Accepted Manuscript Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz, 1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy Jacopo Amalfitano, Luca Giusberti, Eliana Fornaciari, Fabio Marco Dalla Vecchia, Valeria Luciani, Jürgen Kriwet, Giorgio Carnevale PII: S0195-6671(18)30378-1 DOI: https://doi.org/10.1016/j.cretres.2019.02.003 Reference: YCRES 4077 To appear in: Cretaceous Research Received Date: 18 September 2018 Revised Date: 28 December 2018 Accepted Date: 2 February 2019 Please cite this article as: Amalfitano, J., Giusberti, L., Fornaciari, E., Dalla Vecchia, F.M., Luciani, V., Kriwet, J., Carnevale, G., 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. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz, 1835) 2 (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy 3 4 Jacopo Amalfitano1*, Luca Giusberti1,6, Eliana Fornaciari1, Fabio Marco Dalla Vecchia2, Valeria 5 Luciani3, Jürgen Kriwet4 & Giorgio Carnevale5 PT 6 1. Department of Geosciences, University of Padova, Via Gradenigo 6, I-35131 Padova, Italy, e-mail: jacopo.amalfitano@phd.unipd.it, RI 7 luca.giusberti@unipd.it, eliana.fornaciari@unipd.it 8 2. Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer de l'Escola Industrial 23, E-08201 Sabadell, Spain. e-mail: SC 9 fabio.dallavecchia@icp.cat 10 3. Department of Physics and Earth Sciences, University of Ferrara, Scientific and Technological Pole, Via G. Saragat 1, I-44100 Ferrara, Italy, e- 11 mail: lcv@unife.it U 12 4. University of Vienna, Department of Palaeontology, Geozentrum, Althanstrasse 14, 1090 Vienna, Austria, e-mail: juergen.kriwet@univie.ac.at AN 13 5. Department of Earth Sciences, University of Torino, Via Valperga Caluso 35, I-10125 Torino, Italy, e-mail: giorgio.carnevale@unito.it 14 6. Institute of Geosciences and Georesources, CNR, Via Gradenigo 6, 35131 Padova, Italy. 15 *Corresponding author M 16 17 Keywords D 18 Lamniform sharks; Taxonomy; Upper Cretaceous; Paleobiology; Taphonomy TE 19 20 Abstract EP 21 C 22 Cretoxyrhina mantelli was a large pelagic lamniform shark geographically widespread during the AC 23 Late Cretaceous, and well known because of several nearly complete skeletons from the Western 24 Interior Seaway of North America. Here we report 15 partial skeletons belonging to lamniform 25 sharks from the ‘lastame’ lithozone of the Upper Cretaceous Scaglia Rossa Formation of the Lessini 26 Mountains (northeastern Italy). Seven partial but articulated skeletons include tooth sets that allow a 27 confident attribution to Cretoxyrhina mantelli based on dental morphologies. We review the 28 taxonomic history of C. mantelli, evidencing that the taxon was erected by Agassiz (1835) and 29 tracing back four of the original syntypes. Based on calcareous plankton biostratigraphy, the rock in ACCEPTED MANUSCRIPT 30 which the Italian skeletal remains are embedded is constrained to the middle-upper Turonian. Total 31 length estimates of the specimens suggest that the sample includes the largest specimen of 32 Cretoxyrhina mantelli (615-650 cm estimated total lenght) known to date. The placoid scale 33 morphology indicates that C. mantelli most likely was a fast swimmer with a similar ecology as the 34 extant white shark, Carcharodon carcharias. The associated skeletal elements of the specimens PT 35 represent large chondrichthyan deadfalls and the cadavers decayed on the seafloor where they RI 36 remained exposed for several months, as indicated by bioerosional traces, some of which are 37 interpreted as a product of bone-eating worm activities and other bioerosional traces with SC 38 Gastrochaenolites-like structure. The Cretoxyrhina mantelli remains described herein provide new 39 information about the ʻlastameʼ vertebrate assemblage, which seemingly was strongly dominated by 40 chondrichthyans, especially lamniform sharks. U AN 41 M 42 Introduction 43 D 44 The ʻginsuʼ shark, Cretoxyrhina mantelli (Agassiz), is probably one of the best known and TE 45 ecologically relevant top predators of the Late Cretaceous seas. Isolated teeth of this shark have 46 been reported from many localities all around the world (Africa, Asia, Europe, North and South EP 47 America; Cappetta, 2012). Some nearly complete skeletons were discovered in the Smoky Hill C 48 Chalk Member of the Niobrara Chalk in the Western Interior Seaway of North America (Bourdon, AC 49 Everhart, 2011; Shimada, 1993a,b, 1994a-c, 1997a-e, 2008; Shimada et al., 2006). These skeletons 50 indicate that Cretoxyrhina mantelli had a general morphology close to that of the white shark 51 (Carcharodon carcharias Linnaeus, 1758) (Shimada, 1997b; Shimada et al., 2006), being similar or 52 even larger in size. 53 Isolated teeth are so far the most common remains of C. mantelli reported from Italy (e.g., 54 Bassani, 1876; D’Erasmo, 1922), although some partial skeletons were found in the Scaglia Rossa 55 Formation of Lessini Mountains (Verona Province, Veneto Region) during the 1970s (Amalfitano ACCEPTED MANUSCRIPT 56 et al., 2017b,c; Cigala Fulgosi et al., 1980; Dalla Vecchia et al., 2005; Ginevra et al., 2000). The 57 majority of these skeletal remains consist of teeth associated with calcified vertebral centra and 58 fragments of calcified cranial cartilage; in some cases, they include also placoid scales. 59 The goal of this paper is to provide an overview of all the skeletal remains of C. mantelli 60 discovered in northeastern Italy, and to discuss their paleobiological, stratigraphic and taphonomical PT 61 implications. The historical record of the genus Cretoxyrhina in northeastern Italy is also discussed RI 62 and we propose a solution to the nomenclatural conundrum of the exact date of the original erection 63 of Cretoxyrhina mantelli. SC 64 65 Geological background 66 U AN 67 All of the articulated skeletons of Cretoxyrhina mantelli discovered in northeastern Italy M 68 come from the Scaglia Rossa Formation, which crops out and is extensively quarried in several 69 localities of the western Lessini Mountains (Verona Province; Massari and Savazzi, 1981; D 70 Amalfitano et al., 2017a; Ginevra et al., 2000) and the Piave Valley (Belluno Province; Colombara, TE 71 2013). Quarries are opened into two peculiar lithofacies (one in the Lessini Mountains and the other 72 in the Piave Valley) that yielded the totality of the skeletons. They are named ʻlastameʼ (or ʻPietra EP 73 di Prunʼ) and ʻPietra di Castellavazzoʼ. C 74 The ʻlastameʼ (ʻlithozoneʼ 2 of the Scaglia Rossa Formation of Lozar and Grosso, 1997) AC 75 crops out in the surroundings of S. Anna d'Alfaedo village (Verona Province) (Fig. 1), on Loffa 76 Mount (western Lessini Mountains). It consists of a 7-8 m-thick package of limestone that is 77 subdivided into 72 flaser-nodular whitish to reddish layers, rich in planktonic foraminifera and 78 calcareous nannofossils (Amalfitano et al., 2017b). Such layers are laterally continuous with a 79 thickness ranging from 4 to 35 cm and are separated by thin, dark red and shaly layers (marls and 80 clays; Amalfitano et al., 2017b). According to Lozar, Grosso (1997), the ʻlastameʼ spans from the 81 lower Turonian to the lower Santonian, while other authors assigned a Turonian-Coniacian age to it ACCEPTED MANUSCRIPT 82 (e.g., Cigala Fulgosi et al., 1980). A stratigraphic revision of the whole lithozone, however, is 83 currently in progress. The macrofossils of the ʻlastameʼ include several remains of large marine 84 vertebrates (chondrichthyans, bony fishes, marine turtles, and mosasaurs; Amadori et al., 2019; 85 Amalfitano et al., 2017a-c; Capellini, 1884; Chesi, Delfino, 2007; Dalla Vecchia et al., 2005; Cigala 86 Fulgosi et al, 1980; Palci et al., 2013). PT 87 The ʻPietra di Castellavazzoʼ crops out in the surroundings of Castellavazzo village (near RI 88 Longarone, Belluno Province) in the Piave Valley (Fig. 1). The facies resembles the ʻlastameʼ, 89 being a 6-7 m thick section of micritic nodular limestones, with laterally continuous layers ranging SC 90 in thickness from a few cm to 40 cm, separated by dark red clay interlayers (Trevisani, 2009, 2011). 91 Moreover, the ʻPietra di Castellavazzoʼ is subdivided into two sub-units, the lower reddish portion 92 U and the upper grey-greenish portion (Trevisani, 2009, 2011). Some authors referred the ʻPietra di AN 93 Castellavazzoʼ to the early-middle Coniacian based on correlation with the ʻlastameʼ (Coleselli et M 94 al., 1997; Trevisani, 2011), pending revisions, which are currently being carried out (Federico Fanti, 95 pers. comm.). The fossil content of the ‘Pietra di Castellavazzo’, is similar to that of the ʻlastameʼ, D 96 including chondrichthyans and bony fishes, mostly represented by isolated teeth, and occasionally TE 97 by more complete remains (e.g., Bassani, 1876, 1888; Trevisani, 2009, 2011). 98 EP 99 Historical background C 100 AC 101 The first report of isolated teeth of Cretoxyrhina mantelli from the Upper Cretaceous Scaglia 102 Rossa Formation of the Lessini Mountains was that by Bassani (1876). However, teeth of this 103 species have been found in northern Italy from at least the first half of the 19th century, as testified 104 by the presence of four specimens in the collection of Tommaso Antonio Catullo (1782-1869) 105 housed in the Museum di Geology and Paleontology of the University of Padova (pers. obs.; see 106 Tab. 1 for details and D'Erasmo, 1922; Fig. 2 and supplementary material Fig. A.1θ-λ). Other teeth 107 from the ‘Pietra di Castellavazzo’ were initially misidentified as Lamna (=Sphenodus) longidens ACCEPTED MANUSCRIPT 108 (Bassani, 1876; D'Erasmo, 1922). De Zigno (1883: p. 9) reported several localities of the Scaglia 109 Rossa Formation yielding isolated teeth of C. mantelli, including the Valpolicella, Agno Valley, 110 Follina Hills, and the surroundings of the towns of Feltre and Belluno. 111 The first Italian specimen of C. mantelli with articulated vertebral centra and associated 112 teeth was recovered in 1878 from the ‘Pietra di Castellavazzo’ at the Olantreghe quarry near PT 113 Longarone (Bassani, 1888). The fossil, consisting of 122 vertebral centra and 55 teeth, is housed in RI 114 the collection of the Museum of Natural History, University of Pavia and is currently under review 115 (Paolo Guaschi and Federico Fanti, pers. comm.). SC 116 D’Erasmo (1922) published the last synopsis on the C. mantelli record from the Veneto 117 Region as part of a review about fossil fishes of northeastern Italy. Since then, studies on this 118 U lamniform shark in Italy have been neglected, despite the recovery of new relevant specimens AN 119 during the second half of the 20th century. In particular, well-preserved vertebral column segments, M 120 often associated with teeth, were discovered during quarrying into the ʻlastameʼ in the Lessini 121 Mountains. These important findings were only briefly mentioned in short notes (e.g., Cigala D 122 Fulgosi et al., 1980), remaining virtually unknown to the scientific community. The oldest TE 123 stratigraphic record of Cretoxyrhina mantelli from the study area reported in the literature refers to 124 isolated teeth coming from the upper Cenomanian Bonarelli Level (Dalla Vecchia et al., 2005; EP 125 Sorbini, 1976). 126 C AC 127 Material and methods 128 129 Institutional abbreviations 130 CE.A.S.C.: Centro di Analisi e Servizi Per la Certificazione, University of Padova, Padova, Italy; 131 FHSM: Sternberg Museum of Natural History, Hays, Kansas, USA; NHM: The Natural History 132 Museum, London, UK; MCSNV: Museo Civico di Storia Naturale di Verona (Civic Museum of 133 Natural History), Verona, Italy; MGC: Museo Geopaleontologico di Camposilvano ACCEPTED MANUSCRIPT 134 (Geopaleontological Museum), Camposilvano, Italy; MGP-PD: Museo di Geologia e Paleontologia 135 (Museum of Geology and Paleontology), University of Padova, Padova, Italy; MPPSA: Museo 136 Preistorico e Paleontologico di S. Anna d'Alfaedo (Prehistoric and Paleontological Museum), S. 137 Anna d'Alfaedo, Italy. 138 PT 139 Material RI 140 The material reported herein is part of the collections of the Prehistoric and Paleontological 141 Museum of S. Anna d’Alfaedo, the Geopaleontogical Museum of Camposilvano, the Museum of SC 142 Geology and Paleontology, University of Padova, and the Civic Museum of Natural History of 143 Verona. Most of the specimens come from the ʻlastameʼ quarries of Loffa Mount. Limestone slabs 144 U of different size contain calcified vertebral centra, teeth, and, in some cases, fragments of calcified AN 145 cranial cartilage and placoid scales. The isolated teeth come from several localities of Veneto M 146 Region. In Tab. 1, a detailed list of the examined material is provided, reporting the specimen 147 number and its repository, a concise description of its state of preservation, and its locality of D 148 provenance. TE 149 150 Methods EP 151 The specimens were documented with different photographic techniques depending on the 152 size of the slabs (e.g., MPPSA IGVR 36371 and MPPSA IGVR 45305 were photographed with the C AC 153 photogrammetric technique and then the images were elaborated to obtain an orthophoto) using 154 digital cameras (a Canon PowerShot SX720 HS and a Fuji X-E1 mounting 18-55 mm lens). Images 155 and interpretative drawings of the specimens were obtained using the freewares GIMP (v. 2.10.2) 156 and ImageJ (v. 1.47). The synonymy list and open nomenclature follow the standards proposed by 157 Matthews (1973), Bengston (1988) and Sigovini et al. (2016). 158 We refer to papers concerning Cretoxyrhina dentition (e.g., Bourdon, Everhart, 2011; 159 Cappetta, 2012; Shimada, 1997a, 2002) for tooth position identification, general dental characters ACCEPTED MANUSCRIPT 160 and technical terms. The identification of tooth position in lamniform sharks is a debated matter 161 related to the tooth rows arrangement on the palatoquadrate and Meckel’s cartilage, as well as to the 162 structure of these cranial elements (e.g., presence/absence of intermediate bar sensu Shimada, 2002 163 and the two hollows present on each side of the jaw sensu Siverson, 1999). ‘Intermediate teeth’ is 164 probably the most controversial term. Siverson (1999) and Shimada (2002) proposed two PT 165 alternative indications, based on homologous/non-homologous tooth positions. Later on, Cook et al. RI 166 (2011: p. 9-10) summarized the arguments discussed in the debate. We refer to Cook et al. (2011) 167 and Kriwet et al. (2015). We follow Shimada (1997d) for general skeletal anatomy and Newbrey et SC 168 al. (2015) for the vertebral morphology of C. mantelli. The terminology used for placoid scales 169 follows Reif (1985). 170 U As for other fossil chondrichthyans (see Cappetta, 1987, 2012), the diagnosis of AN 171 Cretoxyrhina mantelli is essentially based on dental characters. Herein, we propose an emended M 172 diagnosis of C. mantelli that integrates all of the dental characters reported by Bourdon, Everhart 173 (2011), Cappetta (2012) and Shimada (1997a), in order to complete and better formalize it. D 174 Significant measurements for teeth and vertebral centra were taken using the image analysis TE 175 software ImageJ or directly on the specimens; in particular, the maximum diameter of the largest 176 vertebral centra (posterior precaudal centra, especially mid-trunk, when available), the maximum EP 177 and minimum height of the teeth and the maximum height of the crown measured on the labial side 178 (see supplementary material for further measurements, Tabs. A.1.-A.2). Maximum tooth height was C AC 179 measured in the anterior teeth when possible. We used the equations proposed by Shimada (2008) 180 to estimate the maximum body size of the specimens. The count of incremental bands on vertebral 181 centra for individual age estimation was performed on the more complete preserved centra of the 182 two better preserved specimens (MPPSA-IGVR 36371 and 45305). It was not possible to perform 183 any vertebral section because all the vertebral centra are embedded in the limestone slabs and could 184 not be removed. Only a single vertebra from the specimen MGP-PD 31960 was removed and 185 sectioned, although it did not provide any significant result because of diagenetic alteration. Small ACCEPTED MANUSCRIPT 186 limestone nodules detached from selected specimens (MPPSA-IGVR 36371, 45305 and MGC- 187 IGVR 81375-81376) were processed for micropaleontological analyses. Some nodules were 188 processed following the cold acetolysis method of Lirer (2000) for isolating planktonic foraminifera 189 and placoid scales of the >63 µm fraction, whereas some other nodules were utilized for thin 190 sections and for preparation of smear slides for calcareous nannofossil analysis. Placoid scales were PT 191 picked and counted from the residues >63 µm from selected specimens (MPPSA-IGVR 36371 and RI 192 MGC-IGVR 81375-81376). Representative specimens of the main morphologies of placoid scales 193 were selected and imaged using a SEM (JSM Jeol 6490) at the CE.A.S.C. structure of Università SC 194 degli Studi di Padova. Additionally, we described bioerosional structures that occur on two 195 specimens, MPPSA IGVR 36371 and MGP-PD 31960, following the ichnotaxonomy of Pirrone et 196 al. (2014). U AN 197 M 198 Systematic paleontology 199 D 200 Class Chondrichthyes Huxley, 1880 TE 201 Subclass Elasmobranchii Bonaparte, 1838 202 Cohort Euselachii Hay, 1902 EP 203 Subcohort Neoselachii Compagno, 1977 C 204 Superorder Galeomorphii Compagno, 1973 AC 205 Order Lamniformes Berg, 1937 206 Family Cretoxyrhinidae Glikman, 1958 207 Genus Cretoxyrhina Glikman, 1958 208 209 Type species: Oxyrhina mantelli (Agassiz, 1835) from the Chalk of the Lewes area, East Sussex, 210 England (UK). 211 ACCEPTED MANUSCRIPT 212 Remarks. The taxonomic status of the genus Cretoxyrhina was discussed by Siverson et al. (2013). 213 The authors rejected Zhelzko's (2000) synonymy of Cretoxyrhina with Pseudoisurus. Glikman 214 (1958) erected the new genus Cretoxyrhina with Oxyrhina mantelli (Agassiz) as type species 215 (Siverson, 1996; Siverson et al., 2013). Later, Glikman (1964) replaced Oxyrhina mantelli with 216 Isurus denticulatus (Glickman, 1957) as type species of the genus Cretoxyrhina without PT 217 explanation. As implied by Siverson (1996) and Siverson et al (2013), this represents an invalid RI 218 taxonomic amendment (see Ride et al., 2000, Art. 68.2). The genus Cretoxyrhina currently includes 219 four species: C. mantelli, C. denticulata, C. vraconensis and C. agassizensis. SC 220 221 Cretoxyrhina mantelli (Agassiz, 1835) 222 U Figs. 2-12, 15, supplementary material Figs. A.1α-δ, θ-λ AN 223 M 224 (selected synonyms) 225 D 226 v1822 Squalus mustelus - Mantell, p. 226, pl. 32, fig. 11. TE 227 v1822 Squalus zygaena? - Mantell, p. 227, pl. 32, figs. 4, 7, 8, 10, 26, 28. 228 †1835 Lamna Mantellii Ag. - Agassiz, p. 54. EP 229 p.1838 Oxyrhina Mantellii Agass. - Agassiz, pl. 33, figs. 1-5,7-9 (non 6). 230 1843 Oxyrhina Mantellii Agass. - Agassiz, p. 280. C AC 231 1876 L. (Sphenodus) longidens Agass. - Bassani, p. 296. 232 1876 Ox. Mantellii Agass. - Bassani, p. 298. 233 1876 Ox. subinflata Agass. - Bassani, p. 299. 234 1886 Oxyrhina Mantellii Agassiz - Bassani, p. 144, figs. 1-5. 235 v1886 Oxyrhina subinflata Agassiz - Bassani, p. 145. 236 1888 Oxyrhina Mantellii Agassiz - Bassani, p. 1, pl. 1-3. 237 1889 Oxyrhina mantelli Agassiz - Woodward, p. 376, pl. 17, figs. 9-21. ACCEPTED MANUSCRIPT 238 1894 Oxyrhina mantelli - Eastman, p. 151, pl. 16-18. 239 1911 Oxyrhina mantelli Agassiz - Woodward, p. 202, text-fig. 60, 61, pl. 43, figs. 10-15 (cum 240 syn.). 241 v1922 Oxyrhina Mantelli Ag. - D'Erasmo, p. 36, pl. 3, figs. 1-3 (cum syn.). 242 v1922 Oxyrhina cfr. Mantelli Ag. - D'Erasmo, p. 37, pl. 3, figs. 4-6. PT 243 1957 Cretoxyrhina mantelli - Glickman, p. 569. RI 244 v1976 Isurus mantelli (Agassiz) 1843 - Sorbini, p. 481, pl. 1. 245 1977 Cretoxyrhina mantelli (Agassiz L.) 1843 - Herman, p. 219, pl. 9, fig. 6 (cum syn.). SC 246 v1978 Isurus mantelli - Sorbini, p. 69, fig. 9. 247 v1980 Cretoxyrhina mantelli - Cigala Fulgosi et al., p. 126, text-fig. p. 125 (cum syn.). 248 U v1984 Isurus mantelli - Aspes, Zorzin, p. 14, text-figure p. 15. AN 249 1987 Cretoxyrhina mantelli (Agassiz, 1843) - Cappetta, p. 99, figs. 87e-i. M 250 1992 Cretoxyrhina mantelli (Agassiz, 1843) - Siverson, p. 526, pl. 1, figs. 18-19 (cum syn.). 251 1993 Cretoxyrhina mantelli (Agassiz, 1843) - Welton, Farish, p. 101, text-figs. 1-12 p. 101, 1-2 p. D 252 102. TE 253 1996 Cretoxyrhina mantelli (Agassiz, 1843) - Siversson, p. 819, pl. 1, figs. 1-18. 254 1997a Cretoxyrhina mantelli (Agassiz, 1843) - Shimada, p. 269, figs. 1-10. EP 255 1997b Cretoxyrhina mantelli (Agassiz, 1843) - Shimada, p. 926, figs. 1-2, 5-6. 256 1997c Cretoxyrhina mantelli - Shimada, p. 233, fig. 1. C AC 257 1997d Cretoxyrhina mantelli (Agassiz, 1843) - Shimada, p. 642, figs. 1-11. 258 1997e Cretoxyrhina mantelli (Agassiz, 1843) - Shimada, p. 139, fig. 1. 259 v2000 Cretoxirina mantelli (sic) - Ginevra et al., p. 31, fig. 6.1. 260 v2000 Denti di squalo - Ginevra et al., p. 33, fig. 6.3. 261 v2001 Cretoxyrhina mantelli - Zorzin, p. 98 (including text-fig.). 262 v2003 Isurus mantelli (Agassiz) - Astolfi, Colombara, p. 144, fig. 205. 263 2005 Cretoxyrhina mantelli (Agassiz, 1843) - Siverson, Lindgren, p. 303, fig. 2. ACCEPTED MANUSCRIPT 264 v2005 Cretoxyrhina mantelli - Dalla Vecchia et al., p. 108, fig. 81. 265 2006 Cretoxyrhina mantelli (Agassiz, 1843) - Shimada et al., p. 185, figs.1-6. 266 2008 Cretoxyrhina mantelli (Agassiz) - Shimada, p. 21, figs. 1-2, 6-7. 267 2011 Cretoxyrhina mantelli Agassiz - Bourdon, Everhart, p. 15, figs. 2, 4-7. 268 2012 Cretoxyrhina mantelli (Agassiz, 1838) - Cappetta, p. 236, figs. 216E-I. PT 269 2013 Cretoxyrhina mantelli (Agassiz, 1843) - Cook et al., p. 568, fig. 10. RI 270 2013 Cretoxyrhina mantelli (Agassiz, 1843) - Siverson et al., p. 3, fig. 10. 271 vp.2013 Vertebre di selaceo squaloideo (sic) - Colombara, p. 40 (text-fig.). SC 272 2014 Isurus denticulatus Glickman, 1957 - Diedrich, p. 8, figs. 4, 5A-D, 7A-F, N-V. 273 2015 Cretoxyrhina mantelli (Agassiz, 1843) - Newbrey et al., p. 878, figs. 7A-C, 8. 274 U 2017 Cretoxyrhina mantelli - Everhart, fig. 13.3. AN 275 M 276 Nomenclatural notes 277 Cretoxyrhina mantelli was originally erected by Louis Agassiz in his paleoichthyological treatise D 278 ʻRecherches sur les poissons fossilesʼ (hereinafter ʻRecherchesʼ). The ʻRecherchesʼ were published TE 279 in a series of installments (ʻLivraisonsʼ) over a span of eleven years (1833-1844; Brignon, 2015; 280 Woodward, Sherborn, 1890) and it caused considerable confusion and ambiguity in establishing the EP 281 correct years of erection of several taxa and Cretoxyrhina mantelli is no exception. The date of 282 erection of this species was considered to be either 1838 or 1843 according to various authors (e.g, C AC 283 Cappetta, 1987, 2012; Herman, 1977; Shimada, 1997a-e). In fact, the plate 33 was published in 284 Agassiz (1838, Volume III), whereas the description of the taxon was published in Agassiz (1843, 285 Volume III, text: p. 280), both reporting the name "Oxyrhina Mantellii" and, thus, creating 286 confusion for nowadays nomenclatural rules. Furthermore, Agassiz published 72 supplementary 287 sheets (ʻFeuilletons additionnelʼ) in addition to several ʻLivraisonsʼ, which were later removed from 288 the five volumes on the advice of the author, because they contained redundant information in his 289 opinion (see Brignon, 2015). A few copies of the “Recherches” still containing these ʻFeuilletonsʼ, ACCEPTED MANUSCRIPT 290 nevertheless survived, which are pivotal for nomenclatural purposes because they contain the first 291 mentions of numerous new valid taxon names created by Agassiz (Brignon, 2015). In the 292 ʻFeuilleton additionnelʼ published with the 4th ʻLivraisonʼ in January 1835, Agassiz (1835: p. 54) 293 listed several new taxa identified from the Mantellʼs collection (NHM) and that were previously 294 described and figured by Mantell (1822). Namely, Agassiz (1835: p. 54) unambiguously referred PT 295 isolated shark teeth from this collection (Mantell, 1822: plate 32, figs. 4, 7, 8, 10, 11, 26, 28) to the RI 296 new taxon Lamna Mantellii (Fig. 3A). According to Article 12.1 of the ICZN Code (Ride et al., 297 2000), the name Lamna Mantellii, associated by Agassiz (1835) with previously published figures, SC 298 is available by indication (and Article 12.2.7), and the nominal species therefore takes authorship 299 and date from Agassiz (1835). Furthermore, the seven teeth figured by Mantell (1822) are the 300 U original figured syntypes of C. mantelli (non the ʻtype specimensʼ of Woodward, 1889: p. 377). AN 301 Woodward (1889) listed specimens NHM PV OR 4524, 4527, 4539, 4540 (Figs. 3B-E) as those M 302 specimens figured by Mantell (1822: figs. 8, 11, 26, 28); furthermore, there is a reference of 303 Mantell’s corresponding plate and figure on the label stuck on the specimens (Amalfitano J., D 304 Giusberti L., pers. obs.). For these reasons, these specimens are surely four of the seven figured TE 305 syntypes originally designated by Agassiz (1835), while the remaining figured syntypes (Mantell, 306 1822: figs. 4, 7, 10) are possibly lost or not easily detectable among the teeth of the Mantell’s EP 307 collection. 308 C AC 309 Referred material 310 MPPSA-IGVR 36371*, 45305*, 45324*, 45326*, 45334*, 45344-45345*, MGC-IGVR 311 81375-81376*; MGP-PD 3805, 5404, 6721, 6736, 7342, 7343, 7372, 8498, 8889, 14020, 14029, 312 14034-14039, 14042, 22401-22470, 23527-23529; MCSNV V. 1094, 1095, 11798, 12518, 12519 313 (starred specimens indicate those including other skeletal elements associated with teeth). 314 315 Emended diagnosis ACCEPTED MANUSCRIPT 316 Lamniform shark unique in having the following apomorphic combination of dental 317 characters: dental formula: 4S-2A-4I-11(+x)L / 1s?-2a-1i-15(+x)l; monognatic heterodonty, 318 strongly disjunct in upper teeth, slightly gradational in lower teeth; teeth with usually narrow, blade- 319 like, asymmetrical and distally curved or sloped crown, extending above the mesial branch of root 320 (especially in anterior teeth); cusp sharply pointed; symphyseal teeth strongly asymmetrical, with PT 321 crown higher than wide, and distal root lobe generally more developed than the mesial one; first RI 322 symphyseal teeth distally inclined, other symphyseal teeth mesially curved; anterior teeth nearly 323 symmetrical; lower anterior teeth with very high cusp, higher than wide, particularly the second SC 324 lower anterior tooth; first anterior teeth (upper and lower) usually the most symmetrical, with the 325 second anterior tooth characterized by a slight distal inclination; crown tending to be labiolingually 326 U thicker and more sigmoidal (labially directed) in lower than in upper teeth, which are flatter and AN 327 thinner, with the point of the cusp labially curved; lower teeth with more massive roots than uppers; M 328 intermediate teeth with crown height greater or similar to crown width, cusp inclined distally, with 329 the sole first upper intermediate tooth distally curved; lateral teeth generally with crown wider than D 330 high, slightly to strongly distally inclined, decreasing in size distally, their inclination increases TE 331 distally; enameloid smooth and cutting edges continuous; enameloid may form weak basoapical 332 ripples, mainly on the labial side; lingual face convex, labial face nearly flat or only weakly convex; EP 333 basal part of crown characterized by small oblique heels (shoulders), with continuous cutting edges 334 on the sides of the cusp; both heels may occasionally bear a lateral cusplet in distolateral and C AC 335 commissural teeth; tooth neck marked; root strongly bilobated, becoming more splayed distally; 336 concavity of the basal margin of the root low (lateral teeth) to high (anterior-intermediate teeth); 337 mesial branch of the root usually pointed, the distal one rounded and expanded; lingual 338 protuberance developed, lacking a nutrient groove and usually bearing one nutrient foramen. 339 340 Locality and horizon ACCEPTED MANUSCRIPT 341 All the specimens described here come from the ʻlastameʼ lithofacies of the Scaglia Rossa 342 Formation of Mt. Loffa and other surrounding sites (Lessini Mountains of Verona Province, Italy). 343 Based on calcareous nannofossils and planktic foraminifera biostratigraphy, the specimens are dated 344 to the middle-late Turonian (for more details see Tab. 2) according to the correlation between 345 calcareous plankton zones and stages by Ogg, Hinnov (2012) (UC7-UC9 Zones, Burnet, 1998 and PT 346 Dicarinella primitiva/Marginotruncana sigali Zone, Coccioni, Premoli Silva, 2015). RI 347 348 Description SC 349 We provide below a brief description of the skeletal remains of C. mantelli found in the 350 ʻlastameʼ quarries. The most complete and best-preserved specimens are MPPSA-IGVR 36371 and 351 45305 (Fig. 4). U AN 352 Specimen MPPSA-IGVR 36371 (Fig. 4A) is preserved on nine limestone slabs of different M 353 size, which were originally part of a single bigger slab. The specimen exhibits 37 teeth (23 totally or 354 partially preserved and 14 preserved as impression only), 157 vertebral centra and some fragments D 355 of tessellated cartilage, whose original skeletal position is impossible to identify due to the partial TE 356 preservation. Teeth are scattered on different slabs, and therefore do not provide any evidence of the 357 original tooth arrangement. Most of the teeth are localized in what appears to be the cranial region, EP 358 although some of them are scattered along the vertebral column. The teeth (Fig. 5) display the 359 characteristic features of C. mantelli. There are no lateral cusplets in any preserved tooth. Most of C AC 360 teeth are embedded in the rock, exposing only the labial or lingual side, thereby rendering difficult 361 to establish their original position. At least 16 teeth have complete crowns and roots (e.g., Figs. 5A- 362 B, F). Other teeth are fragmentary. The highest tooth measures 67 mm in total height (52 mm in 363 crown height on the labial side), while the lowest tooth is ca. 28 mm high. 364 The vertebral centra show the typical lamnoid structure (see Applegate, 1967: p. 67), with 365 well-calcified asterospondylic, amphicoelous, and imperforated centra. All centra suffered a 366 taphonomic anteroposterior compression; some show a slight taphonomic distortion and some ACCEPTED MANUSCRIPT 367 others are incomplete or broken. Most of the vertebral centra are disarticulated, moderately 368 scattered near the longitudinal axis of the skeleton and laying on the anterior or posterior articular 369 face. Part of the vertebral column, including more or less 40 posterior caudal centra, is accumulated 370 in a pile along the vertebral column axis. At least twenty-five vertebral centra are extremely 371 fragmentary or preserved as impressions only. The largest vertebral centrum has a maximum PT 372 diameter of 107 mm, while the smallest one has a maximum diameter of ca. 40 mm. The antero- RI 373 posterior length ranges from 14 mm to ca. 40 mm. 374 Remains of tessellated calcified cartilage are very fragmentary (Fig. 6) and show a mosaic SC 375 texture and rough surfaces. Specimen MPPSA-IGVR 36371 was sampled near the cranial region for 376 investigating placoid scales and a total of 526 placoid scales were found. The placoid scales (Figs. 377 U 7A-R) consist of a rhomboidal root and a crown (covered with enameloid) with different shapes AN 378 depending on the position along the body of the shark. Their size does not exceed 500 µm in height M 379 and width. The root may preserve a nutrient foramen at the base (Fig. 7F) and one or two foramina 380 along the posterior side. The crown is constricted near the base, forming a neck. The shape of the D 381 crown can be stubby (e.g., Figs. 7A-F, 7J-L) or slender (e.g., Figs. 7G-I), with a rounded (e.g., Figs. TE 382 7A-C, 7E-F, 7J-L) or pointed cusp (e.g., Figs. 7D, 7G-I, 7M-R). No scales with multiple cusps were 383 found. The anterior face of the crown can be characterized by a more or less expanded, flattened or EP 384 weakly inclined and swollen superior shelf (Figs. 7J, 7M-O), forming a hooked profile in lateral 385 view (Figs. 7B, 7P-Q). The anterior face can be smooth (Figs. 7D, 7G-H, 7J-K) or crossed by thin C AC 386 parallel ridges (or riblets) that are divided by grooves running antero-posteriorly towards the cusp, 387 which can reach the posterior margin of the cusp (Figs. 7A, 7M-R). Ridges, in some cases, may be 388 restricted to the anterior part of the scale, extending apically from the neck up to midway the cusp, 389 while the remaining apical part of the cusp is smooth (Fig. 7A). The posterior face of the crown is 390 convex and smooth, crossed by a central vertical weak keel (Figs. 7C, 7E, 7I, 7L). 391 Specimen MPPSA-IGVR 45305 (Fig. 5B) is preserved on four slabs and comprises 131 392 vertebral centra, 55 teeth, and is associated with fragments of tessellated calcified cartilage. All ACCEPTED MANUSCRIPT 393 teeth are scattered around the cranial region and totally displaced, without any evidence of their 394 original arrangement, but are morphologically similar to those of MPPSA-IGVR 36371 (Figs. 8-9). 395 In two cases (e.g., Fig. 9A), the teeth are still articulated to tessellated calcified cartilage, thereby 396 confirming that these fragments were part of the jaws. It was not possible to identify whether these 397 fragments were part of the palatoquadrate or of the Meckel's cartilage, because one of them PT 398 represents a lateral tooth and the other a commissural one, both not readily identifiable as upper or RI 399 lower. The crown height of the highest tooth measures 42 mm on the labial side, while the smallest 400 one measures only 5 mm. The maximum diameter of the largest centrum is 98 mm, while that of the SC 401 smallest one is ca. 42 mm. The antero-posterior length ranges from 20 to ca. 35 mm. In all teeth, the 402 root is poorly preserved or completely lacking (e.g., Figs. 9B-D). Most vertebral centra are still 403 U connected to each other lying on their lateral side along the axis of the vertebral column, forming an AN 404 arch. The posterior-most centra show a moderate degree of disarticulation. The centra are of the M 405 lamnoid type and show the same taphonomic alterations as observed in MPPSA-IGVR 36371. No 406 placoid scales were found in the samples extracted from this specimen. D 407 Other skeletal remains include segments of partially articulated vertebral columns associated TE 408 with teeth and cranial cartilage fragments (MPPSA-IGVR 45324, MPPSA-IGVR 45326, MPPSA- 409 IGVR 45334, MPPSA-IGVR 45344-45345, MGC-IGVR 81375-81736). All of these specimens EP 410 have dental and vertebral features that are identical to those of MPPSA-IGVR 36371 and 45305, 411 and the teeth are still embedded in the limestone, partially or totally exposed only in labial or C AC 412 lingual views. The vertebral column segments show various degrees of disarticulation, from totally 413 displaced centra (MPPSA-IGVR 45344-45345) to almost articulated sets (e.g., MGC-IGVR 81375- 414 81376). The majority of the vertebral centra lie facing up exposing the articular surfaces, while few 415 others expose the lateral side, as in MPPSA-IGVR 36371 and 45305. 416 Specimen MPPSA-IGVR 45324 (see supplementary material Fig. A.1β) includes five teeth, 417 a single vertebral centrum, and small fragments of tessellated cartilage. The highest complete tooth 418 has a total height of ca. 52 mm, and a crown height measured on the labial side of 37 mm. ACCEPTED MANUSCRIPT 419 Specimen MPPSA-IGVR 45326 (see supplementary Fig. A.1γ) comprises 16 teeth, two 420 vertebral centra and some fragments of tessellated cartilage. Most of the teeth are almost totally 421 embedded in the rock. The crown height of the highest tooth (preserved as impression only), 422 measured on the labial side, is ca. 53 mm. The maximum diameter of the largest vertebral centrum 423 is ca. 97 mm. PT 424 Specimen MPPSA-IGVR 45334 (see supplementary Fig. A.1δ) consists of a slab containing RI 425 two teeth, 14 vertebral centra, and a fragment of tessellated cartilage. The total height of the highest 426 tooth measures ca. 37 mm, and the crown height measured on the labial face 28 mm. The maximum SC 427 diameter of the largest vertebral centrum is 97 mm. 428 Specimen MPPSA-IGVR 45344-45345 (Fig. 10) includes slab and counterslab that contain 429 U 16 teeth, 15 vertebral centra, and a fragment of tessellated cartilage. The teeth solely preserved the AN 430 crown (e.g., Figs. 10B-D). The crown height of the highest tooth measured on the labial side M 431 reaches 46 mm, while that of the smallest one measures ca. 20 mm. The maximum diameter of the 432 largest vertebral centrum is 76 mm, while that of the smallest vertebral centrum measures ca. 60 D 433 mm . TE 434 Specimen MGC-IGVR 81375-81376 (Fig. 11) is preserved on a slab and its counterpart with 435 80 teeth, 33 vertebral centra, and many fragments of tessellated cranial cartilage. The total height of EP 436 the highest tooth is 67 mm, whereas the smallest one measures 12 mm. In this tooth set, the 437 maximum crown height on the labial side is 47 mm. The maximum diameter of the largest vertebral C AC 438 centrum is 86 mm, while that of the smallest one is 45 mm. A total of 142 placoid scales was found 439 in a sample extracted from specimen MGC-IGVR 81375, exhibiting morphologies very similar to 440 those observed in MPPSA-IGVR 36371 (Fig. 12). 441 442 Discussion 443 The shark specimens from the ʻlastameʼ lithozone of the Scaglia Rossa Formation can be 444 unquestionably referred to the genus Cretoxyrhina, because of the presence of narrow and blade- ACCEPTED MANUSCRIPT 445 like crowns. The oblique heels on the side of the cusp, the absence of the lateral cusplets, and the 446 asymmetric, sharp pointed and rather robust crown (if compared to other congeneric species) 447 support the assignment of the specimens to Cretoxyrhina mantelli. Cretoxyrhina mantelli had a 448 cosmopolitan distribution, since it has been reported in Europe, Russia, Africa, North America and 449 Brazil (Cappetta, 2012). The genus Cretoxyrhina includes also the species C. denticulata PT 450 (Glickman, 1957) and C. agassizensis (Underwood, Cumbaa, 2010), both of which are diagnosed RI 451 by the presence of lateral cusplets (with sharp apices in C. agassizensis, and rounded apices in C. 452 denticulata) in many of the lateroposterior teeth (see Newbrey et al., 2015 and Underwood, SC 453 Cumbaa, 2010). C. agassizensis teeth also exhibit a slender and generally straight cusp (also in 454 lateroposterior teeth) than C. mantelli and C. denticulata (see Underwood, Cumbaa, 2010), and 455 U incomplete cutting edges on small juvenile anterior teeth (Newbrey et al., 2015). The latter AN 456 character, however, cannot be taken into account to compare and differentiate adult teeth from the M 457 other species. Some associated dentitions of C. mantelli exhibit almost upright crown on 458 lateroposterior teeth (e.g., Eastman, 1894), but this condition may be related to gynandric D 459 heterodonty (Siverson M., pers. comm.). Two of the three species have partially overlapping TE 460 stratigraphic distributions, because C. denticulata ranges from the lower Cenomanian to the lower 461 middle Cenomanian, C. agassizensis ranges from the upper middle Cenomanian to lower middle EP 462 Turonian (Newbrey et al., 2015), whereas Cretoxyrhina mantelli is reported globally from the upper 463 Cenomanian to the Campanian (Bourdon, Everhart, 2001; Cappetta, 2012; Shimada, 1997e). C AC 464 Another species of Cretoxyrhina, the upper Albian-lower Cenomanian C. vraconensis (Zhelezko, 465 2000), was revised by Siverson et al. (2013) and clearly differs from C. mantelli in having cusplets 466 and different tooth morphologies in corresponding dental positions (for details see Siverson et al., 467 2013: p. 14). The species of the genus Cretoxyrhina might possibly represent chronospecies of a 468 single evolutionary lineage (see Newbrey et al., 2015). This evolutionary lineage was characterized 469 by the progressive reduction of lateral cusplets and the progressive increasing size and robustness of 470 teeth throughout its temporal range (see Cook et al., 2013; Siverson, Lindgren, 2005; Underwood, ACCEPTED MANUSCRIPT 471 Cumbaa, 2010). An increase in tooth size (probably corresponding to an increase in body size; see 472 “Paleobiological remarks” for individual length estimates), a significant decrease of crown height- 473 crown width ratio and a loss of lateral cusplets are recorded in the upper Cenomanian-Coniacian 474 interval (see Shimada, 1997e; Siverson, Lindgren, 2005) and the size of the teeth of the Italian 475 specimens indicates that the teeth from the middle-upper Turonian had already reached a size PT 476 similar to those of the Coniacian specimens. RI 477 Specimens MPPSA-IGVR 36371 and 45305 are among the most complete remains of 478 Cretoxyrhina mantelli in the world. Most of the Cretoxyrhina mantelli skeletons were recovered SC 479 from the Coniacian-lower Campanian Niobrara Chalk in Kansas (USA) and include very well- 480 preserved articulated remains (e.g., FHSM VP-323, FHSM VP-2187; Newbrey et al., 2015; 481 U Shimada, 1997d). Eastman (1894) reported the first nearly complete skeleton from Kansas, which AN 482 belonged to the collections of the Bayerische Staatssammlung für Paläontologie und Geologie of M 483 Munich, Germany, but was unfortunately destroyed during World War II (Diedrich, 2014). 484 Additional incomplete skeletons were found in Europe. These include a set of about 57 teeth D 485 associated with vertebral centra of a single individual from the Upper Chalk (Coniacian-lower TE 486 Campanian) of Grays, southeastern England (NHM PV OR 32346-32347, 39434; Woodward, 1889, 487 1911: fig. 60). Diedrich (2014: fig. 5) described partial skeletal remains of "Isurus denticulatus" (= EP 488 Cretoxyrhina mantelli) consisting of 149 vertebral centra that are mostly disarticulated and 489 associated with five teeth, coming from the upper Turonian of Halle/Westphalia in Germany. C AC 490 In northeastern Italy, Cretoxyrhina mantelli was reported from the upper Cenomanian 491 Bonarelli Level (Sorbini, 1976) up to the uppermost middle and upper Turonian ‘lastame’ of the 492 Scaglia Rossa Formation as documented herein (see Tab. 2). The teeth from the Bonarelli Level are 493 currently under revision, but at least two teeth among those reported by Sorbini (1976) exhibit a 494 series of characters that allow to refer them to C. mantelli (Amalfitano J., pers. obs.). The Italian 495 specimens reported herein represent some of the oldest associated skeletal remains of C. mantelli. ACCEPTED MANUSCRIPT 496 Specimens MPPSA-IGVR 36371 (with 157 preserved centra over an estimated total count 497 of 216 vertebrae given by Shimada et al., 2006) and MPPSA-IGVR 45305 (with 131 vertebrae) are 498 considered two partial vertebral columns associated with disarticulated teeth and fragments of 499 tessellated calcified cartilage. The specimen MGC-IGVR 81375-81376 is interpreted as the anterior 500 portion of the vertebral column with associated teeth and fragments of tessellated cartilage. The PT 501 remaining specimens (MPPSA-IGVR 45324, 45326, 45344-45345) are short anterior portions of RI 502 the vertebral column associated with disarticulated teeth. MPPSA-IGVR 45334 is distinct because it 503 probably includes posterior precaudal vertebral centra. Mid-trunk vertebral centra are usually the SC 504 largest in the vertebral column (see Newbrey et al., 2015). The maximum vertebral diameter is 97 505 mm in MPPSA-IGVR 45334, which is similar to the maximum vertebral diameter in MPPSA- 506 U IGVR 36371 (107 mm) and therefore are most likely large posterior precaudal centra. Further AN 507 taphonomic inferences are provided in the paragraph “Taphonomic remarks”. M 508 509 ?Cretoxyrhina sp. D 510 Figs. 13-14, supplementary material Figs. A.1ε-η TE 511 512 vp.2013 Vertebre di selaceo squaloideo (sic) - Colombara, p. 40 (text-fig.). EP 513 514 Referred material C AC 515 MPPSA-IGVR 45308, MPPSA-IGVR 45319-45320, MPPSA-IGVR 45321-45322, MPPSA- 516 IGVR 45323, MPPSA-IGVR 45337, MGC-IGVR 47789, MGP-PD 31960. 517 518 Locality and horizon 519 All these specimens come from the ʻlastameʼ of Scaglia Rossa Formation of Loffa Mount 520 and other surrounding sites (Lessini Mountains, Verona Province, Italy). 521 ACCEPTED MANUSCRIPT 522 Description 523 The material described here consists of portions of vertebral columns of lamniform sharks 524 with different degrees of disarticulation. The vertebral centra are of lamnoid type sensu Applegate 525 (1967). 526 Specimen MPPSA-IGVR 45308 (Fig. 13A) consists of 55 scattered vertebral centra with PT 527 diameters ranging from 38 to 81 mm. RI 528 Specimens MPPSA-IGVR 45319 and 45320 (Fig. 13B) are preserved on two slabs with 529 portions of vertebral columns that included a total of 25 vertebral centra (12 and 13 centra, SC 530 respectively). Centra are partially embedded in the rock and are moderately articulated, arranged in 531 a nearly linear row mostly showing the lateral side. These elements likely belong to a single 532 U individual as the vertebral diameter gradually decreases along the vertebral rows. AN 533 Specimens MPPSA-IGVR 45321 and MPPSA-IGVR 45322 (see supplementary material M 534 Figs. A.1ε-ζ) represent two portions of the vertebral column containing 17 and 12 centra, 535 respectively. Centrum diameters range from 92 to 98 mm. The portions probably belong to two D 536 distinct individuals, because all centra have similar diameters and there is no decrease in diameter TE 537 along the vertebral rows. 538 EP 539 Specimen MPPSA-IGVR 45323 (Fig. 13C) is a slab with 52 articulated vertebral centra 540 arranged in a linear row except for eight centra that are slightly displaced from the row. The row C AC 541 shows a slight bend toward one extremity. The diameter of the centra ranges from 40 to 85 mm. 542 Specimen MPPSA-IGVR 45337 (see supplementary material Fig. A.1η) is preserved on a 543 small slab with seven vertebral centra arranged in two clusters of three articulated centra plus a 544 single centrum separated from the clusters. Centra are embedded in the rock and measurements 545 cannot be taken. ACCEPTED MANUSCRIPT 546 Specimen MGC-IGVR 47789 (Fig. 14) consists of a partial vertebral column comprising ca. 547 122 vertebral centra that are arranged in an arched row. The maximum diameter of the vertebral 548 centra is 84 mm. 549 Specimen MGP-PD 31960 (Fig. 13D) is a slab containing a segment of a vertebral column 550 with 52 centra. The centra lie on the articular faces and are slightly scattered along the original axis PT 551 of the vertebral column, with only a few centra that are articulated to each other. The diameter of RI 552 the centra is nearly consistent throughout the segment, with a maximum value of 67 mm. 553 SC 554 Discussion 555 The segments of vertebral columns described above can be unambiguously assigned to 556 U lamniform sharks because of the vertebral centra of lamnoid type (Applegate, 1967; Shimada, 2007; AN 557 see also MPPSA-IGVR 36371 above for a brief description of the vertebral type). At least seven M 558 shark specimens of the ‘lastame’ sample with associated teeth and vertebral centra allow a reliable 559 specific attribution to Cretoxyrhina mantelli. The only other lamniform shark from the ‘lastame’ is D 560 Cretodus (Amalfitano et al., 2017b), which is represented therein by a single TE 561 specimen.Furthermore, most of the isolated teeth from the ‘lastame’ belong to Cretoxyrhina 562 mantelli. Therefore, Cretoxyrhina mantelli is so far the most common lamniform shark from the EP 563 ‘lastame’ and the additional vertebral column portions described here are tentatively referred to as 564 ?Cretoxyrhina sp. C AC 565 The arrangement of the vertebral centra on the slabs does not allow determining the original 566 position of the preserved portions within the original vertebral column, with the exception of MGC- 567 IGVR 47789, which is the most complete segment (see Fig. 14). However, the variability of the 568 diameter of the centra along the vertebral column provides some information. MPPSA-IGVR 45308 569 seems to include both caudal and precaudal centra, because of the high range of vertebral diameters 570 from 38 to 81 mm. A similar range in vertebral diameter occurs in MPPSA-IGVR 43323; here the 571 original arrangement of the vertebral column is partly retained, so that the caudal flexion of the ACCEPTED MANUSCRIPT 572 vertebral column, in correspondence of the upper lobe of the caudal fin, is still evident (Fig. 13C). 573 The remaining specimens (MPPSA-IGVR 45319, 45320, 45321, 45322, MGP-PD 31960) show 574 more constant values in vertebral diameter, and probably represent transitional segments between 575 the anterior abdominal and the precaudal portions or between the precaudal sector and the caudal 576 extremity. PT 577 RI 578 Paleobiological remarks 579 Length and longevity estimates SC 580 Qualitative and quantitative data provided by fossil remains of extinct sharks can be useful 581 to infer various parameters related to aspects of their paleobiology. Such inferences are often 582 U hampered by the fact that complete shark skeletons are rare in the fossil record due to low AN 583 fossilization potential of cartilaginous skeletal elements (Shimada, 2008). Shimada (1997c) M 584 provided a general analysis on the periodic marker bands of Cretoxyrhina mantelli vertebral centra 585 and, later, Shimada (2008) expanded his study through a quantitative exploration of the ontogenetic D 586 parameters of C. mantelli (e.g., length at birth, growth rate, and longevity). Other authors addressed TE 587 this issue, comparing the data from C. mantelli skeletons with those of other sharks in order to 588 hypothesize the length at birth, total length, and longevity in various extinct lamniform sharks (e.g., EP 589 Cretalamna hattini in Shimada, 2007; Archaeolamna kopingensis in Cook et al., 2011; and C 590 Cardabiodon ricki in Newbrey et al., 2015). The specimens described herein, especially MPPSA- AC 591 IGVR 36371 and MPPSA-IGVR 45305 provide some significant data for the estimation of their 592 total length (TL) and longevity. Using the equations of Shimada (2008) on the specimen MPPSA- 593 IGVR 36371, that has a maximum crown height measured on the labial side (EH in Shimada, 2008) 594 of 52 mm and a maximum vertebral diameter (CD in Shimada, 2008) of 107 mm, results in 595 estimated TLs of 650 cm and of 615 cm, respectively. These estimated TLs suggest that MPPSA- 596 IGVR 36371 represents one of the largest individuals of Cretoxyrhina mantelli ever found. The size 597 is comparable to the asymptotic (= maximum) length for Cretoxyrhina mantelli (691 cm) proposed ACCEPTED MANUSCRIPT 598 by Shimada (2008), and to the estimated range (640-700 cm) for the largest individual described to 599 date (represented by a well-preserved caudal fin; Shimada et al., 2006). MPPSA-IGVR 45305 (42 600 mm of maximum EH and 98 mm of maximum CD) has an estimated total length of 525 cm based 601 on EH, and 563 cm based on CD. Other relevant specimens are MPPSA-IGVR 45344-45345 and 602 MGC-IGVR 81375-81376, with the maximum EH of 46 and 47 mm corresponding to TLs of 575 PT 603 cm and 587.5 cm, respectively. The other specimens exhibit a comparatively smaller size or are RI 604 more fragmentary and do not include the largest precaudal vertebra or the highest tooth, and for this 605 reason these are not used for size estimates. SC 606 The incremental bands of the vertebral centra (couplets of translucent and opaque bands that 607 correspond to annual growth) allow the estimation of the individual longevity in extant and extinct 608 U elasmobranchs (e.g., Amalfitano et al., 2017c; Cailliet, Goldman, 2004; Goldman et al., 2012; AN 609 Newbrey et al., 2015; Shimada, 2007; Shimada, 2008). These bands can be observed in the M 610 vertebral centra of specimens IGVR 36371 and 45305. Other than the birth band, it is possible to 611 observe 26 incremental bands in MPPSA-IGVR 36371 (Fig. 15A) and 21 in MPPSA-IGVR 45305 D 612 (Fig. 15B), thereby suggesting an estimated age of at least 26 and 21 years, respectively. The TE 613 incremental band counting is consistent with the results obtained from the equation by Shimada 614 (2008) that uses the number of bands to estimate the individual total length. The equation, EP 615 TL=119(BN+1)0.522, in which BN refers to the band number (with 0 referring to the birth band and 616 the outermost band assumed to represent the maximum BN), provides total length estimates (ca. C AC 617 665 cm for MPPSA-IGVR 36371, 597 cm for MPPSA-IGVR 45305) that are only slightly larger 618 than those obtained using the vertebral and dental measurements (615-650 cm for MPPSA-IGVR 619 36371 and 525-565 cm for MPPSA-IGVR 45305). 620 621 Swimming behavior and paleoecology 622 Newbrey et al. (2015) observed that the vertebral centra of Cretoxyrhina mantelli are 623 relatively antero-posteriorly compressed when compared to those of other fossil and extant ACCEPTED MANUSCRIPT 624 lamniform sharks. The short vertebral centra and the high vertebral count led Newbrey et al. (2015) 625 to hypothesize a carangiform swimming mode for Cretoxyrhina mantelli, implying that it was a 626 moderately fast swimmer with high maneuverability. 627 The swimming capabilities of sharks can be estimated also through the morphological 628 analysis of the placoid scales (e.g., Reif, 1985). Scale morphology suggests that Cretoxyrhina PT 629 mantelli was a fast swimming shark (Shimada, 1997d). According to Reif and Dinkelacker (1982), RI 630 keels (= ridges or riblets) and grooves on the scales that run approximately parallel to the body axis, 631 as observed in Cretoxyrhina mantelli, are characteristic of fast swimming sharks (see also Shimada, SC 632 1997d). Reif (1985) recognized six ecological groups of sharks based on their locomotory habits, in 633 which placoid scales have different functions that correlate to the ecology of the various shark 634 U species. Only in two groups, the fast swimming pelagic sharks and the large near-shore AN 635 predators/moderate speed pelagic predators, the morphology of the placoid scales shows an evident M 636 hydrodynamic function. Fast swimming pelagic sharks have flat, usually overlapping scale crowns 637 that form a dense pavement (Reif, 1985). Crowns exhibit a rounded posterior end or short cusps. D 638 The surface of the crown in this group of sharks is ornamented with fine parallel ridges that have TE 639 average distances between 40 and 80 µm with U-shaped grooves separating the ridges. The placoid 640 scales of Cretoxyrhina mantelli show all of these features. When found articulated, such as those EP 641 figured in Shimada (1997d: fig. 8), they have a pavement-like arrangement to reduce hydrodynamic 642 drag. As far as the scale ornamentation is concerned, placoid scales from MPPSA-IGVR 36371 and C AC 643 MGC-IGVR 81375 exhibit a pattern of ridges and grooves nearly identical to that observed by 644 Shimada (1997d), Shimada et al. (2006) and Diedrich (2014) in Cretoxyrhina mantelli from other 645 localities. The distance between the ridges in the specimens described herein ranges between 33 µm 646 and 60 µm, with the mean value falling within the range of fast swimming sharks (Reif, 1985). 647 Moreover, we employed a method utilized by Reif (1985) that was recently used for other fossil 648 sharks (e.g., Marramà et al., 2018), which takes into account the ridge spacing and the scale width. 649 We plotted the average ridge spacing and average crown width from a few individuals (FHSM VP- ACCEPTED MANUSCRIPT 650 2187, MPPSA-IGVR 36371, MGC-IGVR 81375; see Tab. A.3) and compared the results with those 651 of fast pelagic hunting sharks and large near-shore hunters/pelagic predators of moderate speed (see 652 Reif, 1985). The results are shown in Fig. 16, in which Cretoxyrhina mantelli clearly falls within 653 the fast pelagic hunting shark group, in particular the ridge spacing is very similar to that of Isurus 654 oxyrhinchus and Sphyrna tudes. The average crown width, however, differs from those observed in PT 655 these two taxa, but it is more similar to that of Lamna nasus. Comparing the morphology and RI 656 arrangement of placoid scales of Cretoxyrhina mantelli and those of extant lamniform sharks, the 657 scales of Cretoxyrhina mantelli (see Shimada, 1997d: fig. 8) nevertheless differ from those of SC 658 Isurus oxyrhinchus (see Reif, 1985: pl. 20, 21), which is a very fast hunter adapted to chasing fast 659 moving fishes, such as swordfishes and tunas, in that the placoid scales have a looser arrangement. 660 U Actually, the placoid scales arrangement in C. mantelli is more similar to that in Carcharodon AN 661 carcharias (see Reif, 1985: pl. 23). However, the number of ridges in the scales of C. mantelli (up M 662 to nine; Shimada, 1997d) is greater than that observed in I. oxyrhinchus (up to five ridges; Reif, 663 1985) and Carcharodon carcharias (up to three ridges). Reif (1985) hypothesized that new ridges D 664 were added with the expansion of the crown width to maintain constant the distance between the TE 665 ridges. 666 Considering all the aspects discussed herein, the conclusions drawn by other authors about caudal EP 667 fin morphology and metabolic rate estimates (e.g., Ferron, 2017; Kim et al., 2013), and the fossil 668 record of predation attributed to C. mantelli (see Hone et al., 2018; Shimada 1997b; Shimada, C AC 669 Hooks III, 2004), this Cretaceous lamniform shark was likely a fast swimmer with an ecology in 670 some ways similar to that of the living Carcharodon carcharias (see Shimada, 1997d). 671 672 Taphonomic remarks 673 General features of the C. mantelli deadfall from ʻlastameʼ 674 All of the specimen from the ʻlastameʼ documented herein are preserved within the 675 limestone beds or on their bedding surfaces, sometimes draped with calcareous marly sediments. ACCEPTED MANUSCRIPT 676 Lamniform sharks represent the most represented taxonomic group within associated or at least 677 partly articulated vertebrate skeletal remains found in the ‘lastame’, with 15 specimens out of a total 678 of at least 39 determined specimens, including four other chondrichthyans (Ptychodontidae, 679 Cretodus, and Onchosaurus), 13 marine turtles, six mosasaurs and a bony fish.. Other skeletal 680 remains of lamniform sharks consisting of segments of vertebral columns (at least three) are housed PT 681 in the collections of the MCSNV and probably come from the ‘lastame’ lithozone (Zorzin R., pers. RI 682 com.), thereby confirming the great abundance of lamniform shark remains. 683 Skeletal remains of lamniform sharks include mainly portions of vertebral columns, which SC 684 in many cases are associated with disarticulated tooth sets (sensu Shimada, 2005), and sometimes 685 also with fragments of tessellated calcified cartilage and placoid scales. Teeth in some specimens 686 U are accumulated at the anterior end of the vertebral column; sometimes they are associated with AN 687 tessellated cartilage of the mandibular arch, sometimes they are totally displaced and scattered M 688 along the vertebral column. Most of the teeth show fissures on the enameloid produced by 689 compression, and, in many cases, have broken cusps or roots, or may be preserved as impression D 690 only. TE 691 Mineralized cranial elements are not well-preserved, because the cranial skeleton consists of 692 tessellated cartilage, formed by a mosaic of tesserae (composed of both prismatic and globular EP 693 calcification) overlaying a core of extracellular matrix that is not mineralized (Dean, Summers, 694 2006). The decay of the extracellular matrix causes the disarticulation and displacement of the C AC 695 tesserae. Only multilayered elements, which are considerably stiffer (e.g., jaws; Dean, Summers, 696 2006; Maisey, 2013), are characterized by a better state of preservation and occur in some of the 697 specimens described herein (e.g., MPPSA-IGVR 45305). 698 Most vertebral centra described herein show antero-posterior compression; some suffered a 699 slight taphonomic deformation and some others are incomplete, broken or fragmented. Some centra 700 also show diagenetic alteration, such as oxidation (e.g., manganese dendrites; see Fig. 15B). The 701 completeness of the skeletons is clearly biased by the technical difficulty of extracting larger slabs ACCEPTED MANUSCRIPT 702 in the quarries, indicating that most of the skeletons were probably more complete when buried 703 (Amalfitano et al., 2017b). The most complete vertebral column available, MPPSA-IGVR 36371, 704 includes 157 vertebral centra and was fully extracted, as reported by the quarrymans, and 705 subdivided into smaller slabs to facilitate the transport and the preparation (Fig. 4A). 706 The vertebral column segments within the ‘lastame’ sample show different degrees of PT 707 disarticulation, ranging from totally disarticulated and scattered vertebral centra (e.g., MPPSA- RI 708 IGVR 45308, Fig. 13A) to nearly perfectly articulated segments (e.g., MPPSA-IGVR 45305, Fig. 709 4B; MGC-IGVR 47789, Fig. 14), in some cases preserving anatomical details like the caudal SC 710 flexion (e.g., MPPSA-IGVR 43323, Fig. 13C). Other skeletal remains consist of segments of 711 articulated vertebral centra alternated with sectors of disarticulated vertebral centra that, however, 712 U are always scattered around the longitudinal axis of the shark body. There is no evidence of AN 713 preferential orientation of the skeletal elements, imbrication or accumulation caused by tractive M 714 currents or remarkable displacement due to water turbulence (Amalfitano et al., 2017b). Therefore, 715 these skeletal remains can be interpreted as representing various degrees of disarticulation of slowly D 716 decaying deadfall exposed for more or less long-lasting time intervals on the sea bottom under low TE 717 energy conditions (see in analogy Amalfitano et al., 2017b). 718 The more or less rapid decay of non-mineralized tissues, differential sinking in the EP 719 unconsolidated calcareous nannoplankton-foraminiferal ooze that characterized the sea floor during 720 the deposition of the ‘lastame’ and the different exposure intervals on the seafloor before burial C AC 721 could explain the different conditions of the skeletal remains, from the better preserved and fully 722 articulated vertebral columns, MPPSA-IGVR 45305 and MGC-IGVR 47789, to the largely 723 disarticulated vertebral column of MPPSA-IGVR 45308. Some articulated specimens, such as 724 IGVR 36371, 45305 and 47789 (Figs. 4, 14), show an arched arrangement of the vertebral column, 725 which could be explained as a possible product of tetany of the carcasses. The limited displacement 726 of adjacent vertebral centra in partially articulated skeletons was probably caused by the limited 727 decay of the thin fibrous intervertebral cartilages that firmly connect consecutive centra (Cappetta, ACCEPTED MANUSCRIPT 728 1987). Such limited disarticulation of the chondrichthyan axial structures was observed in cases of 729 experimental taphonomy of chondrichthyan material (e.g., Samson et al., 2013). Fully disarticulated 730 skeletons were probably exposed to the biological activity on the seafloor for longer periods. The 731 partial sinking of the carcasses in the carbonate ooze is supported by the presence of centra and 732 teeth partially or totally embedded in the rock and slightly displaced from the main plane of PT 733 arrangement of the skeletal remains on the bedding surface (e.g., MGP-PD 31960, Fig. 17, and RI 734 supplementary material Fig. A.1η). The centra exposing one of the articular surfaces may have had 735 slower sinking, floating for hydrostatic thrust, because they exposed the largest surface to the SC 736 underlying calcareous nannoplankton-foraminiferal ooze, and did not totally sink because of the 737 precocious lithification of the underlying substrate. 738 U AN 739 Bioerosive fossil traces M 740 The prolonged exposure of the carcasses on the seafloor discussed above is also supported 741 by the occurrence of bioerosional trace fossils on the vertebral centra. They were produced by the D 742 activity of organisms that colonized or fed on the deadfalls after partial or total consumption of their TE 743 soft tissues either by scavengers and/or bacteria. These traces have different morphologies and sizes 744 and are detectable only in two specimens, MPPSA-IGVR 36371 and MGP-PD 31960 (Fig. 18). The EP 745 traces on MPPSA-IGVR 36371 are located on the articular surfaces of some vertebral centra and 746 consist of very short, sharp and narrow grooves, V- or U-shaped in cross-section (length 1.5-26 mm C AC 747 and width 0.4-0.9 mm; Figs. 18A-B), with a lined or overlapping pattern, and clustered small 748 circular openings (diameter 0.2-1.4 mm; Figs. 18C-D), with a cylindrical section. The borings on 749 MGP-PD 31960 (Figs. 18E-F) are larger and different from those in MPPSA-IGVR 36371, with a 750 lenticular shape (diameter 1.4-3.7 mm), showing different depths, from superficial to 0.8 mm-deep 751 holes; they are also evidently clustered. 752 The short, sharp, and narrow grooves on some vertebral centra of the specimen MPPSA- 753 IGVR 36371 (e.g., Figs. 18A-B) could be interpreted as tooth marks produced by the scavenging ACCEPTED MANUSCRIPT 754 activity of other sharks or fishes on the deadfall. Many chondrichthyans or bony fishes usually feed 755 on deadfall, being the first stage of ecological succession in a deadfall community - the mobile- 756 scavenger stage (see Smith et al., 2015) - and there are several reports of such activity also in the 757 fossil record (e.g., Schwimmer et al., 1997). Damage on the vertebral centra due to preparation of 758 the specimen can be excluded, because the marks do not exhibit fresh fractures or scratches PT 759 typically caused by a preparation tool. RI 760 Small circular borings on MPPSA-IGVR 36371 vertebral centra are similar to the fossil 761 traces of Osedax, a marine worm (Siboglinidae, Anellida) thriving on vertebrate deadfalls (Rouse et SC 762 al., 2004). Among siboglinids, Osedax has developed a unique metazoan-bacteria symbiosis that 763 exploits the organic material sequestered within the bones of dead vertebrates as an energy source 764 U (Danise, Higgs, 2015). Osedax and Osedax-like fossil traces were found on diverse vertebrate AN 765 remains (e.g., whales, marine birds, bony fishes, marine turtles, and plesiosaurs; Danise, Higgs, M 766 2015; Muñiz et al, 2010; Kiel et al., 2010; Kiel al., 2011; Kiel et al., 2013); the oldest one dates 767 back to the Albian-Cenomanian (Danise, Higgs, 2015). There is no record of Osedax borings in D 768 chondrichthyans, and also experimental studies on extant vertebrate remains (e.g., Rouse et al., TE 769 2011) did not report colonization by Osedax on chondrichthyan remains. However, the use of 770 juvenile Isurus oxyrhinchus vertebrae (diameter 5 mm) in the experiment carried out by Rouse et al. EP 771 (2011) does not provide a definitive support to exclude any possible colonization of adult 772 chondrichthyan deadfalls, which show more calcified skeletal components and, thereby, represent a C AC 773 potential site for Osedax colonization. However, it is not possible to demonstrate whether these 774 borings were actually produced by Osedax worms or not, mostly because the vertebral centra of the 775 larger slabs are still embedded in the rock, thereby preventing a tomographic analysis (like that 776 carried out on other fossil vertebrate remains, e.g. Danise and Higgs, 2015). In any case, coalescent 777 Osedax borings originating from colonization of multiple individuals may often collapse into small 778 irregular pits (Danise S., pers. comm.) surrounded by some individual borings. This feature is ACCEPTED MANUSCRIPT 779 evident in two vertebral centra, supporting a tentative attribution to Osedax colonization and likely 780 representing the first one reported from chondrichthyan remains. 781 The borings on the bioeroded vertebra of specimen MGP-PD 31960 have a 782 Gastrochaenolites-like structure, representing only a superficial boring stage. In fact, these borings 783 resemble the borings produced by pholadid bivalves reported from fossil whale bones affected by PT 784 clavate bivalve borings (see Belaústegui et al., 2012 and references therein). Gastrochaenolites RI 785 borings were also described in coprolites and fish bones from Cretaceous-Paleogene phosphatic 786 conglomerates from Northeastern Mali (Tapanila et al., 2004). All the occurrences known so far SC 787 record borings on transported bone clasts. Belaústegui et al. (2012) provided evidence of the only 788 examples of putative pholadid colonization and borings on an autochthonous carcass after its 789 U deposition on the sea bottom and the removal of the soft tissue cover. However, all the considered AN 790 fossil traces come from shallow marine environments, while the fossil specimen described herein M 791 comes from a basinal setting. Moreover, the borings described here have no traces of the bioglyphs 792 typical of the Gastrochaenolites structures. As a consequence, we cannot ascertain the D 793 Gastrochaenolites affinities of the borings in the bioeroded vertebral centrum of specimen MGP-PD TE 794 31960, but it is the most similar ichnotaxon among those reported on vertebrate bones (see 795 Belaústegui et al., 2012, for a review about bioerosive traces on fossil vertebrate remains). It is EP 796 evident that these traces were produced by opportunistic organisms that colonized the bones or the 797 organically enriched sediments surrounding the carcass. C AC 798 799 Concluding remarks 800 Skeletal remains of lamniform sharks from the Upper Cretaceous of the Lessini Mountains 801 (Verona Province, northeastern Italy) are described. Seven of these skeletal remains can be reliably 802 referred to the ‘ginsu’ shark Cretoxyrhina mantelli based on the associated tooth sets. The 803 remaining specimens are tentatively attributed to ?Cretoxyrhina sp. because of the morphology of 804 the available skeletal elements and information on the fossil assemblage. ACCEPTED MANUSCRIPT 805 Three specimens (MPPSA-IGVR 36371, 45305, MGC-IGVR 81375-81376) were dated in 806 detail through calcareous plankton analysis. The specimens date back to the latest middle-upper 807 Turonian and represent some of the most complete remains of Cretoxyrhina mantelli reported from 808 northeastern Italy, previously known only from isolated teeth and a single partial skeleton reported 809 by Bassani (1888) from the Castellavazzo locality. PT 810 The estimated total lengths of the C. mantelli specimens have revealed that MPPSA-IGVR RI 811 36371 is probably the largest individual known to date of this taxon, attaining a maximum length of 812 615-650 cm. The longevity of two individuals was estimated by counting the incremental bands on SC 813 the vertebral centra resulting in values of about 26 years for the specimen MPPSA-IGVR 36371 and 814 21 years for the specimen MPPSA-IGVR 45305. 815 U The analysis of the morphology of the placoid scales supports the ecological niche of C. AN 816 mantelli as a fast pelagic hunting shark (sensu Reif, 1985) similar to the extant white shark M 817 Carcharodon carcharias. 818 The specimens of C. mantelli from the ʻlastameʼ lithofacies of the Scaglia Rossa Formation D 819 show features that allow to interpret the taphonomic processes that affected the carcasses before TE 820 their final burial. The disarticulation patterns and degree of preservation of some components more 821 prone to degradation (e.g., calcified cartilage) demonstrate that the deadfalls slowly decayed EP 822 exposed on the seafloor under low energy conditions for relatively long time and were visited by 823 different communities of scavenger and colonized by opportunistic feeders as demonstrated by the C AC 824 bioerosive fossil traces. On IGVR 36371 vertebral centra, it is possible to observe what appears to 825 be the first putative Osedax borings on a chondrichthyan skeleton as well as some bite marks made 826 by scavengers, while MGP-PD 31960 vertebral centra contain some peculiar lenticular borings 827 reminiscent of Gastrochaenolithes structures. The evidence of at least two stages of ecological 828 succession of deadfall community, the mobile-scavengers stage (testified by the bite marks) and the 829 enrichment-opportunist stage (i.e. the putative Osedax and lenticular borings) (see Smith et al, 830 2015), demonstrate that the C. mantelli deadfalls could have acted like modern whale deadfalls and ACCEPTED MANUSCRIPT 831 supported an heterotrophic community like other vertebrate carcasses did, as testified by other cases 832 in the Mesozoic fossil record (e.g., Kaim et al., 2008). 833 We also reviewed the nomenclatural history of C. mantelli, establishing that Agassiz (1835) 834 originally erected the species (under the name Lamna Mantellii) when referring to previously 835 published figures by Mantell (1822, plate 32). We also traced back four (PV OR 4524, 4527, 4539, PT 836 4540) of the original syntypes by comparing the figures of pl. 32 of Mantell (1822) with specimens RI 837 from the Mantellʼs Collection of The Natural History Museum, London. The Cretoxyrhina mantelli 838 skeletal remains described herein provide new relevant data to the knowledge of the ʻlastameʼ SC 839 vertebrate assemblage, which is dominated by chondrichthyan remains (15 skeletal remains out of a 840 total of 39 associated or articulated skeletal remains known so far), mostly Cretoxyrhina specimens. 841 U Recent studies (e.g., Amadori et al., 2019; Amalfitano et al., 2017a-c; Palci et al, 2013) have AN 842 revealed the diverse composition of this still poorly studied vertebrate fauna of the marine Upper M 843 Cretaceous of Italy, which is the object of a detailed in progress revision. 844 D 845 Acknowledgements TE 846 We thank Roberto Zorzin and †Anna Vaccari (MCSNV) and Mariagabriella Fornasiero (MGP-PD) 847 for the permission to study the specimens under their care. We are grateful also to Elisa Marchesini EP 848 (Associazione Culturale Officina 3) and Marta Castagna and Francesco Sauro (Associazione Museo C 849 dei fossili della Lessinia) for the help and the access to the specimens exhibited at the PPMSA and AC 850 GMC. Stefano Castelli (Department of Geosciences of University of Padova) is acknowledged for 851 his precious help with photographs. We are also deeply grateful to Guido Roghi (CNR-Padova) for 852 his precious help and photos. We thank Massimo Varese, Luca Deflorian and Daniela Vecchiato for 853 assistance in an early phase of study of some of the specimens here investigated. We deeply thank 854 Todd Cook and Mikael Siverson for their constructive and precious comments and suggestions. 855 This work was carried out in the context of a wide project aimed to study the vertebrate fauna of the 856 Scaglia Rossa Formation from the Veneto Region of NE Italy. Funding for this research was ACCEPTED MANUSCRIPT 857 provided by University of Padova (Progetto di Ateneo CPDA159701/2015 titled ʻReappraisal of 858 two key Fossil-Lagerstätten in Scaglia deposits of northeastern Italy in the context of Late 859 Cretaceous climatic variability: a multidisciplinary approachʼ, assigned to Eliana Fornaciari) and ex 860 60% (L.G.). The research of G.C. was also supported by grants (ex-60% 2017 and 2018) of the 861 Università degli Studi di Torino. PT 862 RI 863 References 864 SC 865 Agassiz, J.L.R., 1835. Rapport sur les poissons fossiles découverts depuis la publication de la U 866 troisième livraison. Feuilleton additionel sur les Recherches sur les poissons fossiles. AN 867 Quatrième livraison (January 1835), pp. 39-64. Imprimerie de Petitpierre et Prince (text), 868 Neuchâtel. M 869 Agassiz, J.L.R., 1838. Recherches sur les poissons fossiles, 11th livraison (November 1838). - 870 Petitpierre (text) and H. Nicolet (plates), Neuchâtel, vol. 2, pl. 42, 43; vol. 3, [73]-140, pl. D 871 1a, 8a, 8b, 15, 17, 19, 20, 24, 25b, 30-35, 39, 40; vol. 5, pl. 60; Feuilleton additionnel, 107- TE 872 116. EP 873 Agassiz, J.L.R., 1843, Recherches sur les poissons fossiles, 15th and 16th livraisons. - Jent et 874 Gassmann, Soleure (text) and H. Nicolet (plates), Neuchâtel, vol. 2, 2nd part, [1]-72, pl. B’, C 875 B’’, Ca, G, 23b, 23c, 31, 33a, 36-38, 42a, 44; vol. 3, 157-390, 382*-382**, 1-32, [33]-[34], AC 876 pl. 1, 18, 22, 22a-b, 26a, 38, 40b-d, 45, 47; vol. 4, pl. 23b; vol. 5, 2nd part, 57-84, pl. B, C, 877 E, H, J, K, 9, 10, 28, 29, 37b, 44-48; Feuilleton additionnel, 139-144. 878 Agassiz, J.L.R., 1833-1844. Recherches sur les poissons fossiles. 5 volumes. Imprimerie de 879 Petitpierre et Prince, Neuchâtel, 1420 pp. doi: dx.doi.org/10.3931/e-rara-11962 880 Amadori, M., Amalfitano, J., Giusberti, L., Fornaciari, E., Luciani, V., Carnevale, G., Kriwet, J., 881 2019. First associated tooth set of a high-cusped Ptychodus (Chondrichthyes, ACCEPTED MANUSCRIPT 882 Elasmobranchii) from the Upper Cretaceous of northeastern Italy, and resurrection of 883 Ptychodus altior Agassiz, 1835. Cretaceous Research 93, 330-345. 884 Amalfitano, J., Giusberti, L., Fornaciari, E., Carnevale, G., 2017a. A reappraisal of the Italian 885 record of the Cretaceous pachycormid fish Protosphyraena Leidy, 1857. Rivista Italiana di 886 Paleontologia e Stratigrafia 123(3), 475-485. PT 887 Amalfitano, J., Dalla Vecchia, F.M., Giusberti, L., Fornaciari, E., Luciani, V., Roghi, G., 2017b. RI 888 Direct evidence of trophic interaction between a large lamniform shark, Cretodus sp., and a 889 marine turtle from the Cretaceous of northeastern Italy. Palaeogeography, SC 890 Palaeoclimatology, Palaeoecology 469, 104-121. 891 Amalfitano, J., Giusberti, L., Dalla Vecchia, F.M., Kriwet, J., 2017c. First skeletal remains of the 892 U giant sawfish Onchosaurus (Neoselachii, Sclerorhynchiformes) from the Upper Cretaceous AN 893 of northeastern Italy. Cretaceous Research 69, 124-135. M 894 Applegate, S.P., 1967. A survey of shark hard parts. In: Gilbert, P.W., Mathewson, R.F., Rail, D.P. 895 (Eds.), Sharks, Skates, and Rays. John Hopkins University Press, Baltimore, 37-67. D 896 Aspes, A., Zorzin, R., 1984. Guida al Museo di Storia Naturale di S. Anna d'Alfaedo (VR). La TE 897 Grafica di Vago, Verona, 35 pp. 898 Astolfi, G., Colombara, F., 2003. La geologia dei Colli Euganei. Edizioni Canova, Padova, 238 pp. EP 899 Bassani, F., 1876. Ittiodontoliti del Veneto. Atti della Società Veneto-Trentina di Scienze Naturali 900 residente in Padova 5, 275-308. C AC 901 Bassani, F., 1886. Sull’età degli strati a pesci di Castellavazzo nel Bellunese. Bollettino della 902 Società Geologica Italiana 4, 143-148. 903 Bassani, F., 1888. Colonna vertebrale di Oxyrhina Mantelli, Agassiz, scoperta nel calcare senoniano 904 di Castellavazzo, nel Bellunese. Estratto dalle Memorie della Società Italiana delle Scienze 905 (detta dei XL) s. III (n. 1), 7, 1-5. ACCEPTED MANUSCRIPT 906 Belaústegui, Z., de Gibert, J.M., Domènech, R., Muñiz, F., Martinell, J., 2012. Clavate borings in a 907 Miocene cetacean skeleton from Tarragona (NE Spain) and the fossil record of marine bone 908 bioerosion. Palaeogeography, Palaeoclimatology, Palaeoecology 323, 68-74. 909 Bengtson, P., 1988. Open nomenclature. Palaeontology 31(1), 223-227. 910 Berg, L.S., 1937. A classification of fish-like vertebrates. Bulletin de l’Académie des Sciences de PT 911 l’URSS 4, 1277-1280. RI 912 Bonaparte, C.L., 1838. Selachorum tabula analytica. Nuovi Annali delle Scienze Naturali 1 (2), 913 195-214. SC 914 Bourdon, J., Everhart, M.J., 2011. Analysis of an associated Cretoxyrhina mantelli dentition from 915 the Late Cretaceous (Smoky Hill Chalk, Late Coniacian) of western Kansas. Transactions of 916 U the Kansas Academy of Science 114(2), 15-32. AN 917 Brignon, A., 2015. Senior synonyms of Ptychodus latissimus Agassiz, 1835 and Ptychodus M 918 mammillaris Agassiz, 1835 (Elasmobranchii) based on teeth from the Bohemian Cretaceous 919 Basin (the Czech Republic). Acta Musei Nationalis Pragae, Series B-Historia Naturalis D 920 71(1-2), 5-14. TE 921 Burnett, J.A. (with contributions from Gallagher, L.T., Hampton, M.J.), 1998. Upper Cretaceous. 922 In: Bown, P.R. (Ed.), Calcareous Nannofossil Biostratigraphy. Kluwer Academic EP 923 Publishing, Dordrecht, Netherlands, 132-199. 924 Cailliet, G.M., Goldman, K.J., 2004. Age determination and validation in chondrichthyan fishes, C AC 925 Chapter 14. In: Carrier, J., Musick, J.A., Heithaus M.R. (Eds.). Biology of Sharks and Their 926 Relatives. CRC Press, Boca Raton, Florida, 399-447. 927 Cappetta, H., 1987. Chondrichthyes II. Mesozoic and Cenozoic Elasmobranchii. In: Schultze, H.-P. 928 (Ed.), Handbook of Paleoichthyology, vol. 3B. Gustav Fischer Verlag, Stuttgart, 193 pp. 929 Cappetta, H., 2012. Chondrichthyes. Mesozoic and Cenozoic Elasmobranchii: Teeth. In: Schultze, 930 H.-P. (Ed.), Handbook of Paleoichthyology, vol. 3E. Verlag Dr. Friedrich Pfeil, Munich, 931 512 pp. ACCEPTED MANUSCRIPT 932 Chesi, F., Delfino, M., 2007. The Italian fossil record of the sea turtles. In: Bologna, M.A., Capula, 933 M., Carpaneto, G.M., Luiselli, L., Marangoni, C., Venchi, A. (Eds.), Atti del 6° Congresso 934 Nazionale della Societas Herpetologica Italica (Roma, 27 Settembre-1 Ottobre 2006). 935 Edizioni Belvedere, Latina, 95-116. 936 Cigala Fulgosi, F., Kotsakis, T., Massari, F., Medizza, F., Sorbini, L., 1980. Il giacimento di S. PT 937 Anna d'Alfaedo. In: Sorbini, L. (Ed.), I vertebrati fossili italiani, Catalogo della Mostra, RI 938 Verona, 123-129. 939 Coccioni, R., Premoli Silva, I., 2015. Revised UpperAlbian-Maastrichtian planktonic foraminiferal SC 940 biostratigraphy and magnetostratigraphy of the classical Tethyan Gubbio section (Italy). 941 Newsletters on Stratigraphy, 48(1), 47-90. 942 U Colombara, F., 2013. Pietre e Marmi del Veneto. Geologia, arte, storia. Edizioni Papergraph, AN 943 Piazzola sul Brenta (Padova), 189 pp. M 944 Colleselli, E., Cornale, P., Siorpaes, C., 1997. Ambiente geologico e caratteri petrografici del 945 materiale. In: Alpago Novello, A. (Ed.), Castellavazzo. Un paese di pietra, la pietra di un D 946 paese. Neri Pozza, 31-44. TE 947 Compagno, L.J.V., 1973. Interrelationships of living elasmobranches. In: Greenwood, P.H., Miles, 948 R.S., Patterson, C. (Eds.), Interrelationships of Fishes. Zoological Journal of the Linnean EP 949 Society 53, suppl. 1. London, 15-61. 950 Compagno, L.J.V., 1977. Phyletic relationships of living sharks and rays. American Zoologist 17, C AC 951 303-322. 952 Cook, T.D., Newbrey, M.G., Murray, A.M., Wilson, M.V., Shimada, K., Takeuchi, G.T., Stewart, 953 J.D., 2011. A partial skeleton of the Late Cretaceous lamniform shark, Archaeolamna 954 kopingensis, from the Pierre Shale of western Kansas, USA. Journal of Vertebrate 955 Paleontology 31(1), 8-21. ACCEPTED MANUSCRIPT 956 Cook, T.D., Wilson, M.V., Murray, A.M., Plint, A.G., Newbrey, M.G., Everhart, M.J., 2013. A 957 high latitude euselachian assemblage from the early Turonian of Alberta, Canada. Journal of 958 Systematic Palaeontology 11(5), 555-587. 959 Dalla Vecchia, F.M., Barbera, C., Bizzarrini, F., Bravi, S., Delfino, M., Giusberti, L., Guidotti, G., 960 Mietto, P., Papazzoni, C., Roghi, G., Signore, M., Simone, O., 2005. Il Cretaceo marino. In: PT 961 Bonfiglio, L. (Ed.), Paleontologia dei Vertebrati in Italia. Memorie del Museo Civico di RI 962 Storia Naturale di Verona, Sezione Scienze della Terra 6, Verona, 101-116. 963 Danise, S., Higgs, N.D., 2015. Bone-eating Osedax worms lived on Mesozoic marine reptile SC 964 deadfalls. Biology letters 11(4), 20150072. 965 Dean, M.N., Summers, A.P., 2006. Mineralized cartilage in the skeleton of chondrichthyan fishes. 966 Zoology 109(2), 164-168. U AN 967 D'Erasmo, G., 1922. Catalogo dei pesci fossili delle Tre Venezie. Memorie dell'Istituto di Geologia M 968 della Regia Università di Padova 6, 1-181. 969 De Zigno, A., 1883. Sui vertebrati fossili dei terreni mesozoici delle Alpi Venete. Memorie della D 970 Regia Accademia di Scienze, Lettere ed Arti di Padova 9 (1883), 1-12. TE 971 Diedrich, C.G., 2014. Skeleton of the Fossil Shark Isurus denticulatus from the Turonian (Late 972 Cretaceous) of Germany—Ecological Coevolution with Prey of Mackerel Sharks. EP 973 Paleontology Journal, Article ID 934235. 974 Eastman, C.R., 1894. Beiträge Kenntniss Gattung Oxyrhina mit besonder Berücksichtigung von C AC 975 Oxyrhina mantelli von Agassiz. Paleontographica 41, 149-192. 976 Everhart, M.J., 2017. Oceans of Kansas. 2nd Edition. Indiana University Press, Bloomington, 427 977 pp. 978 Ferrón, H.G., 2017. Regional endothermy as a trigger for gigantism in some extinct macropredatory 979 sharks. PLoS ONE 12(9), e0185185. ACCEPTED MANUSCRIPT 980 Glickman, L.S., 1957. [Genetic Relations of the Lamnidae and Odontaspidae and new genera of 981 lamnids from the Upper Cretaceous]. Trudy Geologicheskogo Muzeia Akademiia Nauk 982 SSSR 1, 110-117. (in Russian). 983 Glikman, L.S., 1958. [Rates of evolution in lamnoid sharks]. Doklady Akademii Nauk SSSR 123, 984 568-571. (in Russian). PT 985 Goldman, K.J., Cailliet, G.M., Andrews, A.H., Natanson, L.J., 2012. Assessing the age and growth RI 986 of chondrichthyan fishes. In: Carrier, J.C., Musick, J.A.,. Heithaus, M.R (Eds.). Biology of 987 Sharks and their Relatives. Second Edition. CRC Press, Boca Raton, Florida, Chapter 14, SC 988 423-451. 989 Hay, O.P., 1902. Bibliography and catalogue of the fossil Vertebrata of North America. Bulletin of 990 U the US Geological Survey. US Government Printing Office 179, Washington, p. 882. AN 991 Hone, D.W.E., Witton, M.P., Habib, M.B., 2018. Evidence for the Cretaceous shark Cretoxyrhina M 992 mantelli feeding on the pterosaur Pteranodon from the Niobrara Formation. PeerJ 6, e6031. 993 Huxley, T., 1880. A Manual of the Anatomy of Vertebrated Animals. D. Appleton, New York. D 994 Kaim, A., Kobayashi, Y., Echizenya, H., Jenkins, R.G., Tanabe, K., 2008. Chemosynthesis-based TE 995 associations on Cretaceous plesiosaurid carcasses. Acta Palaeontologica Polonica 53(1), 97- 996 104. EP 997 Kiel, S., Goedert, J.L., Kahl, W.A., Rouse, G.W., 2010. Fossil traces of the bone-eating worm 998 Osedax in early Oligocene whale bones. Proceedings of the National Academy of Sciences C AC 999 107(19), 8656-8659. 1000 Kiel, S., Kahl, W.A., Goedert, J.L., 2011. Osedax borings in fossil marine bird bones. 1001 Naturwissenschaften 98(1), 51-55. 1002 Kiel, S., Kahl, W.A., Goedert, J.L., 2013. Traces of the bone-eating annelid Osedax in Oligocene 1003 whale teeth and fish bones. Paläontologische Zeitschrift 87(1), 161-167. 1004 Kim, S.H., Shimada, K., Rigsby, C.K., 2013. Anatomy and evolution of heterocercal tail in 1005 lamniform sharks. The Anatomical Record 296(3), 433-442. ACCEPTED MANUSCRIPT 1006 Kriwet, J., Mewis, H., Hampe, O., 2015. A partial skeleton of a new lamniform mackerel shark 1007 from the Miocene of Europe. Acta Palaeontologica Polonica 60 (4), 857–875 1008 Lioy, P., 1865. Sopra alcuni avanzi di Plagiostomi fossili del Vicentino e specialmente 1009 sull'Alopiopsis Plejodon, Lioy (Galeus Cuvieri, Ag.). Atti della Società Italiana di Scienze 1010 Naturali 8, 398-409. PT 1011 Lirer, F., 2000. A new technique for retrieving calcareous microfossils from lithified lime deposits. RI 1012 Micropaleontology 46 (4), 365-369. 1013 Lozar, F., Grosso, F., 1997. Biostratigrafia della successione cretacica del margine dei Lessini SC 1014 occidentali (Provincia di Verona, Italia). Bollettino del Museo Regionale di Scienze Naturali 1015 di Torino 15 (1), 111-136. 1016 U Maisey, J.G., 2013. The diversity of tessellated calcification in modern and extinct chondrichthyans. AN 1017 Revue de Paléobiologie 32, 355-371. M 1018 Mantell G., 1822. The fossils of the South Downs; or illustrations of the geology of Sussex. Lupton 1019 Relfe, London, XIV + [XV-XVI] + 328 pp. http://doi.org/10.3931/e-rara-16021 D 1020 Marramà, G., Carnevale, G., Kriwet, J., 2017. New observations on the anatomy and paleobiology TE 1021 of the Eocene requiem shark †Eogaleus bolcensis (Carcharhiniformes, Carcharhinidae) from 1022 Bolca Lagerstätte, Italy. Comptes Rendus Palevol 17, 443-459. EP 1023 Massari, F., & Savazzi, E., 1981. Driftwood transportation of exotic pebbles in the Upper 1024 Cretaceous Scaglia Rossa veneta (Mt. Loffa, Southern Alps) suggested by Teredinid tubes. C AC 1025 Neues Jahrbuch für Geologie und Paläontologie Monatshefte 5, 311-320. 1026 Matthews, S. C., 1973. Notes on open nomenclature and on synonymy list. Palaeontology 16, 713- 1027 719. 1028 Muñiz, F., De Gibert, J.M., Esperante, R., 2010. First trace-fossil evidence of bone-eating worms in 1029 whale carcasses. Palaios 25(4), 269-273. ACCEPTED MANUSCRIPT 1030 Newbrey, M.G., Siverson, M., Cook, T.D., Fotheringham, A.M., Sanchez, R.L., 2015. Vertebral 1031 morphology, dentition, age, growth, and ecology of the large lamniform shark Cardabiodon 1032 ricki. Acta Palaeontologica Polonica 60(4), 877-897. 1033 Ogg, J.G., Hinnov, L., 2012. The Cretaceous Period. In: Gradstein, F.M., Ogg, J.G., Shmitz M.D., 1034 Ogg, G.M. (Eds.), The Geological Time Scale 2012, Elsevier, 793-853. PT 1035 Palci, A., Caldwell, M.W., Papazzoni, C.A., 2013. A new genus and subfamily of mosasaurs from RI 1036 the Upper Cretaceous of northern Italy. Journal of Vertebrate Paleontology 33(3), 599-612. 1037 Pirrone, C.A., Buatois, L.A., Bromley, R.G., 2014. Ichnotaxobases for bioerosion trace fossils in SC 1038 bones. Journal of Paleontology 88(1), 195-203. 1039 Reif, W.E., 1985. Squamation and ecology of sharks. Courier Forschungsinstitut Senckenberg 78, 1040 1-255. U AN 1041 Reif, W.E., Dinkelacker, A., 1982. Hydrodynamics of the squamation in fast swimming sharks. M 1042 Neues Jarbuch für Geologie und Pälaontologie, Abhandlungen 164: 184-187. 1043 Ride, W.D.L, Cogger, H.G., Dupuis, C., Kraus, O., Minelli, A., Thompson, F.C., Tubbs, P.K., D 1044 (eds.)., 2000. International Code of Zoological Nomenclature. 4th edition. The International TE 1045 Trust for Zoological Nomenclature 1999, London, 306 pp. 1046 Rouse, G.W, Goffredi, S.K., Vrijenhoek, R.C., 2004. Osedax: bone-eating marine worms with EP 1047 dwarf males. Science 305, 668-671. 1048 Rouse, G.W., Goffredi, S.K., Johnson, S.B., Vrijenhoek, R.C., 2011. Not whale-fall specialists, C AC 1049 Osedax worms also consume fishbones. Biology letters 7(5), 736-739. 1050 Sansom, R.S., Gabbott, S.E., Purnell, M.A., 2013. Atlas of vertebrate decay: a visual and 1051 taphonomic guide to fossil interpretation. Palaeontology 56(3), 457-474. 1052 Schwimmer, D.R., Stewart, J.D., Williams, G.D., 1997. Scavenging by sharks of the genus 1053 Squalicorax in the Late Cretaceous of North America. Palaios 12, 71-83. 1054 Shimada K., 1994c. Scales of the Late Cretaceous shark, Cretoxyrhina mantelli, from western 1055 Kansas. Journal of Vertebrate Paleontology 14 (Suppl. to no. 3), p. 46A. ACCEPTED MANUSCRIPT 1056 Shimada, K., 2002. Dental homologies in lamniform sharks (Chondrichthyes: Elasmobranchii). 1057 Journal of Morphology 251(1), 38-72. 1058 Shimada, K., 2005. Types of tooth sets in the fossil record of sharks, and comments on 1059 reconstructing dentitions of extinct sharks. Journal of Fossil Research 38 (2), 141-145. 1060 Shimada, K., 2007. Skeletal and dental anatomy of lamniform shark, Cretalamna appendiculata, PT 1061 from Upper Cretaceous Niobrara Chalk of Kansas. Journal of Vertebrate Paleontology RI 1062 27(3), 584-602. 1063 Shimada, K., 2008. Ontogenetic parameters and life history strategies of the Late Cretaceous SC 1064 lamniform shark, Cretoxyrhina mantelli, based on vertebral growth increments. Journal of 1065 Vertebrate Paleontology 28(1), 21-33. 1066 U Shimada, K., 1993a. Preliminary report on the Upper Cretaceous shark, Cretoxyrhina mantelli, from AN 1067 western Kansas. Journal of Vertebrate Paleontology 13 (Suppl. to no. 3), p. 57A. M 1068 Shimada, K., 1993b. Jaws of the Upper Cretaceous shark, Cretoxyrhina mantelli, from western 1069 Kansas. Geological Society of America (Abstract with Programme) 25(6), p. 58. D 1070 Shimada, K., 1994a. Functional morphology of the dentition of the Late Cretaceous shark, TE 1071 Cretoxyrhina mantelli. Abstract from the annual meeting of the Kansas Academy of 1072 Sciences 13, p. 35. EP 1073 Shimada, K., 1994b. Paleobiology of the Late Cretaceous shark, Cretoxyrhina mantelli 1074 (Lamniformes: Cretoxyrhinidae), from Kansas (Unpublished M.S. thesis). Hays, Kansas, C AC 1075 Fort Hays State University, 169 pp. 1076 Shimada, K., 1997a. Dentition of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli, from 1077 the Niobrara Chalk of Kansas: Journal of Vertebrate Paleontology 17, 269-279. 1078 Shimada, K., 1997b. Paleoecological relationships of the Late Cretaceous lamniform shark, 1079 Cretoxyrhina mantelli (Agassiz): Journal of Paleontology 71, 926-933. 1080 Shimada, K., 1997c. Periodic marker bands in vertebral centra of the Late Cretaceous lamniform 1081 shark, Cretoxyrhina mantelli. Copeia 1, 233-235. ACCEPTED MANUSCRIPT 1082 Shimada, K., 1997d. Skeletal anatomy of the Late Cretaceous lamniform shark, Cretoxyrhina 1083 mantelli, from the Niobrara Chalk in Kansas: Journal of Vertebrate Paleontology 17, 642- 1084 652. 1085 Shimada, K., 1997e. Stratigraphic record of the Late Cretaceous lamniform shark, Cretoxyrhina 1086 mantelli (Agassiz), Kansas: Transactions of Kansas Academy of Science 100, p. 139-149. PT 1087 Shimada, K., Hooks III, G.E., 2004. Shark-bitten protostegid turtles from the Upper Cretaceous RI 1088 Mooreville Chalk, Alabama. Journal of Paleontology, 78(1), 205-210. 1089 Shimada, K., Cumbaa, S.L., Van Rooyen, D., 2006. Caudal fin skeleton of the Late Cretaceous SC 1090 shark, Cretoxyrhina mantelli (Lamniformes: Cretoxyrhinidae) from the Niobrara Chalk of 1091 Kansas. Bulletin of the New Mexico Museum of Natural History 35, 185-192. 1092 U Sigovini, M., Keppel E., Tagliapietra, D., 2016. Open Nomenclature in the biodiversity era. AN 1093 Methods in Ecology and Evolution 7, 1217-1225. M 1094 Siverson, M., 1992. Biology, dental morphology and taxonomy of lamniform sharks from the 1095 Campanian of the Kristianstad Basin, Sweden. Palaeontology 35, 519-554. D 1096 Siverson, M., 1996. Lamniform sharks of the mid Cretaceous Alinga Formation and Beedagong TE 1097 Claystone, western Australia. Palaeontology 39, 813-849. 1098 Siverson, M., Lindgren, J., 2005. Late Cretaceous sharks Cretoxyrhina and Cardabiodon from EP 1099 Montana, USA. Acta Palaeontologica Polonica 50, 301-334. 1100 Siverson, M., Ward, D.J., Lindgren, J., Kelley, L.S., 2013. Mid-Cretaceous Cretoxyrhina C AC 1101 (Elasmobranchii) from Mangyshlak, Kazakhstan and Texas, USA. Alcheringa: An 1102 Australasian Journal of Palaeontology 37(1), 87-104. 1103 Smith, C.R., Glover, A.G., Treude, T., Higgs, N D., Amon, D. J., 2015. Whale-fall ecosystems: 1104 recent insights into ecology, paleoecology, and evolution. Annual Review of Marine Science 1105 7, 571-596. 1106 Sorbini, L., 1976. L'ittiofauna cretacea di Cinto Euganeo (Padova-Nord Italia). Bollettino del 1107 Museo Civico di Storia Naturale di Verona 3, 479-567. ACCEPTED MANUSCRIPT 1108 Sorbini L., 1978. Amatori di fossili e recenti scoperte paleontologiche in provincia di Verona. La 1109 Lessinia, Ieri, Oggi e Domani. Quaderno Culturale n. 1 1978, 65-70. 1110 Tapanila, L., Roberts, E.M., Bouaré, M.L., Sissoko, F., O'Leary, M.A., 2004. Bivalve borings in 1111 phosphatic coprolites and bone, Cretaceous-Paleogene, northeastern Mali. Palaios 19, 565- 1112 573. PT 1113 Trevisani, E., 2009. New shark from the Upper Cretaceous of the Venetian Prealps (Castellavazzo, RI 1114 Belluno province, Northern Italy). In: International Conference on Vertebrate 1115 Palaeobiogeography and continental bridges across Tethys, Mesogea, and Mediterranean SC 1116 Sea. Abstract Book, p. 89-92. 1117 Trevisani, E., 2011. Resti di uno squalo lamniforme dal marmo di Castellavazzo (Cretaceo 1118 U Superiore, Castellavazzo, provincia di Belluno). Frammenti, conoscere e tutelare la natura AN 1119 bellunese 3, 87- 92. M 1120 Underwood, C.J., Cumbaa, S.L., 2010. Chondrichthyans from a Cenomanian (Late Cretaceous) 1121 bonebed, Saskatchewan, Canada. Palaeontology 53(4), 903-944. D 1122 Welton, B.J., Farish, R.F., 1993. The Collector’s Guide to Fossil Shark and Rays from the TE 1123 Cretaceous of Texas. Before Time Edition, Lewisville, 204 pp. 1124 Woodward, A.S., 1889. Catalogue of the Fossil Fishes in the British Museum (Natural History). EP 1125 Part I containing the Elasmobranchii. London, Printed by order of the Trustees, 1889-1901. 1126 474 pp. C AC 1127 Woodward, A.S., 1911. The Fossil Fishes of the English Chalk. Part VI. Palaeontographical 1128 Society, London 63, 185-224. 1129 Woodward, A.S., Sherborn, C.D., 1890. Dates of publication of 'Recherches sur les Poissons 1130 fossiles'…par L. Agassiz (contributed by Brown, W. H.) [xxv-xxix]. In: Woodward AS, 1131 Sherborn CDA Catalogue of British Fossil Vertebrata, London, Dulau & Co., 37 Soho 1132 Square, W. ACCEPTED MANUSCRIPT 1133 Zhelezko, V.I., 2000. The evolution of the tooth system of the shark genus of Pseudoisurus 1134 Glikman, 1957. Materials for the stratigraphy and paleontology of the Urals 4, 136-141. 1135 Zorzin, R., 2001. Fossili. Per conoscere il nostro passato. Demetra S.r.l., Verona, 128 pp. 1136 PT RI U SC AN M D TE C EP AC ACCEPTED MANUSCRIPT 1137 Figure captions 1138 Fig. 1. Location of the sites. The sites that yielded the skeletal remains of Cretoxyrhina mantelli 1139 described herein (Sant'Anna D'Alfaedo, Verona province) and Castellavazzo, near Longarone 1140 (Belluno province) are the only other site that yielded Cretoxyrhina mantelli skeletons. Scale bar = PT 1141 100 km. [single-column width] 1142 Fig. 2. Cretoxyrhina mantelli (Agassiz, 1835). Isolated teeth housed in the historical collections RI 1143 of the Museum of Geology and Paleontology of the University of Padova. A. MGP-PD 5404 as SC 1144 figured by D'Erasmo (1922: pl. 3, figs. 4-6), coming from Spilecco (VR) and erroneously attributed 1145 to the Eocene. B. photos of MGP-PD 5404 in lingual, lateral and labial view. C. MGP-PD 6721 as U 1146 figured by D'Erasmo (1922: pl. 3, fig. 1). D. photos of MGP-PD 6721 in lingual, lateral and labial AN 1147 view (Achille De Zigno’s collection). E. MGP-PD 6736 as figured by D'Erasmo (1922: pl. 3, figs. 1148 2-3). F. photos of MGP-PD 6736 in lingual, lateral and labial view (Achille De Zigno’s collection). M 1149 Scale bars = 10 mm. [2-column width] D 1150 Fig. 3. Syntypes of Cretoxyrhina mantelli (Agassiz, 1835). A. Isolated teeth figured by Mantell TE 1151 (1822: pl. 32, figs. 4, 7, 8, 10, 11, 26, 28) to which Agassiz (1835, p. 54) referred when created 1152 Lamna Mantellii (subsequently Oxyrhina mantelli and Cretoxyrhina mantelli). Woodward (1889) EP 1153 identified the teeth illustrated by Mantell (1822) in fig. 8 (NHM PV OR 4539), 11 (NHM PV OR 1154 4540), 26 (NHM PV OR 4527) and 28 (NHM PV OR 4524) of plate 32. Excerpt of the plate 32 of C 1155 Mantell (1822). ETH-Bibliothek Zürich, Rar 2452, http://doi.org/10.3931/e-rara-16021/Public AC 1156 Domain Mark. B-E. Photos of four of the seven original syntypes of Cretoxyrhina mantelli 1157 (Agassiz, 1835). B. NHM PV OR 4524. C. NHM PV OR 4527. D. NHM PV OR 4539. E. NHM 1158 PV OR 4540. Scale bar = 10 mm. Collections of the Natural History Museum, London (CC-BY). 1159 Photos courtesy of Natural History Museum, London. Dataset: Collection specimens. Resource: 1160 Specimens. Natural History Museum Data Portal (data.nhm.ac.uk). 1161 https://doi.org/10.5519/0002965. Retrieved: 10:32 25 Jul 2018 (GMT). [single-column width] ACCEPTED MANUSCRIPT 1162 Fig. 4. Cretoxyrhina mantelli (Agassiz, 1835). Orthophotos of the specimens MPPSA-IGVR 1163 36371 and 45305. A. MPPSA-IGVR 36371. B. MPPSA-IGVR 45305. Scale bar = 100 cm. [2- 1164 column width] 1165 Fig. 5. Cretoxyrhina mantelli (Agassiz, 1835). Teeth of the specimen MPPSA-IGVR 36371. A. 1166 First upper anterior tooth (A1) (left palatoquadrate), labial view. B. Second lower anterior tooth (a2) PT 1167 (left Meckel’s cartilage), lingual view. C. Intermediate tooth/third lower anterior? tooth (i?/a3), RI 1168 labial view. D. First upper lateral tooth (L1) (right palatoquadrate), lingual view. E. Second upper 1169 lateral tooth (L2) (right palatoquadrate), lingual view. F. Lateral tooth (L/l?), labial view). Arrows SC 1170 indicate the small oblique heel (or shoulders). Scale bars = 10 mm. [2-column width] 1171 Fig. 6. Cretoxyrhina mantelli (Agassiz, 1835). Tessellated calcified cartilage of MPPSA-IGVR 1172 U 36371. Photo of a fragment of cranial calcified cartilage associated to a tooth. Scale bar = 50 mm. AN 1173 [single-column width] M 1174 Fig. 7. Cretoxyrhina mantelli (Agassiz, 1835). SEM images of placoid scale from MPPSA- 1175 IGVR 36371. A. Frontal view. B. Lateral view. C. Posterior view. D. Anterior view. E. Posterior D 1176 view. F. Inferior view. G. Anterior view. H. Anterior view. I. Posterior view. J. Superior view. K. TE 1177 Frontal view. L. Posterior view. M. Superior view. N. Superior view. O. Superior view. P. Lateral 1178 view. Q. Lateral view. R. Anterior view. Scale bars = 100 µm. [single-column width] EP 1179 Fig. 8. Line drawing of the head region of the specimen of Cretoxyrhina mantelli (Agassiz, 1180 1835) MPPSA-IGVR 45305. Colors map: dark gray = vertebral centra; gray = teeth; light gray = C AC 1181 rocky matrix; white = fragments of tessellated cartilage. The barred area represents a glued 1182 fragment which contains the counterpart of the cranial cartilage indicated by the arrow. The dashed 1183 line delimits the area where can be detected the fragments (tesserae) of cranial calcified cartilage. 1184 Scale bar = 100 cm. [2-column width] 1185 Fig. 9. Cretoxyrhina mantelli (Agassiz, 1835). Teeth from MPPSA-IGVR 45305. A. Teeth 1186 articulated to a fragment of tessellated mandibular cartilage. B. Two lateral teeth with poorly 1187 preserved roots. C-D. Partially associated tooth sets. Scale bars = 10 mm. [single column width] ACCEPTED MANUSCRIPT 1188 Fig. 10. Cretoxyrhina mantelli (Agassiz, 1835). MPPSA-IGVR 45344-45345. A. MPPSA-IGVR 1189 45345 (left slab) and MPPSA-IGVR 45344 (right counterslab). Scale bar = 500 mm. B. Detail of 1190 the labial side of the teeth. Scale bar = 10 mm. C. Detail of the labial side of teeth. Scale bar = 10 1191 mm. D. Detail of teeth on the lingual side. Scale bar = 10 mm. [2-column width] 1192 Fig. 11. Cretoxyrhina mantelli (Agassiz 1835). MGC-IGVR 81375. A. Photo of MGC-IGVR PT 1193 81375. B. Line drawing of MGC-IGVR 81375. Colors map: dark gray = vertebral centra; gray = RI 1194 teeth; light gray = rocky matrix; white = fragments of tessellated cartilage. Dashed lines delimit 1195 imprints of teeth and vertebral centra. Scale bars = 500 mm. [1,5-column width] SC 1196 Fig. 12. Cretoxyrhina mantelli (Agassiz, 1835). Placoid scale from MGC-IGVR 36371. SEM 1197 imaging of selected placoid scales, from different views, coming from the residual of detached 1198 U samples. A-D. Anterior views. E-H. Lateral views. I. Anterior view. J-K. Posterior views. L. AN 1199 Inferior view. [single column width] M 1200 Fig. 13. ?Cretoxyrhina sp. Incomplete vertebral columns. A. Orthophoto of MPPSA-IGVR 1201 45308. B. Photo of MPPSA-IGVR 45319 (right) and MPPSA-IGVR 45320 (left). C. Photo of D 1202 MPPSA-IGVR 45323. D. Orthophoto of MGP-PD 31960. The asterisk indicates the vertebral TE 1203 centrum removed and showing bioerosive fossil traces (see Taphonomic remarks and Figs. 18E-F). 1204 Scale bars = 500 mm. [1,5-column width] EP 1205 Fig. 14. ?Cretoxyrhina sp. Vertebral column. Photo of MGC-IGVR 47789. Scale bar = 100 cm. 1206 [single column width] C AC 1207 Fig. 15. Cretoxyrhina mantelli (Agassiz, 1835). Count of incremental bands of individuals 1208 MPPSA-IGVR 36371 and 45305. A. Vertebral centrum from MPPSA-IGVR 36371. The black dot 1209 indicates the vertebral fulcrum, the black stars indicate the incremental bands (26). B. Vertebral 1210 centrum from MPPSA-IGVR 45305. The black dot indicates the vertebral fulcrum, the black stars 1211 indicate the incremental bands (21). Scale bars = 10 mm. [1,5-column width] 1212 Fig. 16. Correlation diagram of scale crown width (x axis) and ridge distance (y axis) (in µm). 1213 The black dashed line indicates the slope of 1,0. The red labeled taxa are from the group of fast ACCEPTED MANUSCRIPT 1214 pelagic hunting sharks, while the blue labeled taxa are from the group of large nearshore 1215 predators/moderate speed pelagic predators. Note that the Cretoxyrhina mantelli average (green 1216 star) falls in the clouds of correlation of fast pelagic hunting sharks, in the lower part of the 1217 diagram. The diagram is modified after Reif (1985). [single column width] 1218 Fig. 17. ?Cretoxyrhina sp. MGP-PD 31960. Photo of a section of a vertebral centrum fully PT 1219 embedded in the rocky matrix. Scale bar = 50 mm. [single column width] RI 1220 Fig. 18. Cretoxyrhina mantelli (Agassiz, 1835) and ?Cretoxyrhina sp. bioerosive fossil traces on 1221 vertebral centra. A. Photo of a vertebra with short grooves from MPPSA-IGVR 36371. B. Same SC 1222 as in A, with the grooves highlighted by dashed lines. C. Photo of small circular openings and 1223 collapsed areas on a vertebra from IGVR 36371, indicated by the black arrows, here interpreted as 1224 U putative Osedax borings. D. Another photo of a vertebral centrum from MPPSA-IGVR 36371 with AN 1225 putative Osedax borings. E. Photo of a vertebra from MGP-PD 31960, with lenticular borings with M 1226 a Gastrochaenolites-like structure. F. Detail of the lenticular borings from the vertebra of MGP-PD 1227 31960. Scale bars = 10 mm. [2-column width] D 1228 TE 1229 In-text table captions 1230 Tab. 1. List of the specimens examined during the study. Institutional abbreviations: MCSNV: EP 1231 Civic Museum of Natural History of Verona, Italy; MGC: Geopaleontological Museum of C 1232 Camposilvano, Italy; MGP-PD: Museum of Geology and Paleontology, University of Padova, AC 1233 Italy, C: Catullo’s Collection, Z: De Zigno’s Collection; MPPSA: Prehistoric and Paleontological 1234 Museum of S. Anna d'Alfaedo, Italy. The provenance locality reported in the historical labels of 1235 MGP-PD 14029 and 14034-14039 is ʻForni di Zoldoʼ (Belluno) but the lithology of the matrix 1236 embedding the specimens indicates that they come from the surroundings of Castellavazzo (ʻPietra 1237 di Castellazazzoʼ). ACCEPTED MANUSCRIPT 1238 Tab. 2. Biostratigraphic data from Cretoxyrhina mantelli specimens. Main calcareous plankton 1239 taxa and biostratigraphic classification of samples obtained from three partially articulated 1240 specimens of Cretoxyrhina mantelli (Agassiz, 1835). 1241 PT RI U SC AN M D TE C EP AC ACCEPTED MANUSCRIPT 1242 Jacopo Amalfitano conceived and designed the experiments, performed the experiments, analyzed 1243 the data, authored or reviewed drafts of the paper, prepared figures and/or tables, approved the final 1244 draft. 1245 1246 Luca Giusberti conceived and designed the experiments, performed the experiments, analyzed the 1247 data, contributed reagents/materials/analysis tools, authored or reviewed drafts of the paper, 1248 approved the final draft. 1249 PT 1250 Eliana Fornaciari performed the micropaleontological analyses, analyzed the data, contributed 1251 reagents/materials/analysis tools, authored or reviewed drafts of the paper, prepared supplementary 1252 tables, approved the final draft. RI 1253 1254 Fabio Marco Dalla Vecchia authored or reviewed drafts of the paper, approved the final draft. SC 1255 1256 Valeria Luciani performed the micropaleontological analyses, analyzed the data, authored or U 1257 reviewed drafts of the paper, prepared supplementary tables, approved the final draft. 1258 AN 1259 Giorgio Carnevale authored or reviewed drafts of the paper, approved the final draft. 1260 M 1261 Jürgen Kriwet authored or reviewed drafts of the paper, approved the final draft. 1262 D TE C EP AC ACCEPTED MANUSCRIPT Inventory number Preservation Provenance locality partial skeleton (teeth, articulated vertebrae, cranial cartilage fragments, MPPSA-IGVR 36371 Loffa Mount (S. Anna d'Alfaedo) placoid scales) partial skeleton (teeth, articulated vertebrae, cranial cartilage fragments, MPPSA-IGVR 45305 Loffa Mount (S. Anna d'Alfaedo) placoid scales) MPPSA-IGVR 45308 disarticulated vertebrae Loffa Mount (S. Anna d'Alfaedo) MPPSA-IGVR 45319-45320 portion of vertebral column Loffa Mount (S. Anna d'Alfaedo) MPPSA-IGVR 45321-45322 portion of vertebral column Loffa Mount (S. Anna d'Alfaedo) MPPSA-IGVR 45323 portion of vertebral column Loffa Mount (S. Anna d'Alfaedo) teeth associated with a portion of the vertebral column and fragments of MPPSA-IGVR 45324 Loffa Mount (S. Anna d'Alfaedo) cranial cartilage teeth associated with a portion of the vertebral column and fragments of MPPSA-IGVR 45326 Loffa Mount (S. Anna d'Alfaedo) cranial cartilage PT teeth associated with a portion of the vertebral column and fragments of MPPSA-IGVR 45334 Loffa Mount (S. Anna d'Alfaedo) cranial cartilage MPPSA-IGVR 45337 portion of vertebral column Loffa Mount (S. Anna d'Alfaedo) teeth associated with a portion of the vertebral column and fragments of MPPSA-IGVR 45344-45345 Loffa Mount (S. Anna d'Alfaedo) cranial cartilage RI MGC-IGVR 47789 articulated vertebral column Loffa Mount (S. Anna d'Alfaedo) partial anterior portion of skeleton (teeth, articulated vertebral centra, MGC-IGVR 81375-81376 Loffa Mount (S. Anna d'Alfaedo) cranial cartilage fragments, placoid scales) MGP-PD 3805 isolated tooth Castellavazzo? SC MGP-PD 5404 isolated tooth Spilecco (Verona) MGP-PD 6721 Z isolated tooth Mazzurega (Verona) MGP-PD 6736 Z isolated tooth Mazzurega (Verona) MGP-PD 7342 C isolated tooth Castellavazzo (Belluno) MGP-PD 7343 C isolated tooth Castellavazzo (Belluno) U MGP-PD 7372 C isolated tooth Mt. Belvedere (Belluno) MGP-PD 8498 C isolated tooth Castellavazzo (Belluno) MGP-PD 8889 isolated tooth Cerè Alto-Valdagno (Vicenza) AN MGP-PD 14020 isolated tooth Valle del Cismon (Belluno) MGP-PD 14029 isolated tooth Castellavazzo (Belluno) MGP-PD 14034-14039 isolated teeth Castellavazzo (Belluno) MGP-PD 14042 isolated tooth Castellavazzo (Belluno) MGP-PD 22401-22470 isolated teeth Castellavazzo (Belluno) M MGP-PD 23527-23529 isolated teeth Belluno province MGP-PD 31960 portion of vertebral column Loffa Mount (S. Anna d'Alfaedo) MCSNV V. 1094 isolated tooth Spilecco (Verona) MCSNV V. 1095 isolated tooth Spilecco (Verona) D MCSNV V. 11798 isolated tooth Spilecco (Verona) MCSNV V. 12518 isolated tooth Caprino Valley (Lubiara, Verona) MCSNV V. 12519 isolated tooth Prun (Verona) TE C EP AC ACCEPTED MANUSCRIPT PLANKTON BIOSTRATIGRAPHY NANNOs FORAMs SAMPLE DESCRIPTION LOCALITY LITHOFACIES Calcareous nannofossil assemblage Planktic foraminiferal assemblage (Burnett, 1998). (Coccioni & Premoli Silva, 2015) Marginotruncana sigali (RR); Dicarinella primitiva/ UC7-UC9 Zone. M. schneegansi (RR); M. renzi (F); Marginotruncana sigali Zone: Watznaueria spp. (C); Chiastozygus PT Concomitant presence of Q. M. pseudolinneiana (RR); occurrence of spp. (RR); Monte Loffa gartneri and absence of Micula M coronata (RR); marginotruncanids and Eiffellithus spp. (RR); (S. Anna Lastame (Scaglia staurophora. M. marianosi (RR); dicarinellids in absence of MPPSA-IGVR 36371 Partially articulated skeleton Eprolithus octopetalus (RR); d'Alfaedo, Rossa Fm.) The scarcity in calcareous Dicarinella Iimbricata (RR); Helvetoglobotruncana helvetica Quadrum gartneri (R) RI Verona) nannfossil content makes it hard Heterohelix spp. (C); and Dicarinella concavata, Quadrum intermedium (RR); to constrain even more the Muricohedbergella spp. (R); markers of the total range zones Prediscosphaera spp. (RR) biostratigraphic interval Macroglobigerinelloides spp. (R); underlying and overlying. Calcisphaerulids (A) SC Marginotruncana sigali (R); Watznaueria spp. (C/A); M. schneegansi (R); M. renzi (F); Dicarinella primitiva/ UC8b-UC9 Zone. Zeugrhabdotus bicrescenticus (R); M coronata (RR); Marginotruncana sigali Zone: Concomitant presence of L. Eprolithus octopetalus (RR); Dicarinella imbricata (RR); occurrence of quadrifidus and absence of Monte Loffa Chiastozygus spp. (RR); Retecapsa spp. D. canaliculata (F); marginotruncanids and U Micula staurophora. (S. Anna Lastame (Scaglia (R); Whiteinella sp. (RR) dicarinellids in absence of MPPSA-IGVR 45305 Partially articulated skeleton The vacancy of Lithastrinus d'Alfaedo, Rossa Fm.) Rhagodiscus achlyostaurion (RR); Heterohelix reussi(C); Helvetoglobotruncana helvetica septenarius could be an AN Verona) Lucianorhabdus quadrifidus (RR); H. moremani (RR); and Dicarinella concavata, ecological exclusion. Quadrum gartneri (RR); Muricohedbergella planispira (R); markers of the total range zones Hence, its stratigraphical absence Prediscosphaera spp. (F) M. delrioensis (F); underlying and overlying. cannot be confirmed. Macroglobigerinelloides spp. (C) Calcisphaerulids (AA) M Marginotruncana sigali (R); M. schneegansi (R); M. renzi (F); M. marianosi (C) Dicarinella primitiva/ M. pseudolinneiana (RR); UC7-UC9 Zone. Marginotruncana sigali Zone: D Watznaueria spp. (C); M. coronata (RR); Concomitant presence of Q. occurrence of Chiastozygus spp. (RR); Retecapsa spp. M. undulata (R); Monte Loffa gartneri and absence of Micula marginotruncanids and (R); Dicarinella imbricata (C); TE Cephalic portion of partially (S. Anna Lastame (Scaglia staurophora. dicarinellids in absence of MGC-IGVR 81375 Quadrum gartneri (RR); D. canaliculata (R); articulated skeleton d'Alfaedo, Rossa Fm.) The scarcity in calcareous Helvetoglobotruncana helvetica Prediscosphaera spp. (F) Heterohelix reussi (R); Verona) nannfossil content makes it hard and Dicarinella concavata, Eiffellithus spp. (RR); H. moremani (RR); to constrain even more the markers of the total range zones Muricohedbergella planispira (RR); biostratigraphic interval underlying and overlying. M. delrioensis (C) EP M. delrioensis (C); Macroglobigerinelloides spp. (RR) Calcisphaerulids (C) A= abundant; C= common; F= few; R= rare; RR= very rare C AC EP TE D M AN U CC EP TE D M AN US C AC C EP TE D M AN US C RI P EP TE D M AN U AC C EP TE D M AN US C RI PT CE PT ED M AN US C R AC C EP TE D M AN US C RI PT PT ED M AN TE D M A ACCEPTED MANUSCRIPT PT C RI US AN M D TE EP C AC AC C EP TE D M AN US C RI P PT ED M AN AC C EP TE D M AN US C RI PT AC C EP TE D M AN US C RI P PT ED M AN CE PT ED M AN US C R AC C EP TE D M AN US C RI PT CE PT ED M AN US C R ACCEPTED MANUSCRIPT Large deadfalls of the ʻginsuʼ shark Cretoxyrhina mantelli (Agassiz, 1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy • Description of partial skeletal remains of the ʻginsuʼ shark Cretoxyrhina mantelli • Nomenclatural remarks about the species C. mantelli PT • Emendation of the diagnosis of the species C. mantelli • Paleobiological and taphonomic remarks about the deadfalls of C. mantelli • Micropaleontological analyses on the C. mantelli specimen matrix RI U SC AN M D TE C EP AC