Palaeogeography, Palaeoclimatology, Palaeoecology 469 (2017) 104–121 Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Direct evidence of trophic interaction between a large lamniform shark, Cretodus sp., and a marine turtle from the Cretaceous of northeastern Italy Jacopo Amalfitano a,⁎, Fabio Marco Dalla Vecchia b, Luca Giusberti a, Eliana Fornaciari a, Valeria Luciani c, Guido Roghi d a Department of Geosciences, University of Padova, Via Gradenigo 6, I-35131 Padova, Italy b Soprintendenza per i Beni Archeologici del Friuli Venezia Giulia, Nucleo Operativo di Udine, Via Zanon 22, I-33100 Udine, Italy c Department of Physics and Earth Sciences, Ferrara University, Polo Scientifico Tecnologico, Via G. Saragat 1, I-44100 Ferrara, Italy d Institute of Geosciences and Earth Resources, CNR, Via Gradenigo 6, I-35131 Padova, Italy article info abstract Article history: In the nineties of the 20th century, a large and partially articulated skeleton of a lamniform shark was discovered in Received 27 July 2016 Upper Cretaceous hemipelagic beds of the Venetian Prealps of northeastern Italy. The shark, dating back to the mid- Received in revised form 21 December 2016 dle Turonian, is here ascribed to Cretodus and represents the first record of this genus in Italy. The fossil is the most Accepted 29 December 2016 complete specimen of Cretodus so far discovered and includes 120 teeth, 86 vertebral centra and many placoid Available online 02 January 2017 scales. Closely associated with the remains of the shark (estimated total length over 6.5 m) is a pellet-like accumu- lation of partially broken bones belonging to a large chelonioid turtle (about 2 m of estimated total length). Some of Keywords: Lamniform shark the bones show evidence of damages referable to bites and possible acid etching. Because of this, and because of Swallowed marine turtle their position in correspondence of the abdominal region of the shark, the turtle remains are interpreted as stomach Predation/Scavenging content. The taphonomy of this association is discussed and compared with other fossil records of shark predation/ Trophic interaction scavenging and with lamniform shark-chelonioid turtle interactions in modern marine environment. The Italian fos- Taphonomy sil represents the second evidence of a turtle swallowed by a shark in the fossil record and a direct evidence of the Cretaceous possible dietary preference of Cretodus, adding some evidence for discerning scavenging from predatory lifestyle. © 2017 Elsevier B.V. All rights reserved. 1. Introduction However, some almost complete vertebral columns of lamniform sharks, often associated with teeth, were discovered between the end The Upper Cretaceous-lower Eocene Scaglia Rossa Formation widely of 19th century and the beginning of 21st century during quarrying ac- crops out in the Venetian Prealps of northeastern Italy. It mainly consists tivities in peculiar lithofacies of Scaglia Rossa (called ʻlastameʼ and of pink to reddish limestones, marly limestones and marls rich in plank- ʻpietra di Castellavazzoʼ). These important finds have been mostly re- tonic foraminifera and deposited in hemipelagic setting (Channell and ferred to Cretoxyrhina mantelli, but after the pioneering paper by Medizza, 1981; Massari et al., 1983). This formation is generally poor Bassani (1888), they were only briefly mentioned in short notes (e.g., in macrofossils, but it is renowned since the 19th century for findings Cigala Fulgosi et al., 1980; Roghi, 2010; Trevisani, 2011), remaining of echinoids, mollusks and rarer marine vertebrates (e.g., Astolfi and practically unknown to the scientific community. Colombara, 2003; Cigala Fulgosi et al., 1980; Dalla Vecchia et al., 2005; Here we report a previously undescribed large lamniform shark as- Giusberti et al., 2005 and references therein; Palci et al., 2013; sociated with remains of a large turtle coming from the Upper Creta- Trevisani and Cestari, 2007). Fossil vertebrates come almost exclusively ceous of Lessini Mountains, near the village of Sant'Anna d'Alfaedo from the Cretaceous portion of Scaglia Rossa and are mainly represented (Verona Province, northeastern Italy; Fig. 1). The goal of this paper is by isolated teeth and vertebrae of chondrichthyans (e.g., Bassani, 1877; to describe in detail and identify this exceptional fossil discussing the Bassani, 1885; Cigala Fulgosi et al., 1980; D'Erasmo, 1922; De Zigno, taphonomic and paleoecological implications of the shark-turtle associ- 1883; Lioy, 1865; Nicolis, 1882; Pellegrini, 1883; Sirna et al., 1994). ation. This study, including precise dating of the fossil through calcare- ous plankton analysis, was carried out in the context of a broader ⁎ Corresponding author. project aimed to describe and revise the entire vertebrate marine E-mail address: jacopo.amalfi[email protected] (J. Amalfitano). fauna from the Scaglia Rossa Formation of northern Italy. http://dx.doi.org/10.1016/j.palaeo.2016.12.044 0031-0182/© 2017 Elsevier B.V. All rights reserved. J. Amalfitano et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 469 (2017) 104–121 105 Fig. 1. Location of the site. The quarry that yielded the specimen IGVR 91032 in the surrounding of Sant'Anna D'Alfaedo (Verona) is indicated by an asterisk. 2. Geological and paleontological context compaction, as suggested by the presence of solution seams (Massari and Savazzi, 1981). The ʻlastameʼ facies is highly reminiscent of the nod- The specimen comes from the ʻGiovanni and Gianfranco Benedetti's ular Rosso Ammonitico Formation of Middle-Late Jurassic age (Massari Quarryʼ that is located on the western slope of Mt. Loffa, near the village and Savazzi, 1981). It deposited on the ʻTrento Plateauʼ, which was a ba- of Sant'Anna d'Alfaedo (Fig. 1), which is approximately 30 km N of Ve- sinal high (Winterer and Bosellini, 1981) some hundred kilometers rona and 50 km W of the famous Eocene Lagerstätten of Bolca away from an emergent area drained by rivers that was located to the (Papazzoni et al., 2014). The hemipelagic limestone of the Scaglia north/north-west, probably in the Upper Austroalpine domain (Cigala Rossa is extensively quarried around Sant'Anna d'Alfaedo and in other Fulgosi et al., 1980; Massari and Medizza, 1973; Massari and Savazzi, localities of western Lessini Mountains for building stone (e.g., Cigala 1981). Fulgosi et al., 1980; Ginevra et al., 2000; Massari and Savazzi, 1981). In strict analogy with ʻRosso Ammoniticoʼ, ʻlastameʼ shows clear evi- The quarried interval, which is locally known as ʻlastameʼ, is a peculiar dence of low sedimentation rates, such as reduced thickness compared lithofacies of Scaglia Rossa (ʽlithozoneʼ 2 of Scaglia Rossa; Lozar and to its temporal range, local invertebrate macrofossil concentrations and Grosso, 1997) and ranges from lower Turonian to lower Santonian local hardgrounds or incipient hardgrounds (Massari and Savazzi, 1981; (Lozar and Grosso, 1997; Palci et al., 2013). ʻLastameʼ is about 7–8m Trevisani and Cestari, 2007). Paleodepth interpretation of Lessinian thick and comprises 70 layers of flaser-nodular whitish to reddish lime- ʻlastameʼ is quite controversial and debated (ranging from 50 to 100 m stones that are laterally continuous with a thickness ranging from 4 to to full bathyal; Trevisani and Cestari, 2007; Venturini, 2012) and is in 35 cm (Lozar and Grosso, 1997; Massari and Savazzi, 1981; Roghi and need of a careful revision based on integrated sedimentological and Romano, 2009). According to the finder, Gianfranco Benedetti, the slab macro-micropaleontological studies. that contains the specimen forming the focus of this study comes from Fossil macroinvertebrates from ʻlastameʼ consist of ammonites, the middle of the section. The strata of ʻlastameʼ are subhorizontal and inoceramids, rudists and irregular echinoids. Skeletons of large marine planar and are separated by thin, dark red and shaly layers (marls and vertebrates were occasionally recovered from different levels within clays). The bedding surfaces are firmgrounds, representing long inter- the quarried interval (Cigala Fulgosi et al., 1980). They include several vals of non-deposition (Trevisani and Cestari, 2007). Large echinoid condrichthyans (Cretoxyrhina mantelli, Ptychodus spp. and a burrows occur on some bedding surfaces (Trevisani and Cestari, sclerorhynchid, Amalfitano et al., 2017; Cigala Fulgosi et al., 1980; 2007). Early lithification of the nodules of ʻlastameʼ has been inferred Trevisani and Cestari, 2007); marine turtles (Protosphargis veronensis from their lack of compactional deformation, while the matrix suffered and other taxa, Chesi, 2008; Villa et al., 2014) and mosasaurs 106 J. Amalfitano et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 469 (2017) 104–121 (Romeosaurus fumanensis and R. sorbinii; Palci et al., 2013). Despite the 3.1. Materials importance of these finds, recent paleontological studies have been oc- casional, including just the revisions of the rudists (Trevisani and The specimen IGVR 91032 (Fig. 2) is housed in the Museum of Cestari, 2007) and mosasaurs (Palci et al., 2013). Most of the vertebrate Prehistory and Paleontology of Sant'Anna d'Alfaedo. It consists of the remains recovered from the quarries of ʻlastameʼ near Mt. Loffa are main slab (slab A; Fig. 2), which preserves the teeth, the proximal housed in the Museum of Prehistory and Paleontology of Sant'Anna segment of the vertebral column and the partial cranial calcified carti- d'Alfaedo