017a; Ginevra et al., 2000) and Valley (Fig. 1). The facies resembles the lastame , being a 6 7m 87 23 88 the Piave Valley (Belluno Province; Colombara, 2013). Quarries are thick section of micritic nodular limestones, with laterally contin- 24 opened into two peculiar lithofacies (one in the Lessini Mountains uous layers ranging in thickness from a few cm to 40 cm, separated 89 25 90 and the other in the Piave Valley) that yielded the totality of the by dark red clay interlayers (Trevisani, 2009, 2011). Moreover, the ʻ ʼ 26 skeletons. They are named ʻlastameʼ (or ʻPietra di Prunʼ) and ʻPietra Pietra di Castellavazzo is subdivided into two sub-units, the lower 91 27 92 di Castellavazzoʼ. reddish portion and the upper grey-greenish portion (Trevisani, 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 122 58 123 59 124 60 125 61 126 62 127 63 128 64 Fig. 1. Location of the sites. The sites that yielded the skeletal remains of Cretoxyrhina mantelli described herein (Sant'Anna D'Alfaedo, Verona province) and Castellavazzo, near 129 65 Longarone (Belluno province) are the only other site that yielded Cretoxyrhina mantelli skeletons. Scale bar ¼ 100 km [single-column width]. 130 Please cite this article as: Amalfitano, J et al., Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz,1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy, Cretaceous Research, https://doi.org/10.1016/j.cretres.2019.02.003 YCRES4077_proof ■ 15 February 2019 ■ 3/26 J.