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Paleodays 2019 La Società Paleontologica Italiana a e

Parte 2: Guida all’escursione

XIX Riunione annuale SPI Ente GeoPaleontologico di Pietraroja (21)22-24(25) Maggio 2019

a cura di Rook L. & Pandolfi L. Paleodays 2019. La Società Paleontologica Italiana a Benevento e Pietraroja

XIX Riunione annuale della Società Paleontologica Italiana Benevento/Pietraroja, (21)22-24(25) Maggio 2019

Comitato Organizzatore Ente GeoPaleontologico di Pietraroja: G. Santamaria, G. Festinese, P. Forte, G. Lioni, A.V. Maturo, R. Melillo, L. Prencipe, A. Torrillo, F.O. Amore, S. Foresta, C. Dal Sasso, V. Morra, L. Rook

Comitato Scientifico F.O. Amore, L. Angiolini, A. Bartiromo, M. Bernardi, G. Carnevale, M. Cherin, M. Chiari, G. Crippa, C. Dal Sasso, A. Ferretti, E. Ghezzo, L. Jaselli, L. Pandolfi, P. Raia, L. Rook

Con il supporto di Ente Geopaleontologico di Pietraroja Univeristà degli Studi del Sannio Dipartimento di Scienze e Tecnologie, Univeristà degli Studi del Sannio Dipartimento di Scienze della Terra, Università degli Studi di Firenze Confindustria, BN Provincia di Benevento di Pietraroja, BN

Con il patrocinio di Ministero dell’Ambiente e della Tutela del Territorio e del Mare MATTM Ministero per i Beni e le Attività Culturali MiBAC Istituto Superiore per la Protezione e la Ricerca Ambientale ISPRA Federculture Regione Provincia di Benevento Comune di Pietraroja Parco regionale del Museo Civico di Storia Naturale di Milano Società Geologica Italiana Università degli Studi di Firenze Università degli Studi del Sannio Università degli Studi di Napoli “Federico II” Università degli Studi di Napoli “Suor Orsola Benincasa” Università degli Studi di Napoli “L’Orientale” Università degli Studi della Campania Luigi Vanvittelli

Progetto grafico logo di copertina M. Repola, R. D’Uva

Rook L. & Pandolfi L. (a cura di) 2019. Paleodays 2019. La Società Paleontologica Italiana a Benevento e Pietraroja. Parte 2: Guida all’e- scursione della XIX Riunione annuale SPI (Società Paleontologica Italiana). 24 pp. Ente GeoPaleontologico di Pietraroja (Benevento). ISBN 979-12-200-4867-5 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Scipionyx samniticus & Pietraroja Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Restoration of , according to the Milanese palaeoartist Davide Bonadonna. (© Davide Bonadonna)

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Scipionyx samniticus

Cristiano Dal Sasso & Simone Maganuco

Scipionyx samniticus abruptly entered the limelight when, on the occasion of its formal naming, it made the cover of Nature (Dal Sasso & Signore, 1998), attracting not only the interest of palaeontologists but also popular imagination throughout the world. In fact, the Scipionyx is a striking, articulated, juvenile coelurosaur with a unique combination of osteological characters and superbly fossilised soft tissues (Fig. 1). The latter render Scipionyx one of the best-preserved known, and a unique specimen within the fossil record of Mesozoic vertebrates. Scipionyx samniticus was the first fossil body unearthed in ; thus, its discovery was a major event in the history of Italian palaeontology.

The finding The story of the discovery of the Scipionyx fossil is original in itself (for a detailed account, see Dal Sasso, 2001, 2004). In the spring of 1981, Giovanni Todesco unearthed it at Le Cavere. The collector cleaned the fossil as well as he could and stored it in the basement of his house. It remained the- re until 1993, when Todesco showed it to professional pa- laeontologists, who identified the tiny reptile as a dinosaur. In accordance with Italian law, the specimen was handed over to the Superintendence of the site of provenance, and the popular magazine Oggi dubbed the dinosaur “Ciro” (a typical Neapolitan name). In 1994, the Museo di Storia Na- turale di Milano (MSNM) obtained permission to properly prepare and study the fossil. It was only during this prepa- ration work, that the extraordinary degree of preservation of the specimen’s soft tissue was fully realised. This later became the main focus of the paper published in Nature on its formal description (Dal Sasso & Signore, 1998), and, subsequently, of a detailed monograph (Dal Sasso & Maga- Fig. 1 - Overall view of the holotype (and only known specimen) of nuco, 2011), which is most complete publication existing Scipionyx samniticus. Most of the soft tissues are visible to the naked eye to date on Scipionyx samniticus. on account of their distinctive ochre colour. Other organic remains are preserved as thin films, that can be seen only under ultraviolet-induced Osteology, ontogenetic stage and phylogenetic rela- fluorescence. Scale bar = 2 cm. (After Dal Sasso & Maganuco, 2011. © tionships SABAP-CE-BN, centro operativo di Benevento, courtesy MIBAC. Photo The diagnosis of this taxon includes five premaxillary te- Roberto Appiani & Leonardo Vitola) eth, a sinusoidal ridge of the supratemporal fossa, a distally squared descending process of the squamosal, lower tooth row extending farther back than the upper row, and the absence of an external mandibular fenestra (Fig. 2). Among the relevant postcranial skeletal features are fan-shaped dorsal neural spines with beak-like attachments, hair-like cervical ribs, dorsal ribs with cup-like sternal at- tachments, carpus composed of only two stacked, well-os- sified bones, manual digit III longer than digit I, a cranially notched iliac preacetabular blade and a distally squared ischial obturator process. The holotype (and only know specimen) of Scipionyx sam- niticus is clearly a very immature individual, probably less than three weeks old at the time of death. This is indicated by a long list of juvenile characters, such as the presen- ce of a frontoparietal fontanelle (Fig. 2), a short and deep antorbital region, tooth replacement not yet started, pe- culiar scarred bone surfaces, non-sutured girdle elements Fig. 2 - and of Scipionyx samniticus. The large circular and closure of the neurocentral sutures not yet started in orbits, the short snout, and the fronto-parietal fontanelle (i.e., the U-shaped gap on the cranial vault) point out to the very young age of any vertebra, and -last but not least- anterior abdominal the . (After Dal Sasso & Maganuco, 2011. © SABAP-CE-BN, centro displacement of the intestine, with an empty space in the operativo di Benevento, courtesy MIBAC. Photo Roberto Appiani) pelvic area suggesting the presence of a yolksac. 5 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Fig. 3 - Map of the soft tissues preserved in the holotype of Scipionyx samniticus, obtained by combining observations under optical microscopy, ultraviolet light and scanning electron microscopy. (After Dal Sasso & Maganuco, 2011. © Museo di Storia Naturale di Milano. Drawing Marco Auditore) According to Dal Sasso & Maganuco (2011), the recon- structed total body length of the holotype of Scipionyx does not exceed 50 cm. After having re-articulated its elements and corrected for the deformation, the pel- vis of Scipionyx results comparatively slightly narrower mediolaterally than that of , the three-dimen- sionally preserved pelvis of which has a width appro- ximately 30 mm wide across the (Martill et al., 2000). Comparison with usual body proportions of small extinct and extant coelurosaurs, including , permitted to tentatively estimate a weight in life of no more than 0.2 kg. Phylogenetic analysis of (90 taxa, 360 characters), evaluating also the ontogeny-related cha- racters, identified Scipionyx as a basal member of a monophyletic , which resulted to be more derived than (Dal Sasso & Ma- ganuco, 2011).

Soft tissue anatomy and taphonomy Fig. 4 - The duodenal loop of the intestine of Scipionyx samniticus still pre- Scipionyx superbly preserves a unique variety of fossi- serves the circular folds of the mucosa (arrows). The iron-rich reddish halo to the right comes from the decay of the (After Dal Sasso & Maganuco, lised internal organs and soft tissues (Fig. 3), so that a 2011. © SABAP-CE-BN, centro operativo di Benevento, courtesy MIBAC. real paleo-autopsy has been possible (for a complete Photo Leonardo Vitola) account, see Dal Sasso & Maganuco, 2011). External

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soft tissues are beautifully represented by the horny (keratinised) manual claws. The internal tissues in- clude axial , axial and appendicular arti- cular cartilage, neck muscles and connective tissue, part of the , oesophageal remains, traces of the liver and other blood-rich organs, the entire intestine, mesenteric blood vessels, and pelvic and hind limb muscles. The intestine is the largest, most complete and most visible internal organ of Scipio- nyx (Fig. 4): its three-dimensional duodenal loops are lumpy and shiny, reminiscent of the aspect one would see after dissecting a modern animal; the shows a typical lower density of mucosal folds and garland-like coils in the dorsal portion of the abdomen; cranial to the pelvic girdle, a constric- tion possibly marks a ileorectal valve; the makes the terminal enlarged tract of the gut that, after passing through the pubic and the ischial fora- Fig. 5 - SEM imaging of a phosphatised capillary-sized blood vessel in a mina, reaches the base of the tail. microsample of the rectum of Scipionyx. The vacuolar aspect of the matrix is Most of the soft tissues can be easily identified to consistent with the tissue having been replaced by pseudomorph phosphati- the naked eye by their ochre colour, whereas other sed bacteria, many of which are preserved as hollow spheres. (After Dal Sasso organic remains are preserved as thin films that are & Maganuco, 2011. © Museo di Storia Naturale di Milano, courtesy MIBAC. visible only under ultraviolet-induced fluorescence Photo Michele Zilioli) (UV). Scanning electron microscopy (SEM) analyses have revealed an exceptional three-dimensional preservation of the soft tissues and astonishing information at a cellular and even subcellular level (Fig. 5-6), such as the sarcomere-related banded pattern observable within every single myofibre of the M. caudofemoralis longus. SEM element microanalysis has also confirmed the haematic origin of the reddish macula referred to the liver and other blood-rich organs (Fig. 4). The exceptional preser-vation of labile soft tissue indicates that, after death, the carcass of this theropod hatchling was subjected to very little decay and rapid authigenic mineralisation (Briggs, 2003) in the presence of a high concentration of phosphates. In facts, the diagenetic processes were halted very early after Scipionyx died, so that mineralisation outpaced degradation. Authi- genic mineralization occurs through early infiltration and permeation of soft tissue by mineral-charged water and differs from petri- fication, which is a replacement process. The contemporary presence of substrate-, inter- mediate- and microbial-microfabrics (sensu Wilby & Briggs, 1997) in different anatomical portions of Scipionyx likely reflects the diffe- rent rates at which microbes gained access to the subcutaneous tissues of the organism.

Palaeobiology and physiology Outstandingly, the degree of preservation of the soft tissues has permitted an analysis of the relative position of the food remains in the digestive apparatus and, thus, recon- struction of a feeding chronology for this specimen, an insight that is usually impos- sible to obtain for fossil vertebrates. In fact, Scipionyx’s gut has been found to contain allogenous bones from a lepidosaurian rep- tile in the stomach region, -like polygo- nal squamae in the , fish scales in the rectum, and a variety of tiny remains in Fig. 6 - Above: the cells of the caudofemoral muscle of Scipionyx, still forming a com- several points of the intestine (Fig. 13). This pact bundle. Below: close-ups of the same muscle as seen under scanning electron is compelling evidence that Scipionyx fed on microscope, revealing exceptional fossilisation of the banded pattern (sarcomera) both and fish. The relatively large size within every single cell. In taphonomical terms, such a level of preservation is related of a leg of lizard found in the stomach sug- to the formation of a substrate microfabric (sensu Wilby & Briggs, 1997). (After Dal Sasso & Maganuco, 2011. © Museo di Storia Naturale di Milano, courtesy MIBAC. gests that the little dinosaur has been fed by Photo Michele Zilioli) 7 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Fig. 7 - Reconstruction of the skeleton of Scipionyx (missing bones in light gray) and of its prey, drawn to scale. The circled numbers indicate the sequence of intake, based on the gut contents of the baby dinosaur (After Dal Sasso & Maganuco, 2011. © Museo di Storia Naturale di Milano. Drawing Marco Auditore) their parents with pieces of prey captured and dismembered specifically to feed the nestlings. The digestive physiology of Scipionyx seems different from that of extant . fully dissolve small- to medium- sized bones and regurgitate large undigested bones; similarly, carnivorous birds regurgitate bones and other hard prey items within their pellets, producing faeces that are devoid of solid inclusions, as well. Possibly, theropod dinosaurs also regurgitated the largest chunks of bone, but Scipionyx tells us that the swallowed bones were mostly guided through the intestine, like in carnivorous mammals and lepidosaurian reptiles, that have pyloric openings larger than in crocodiles and produce faeces containing bones and other incompletely digested hard tissues. This gives important confirmation to the supposedly theropod origin of some bone-bearing described in recent (Chin et al., 2003; Chin & Bishop, 2007). In turn, the remains or imprints purported by some authors to be of the diaphragmatic muscles (Ruben et al., 1999) are, in fact, a calcite nodule of amorphous microstructure, inconsistent with the preservation of other muscle tissue in this specimen (Dal Sasso & Maganuco, 2011). This evidence, and other anatomical observations on bones and soft tissues, deny the hypothesis of a crocodilian-like hepatic-piston assisted breathing mechanism in Scipionyx. The amount and detail of information gained from this single specimen make the Pietraroja Plattenkalk a unique fossil locality. In contrast to the Chinese Jehol Group, which is a lacustrine/volcanic freshwater deposit that has preserved dinosaurs with delicate integumentary structures, such as filaments, feathers and bristles, the Italian shallow marine Lagerstätte has preserved internal organs. This is unprecedented not only for a dinosaur, but also for any other Mesozoic terrestrial vertebrate.

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The Pietraroja region, a geopaleontological overview

Filomena Ornella Amore, Antonello Bartiromo, Cristiano Dal Sasso

Geological setting

Filomena Ornella Amore

The Matese massif represents part of the Central-Southern Apennine, a segment of the thrust belt-foredeep-foreland system that originated during the Cenozoic (Fig.1). The area experienced compressive and extensional tectonic events, mainly related to the deformation and subduction of the continental margin of the Adria Plate and to the opening of the Tyrrhenian Sea, which designed the present-day structural features of the Southern Apennines (Patacca & Scandone, 1989; Doglioni, 1991; Patacca & Scandone, 2007; Bonardi et al., 2009). The Matese Mountains are formed by the pre-orogenic Mesozoic to Tertiary successions, constituted by shallow water to slope carbonates, 3-4 km thick, topped with middle-upper Tortonian turbiditic and silicoclastic sediments of the foredeep basin (D’Argenio et al., 1973; Ciampo et al., 1983; Lirer et al., 2005). Different palaeogeographic reconstructions are available for this part of the Apennines (D’Argenio et al., 1973; Mostardini & Mer- lini, 1986; Bonardi et al., 1988; Sgrosso, 1988a,b; Pescatore, 1988; Patacca et al., 1992; Pescatore et al., 1999, Patacca & Scandone, 2007; Vitale & Ciarcia, 2018), however there is a general agreement in regarding the present-day Pietraroja area as part of a shal- low-water carbonate domain which, during the Early , experienced tropical-subtropical climatic conditions. Begin- ning with the Late Aptian-Early , margin retrogradation and broad tectonic uplifts took place, with the establishment of more open marine conditions, including deeper margins connecting the shelf to the basin areas (Vigorito et al., 2003; Carannan- te et al., 2004). The deposition of the fossiliferous upper Plattenkalk deposits of Pietraroja preceded the mid-Cretaceous» strati- graphic gaps and the beginning of karst processes, associated to the formation of bauxite bedrocks (see the Stop of Regia Piana). The bauxite bodies are unconformably covered by late Turonian, shallow water rudist-rich sediments (Carannante et al., 1988; Carannante10 S. VITALE et AND al., S. CIARCIA 2009). A paraconformity surface separates the Cretaceous carbonate platform succession by overlaid Middle Miocene calcarenites (Cusano Fm.); these suggest a neritic marine envi- ronment evolving, upwards, in the deep and open sea environments of marly limestone and clayey marl, rich in planktonic foraminifers of Longano Fm, Serravallian-Torto- nian in age. The Middle-Upper Tortonian sediments of Pietraroia Fm. indicate the onset of the silici- clastic sedimentation in the area. Between the upper Tortonian and the lower Messinian, the area was involved in the orogenic transport, the front and the deformation mi- grated eastward, thrusting the Ma- tese toward the Apulian domains. From the Late Pliocene to the Early Pleistocene the area was affected by intense fragmentation due to normal faults and by the subse- quent uplift. Since the Middle Plei- stocene an extensional tectonic regime occurred and brought the Matese Massif to its present ele- vation and structural setting of a horst bounded by the tectonic de- pressions of the Boiano basin and Valley (Carannante et al., 2006; Amato et al., 2011; Aucelli Fig. 1 - Schematic geological map of the southern Apennines (after Vitale & Ciarcia, 2018) 2013). Figure 1. Schematic geological map of the southern Apennines (after Vitale, Amore, et al., 2017). 9 successions, dissected by several Quaternary structural Europa/AlKaPeCa plates (e.g. Carminati, Lustrino, & depressions, especially along the Tyrrhenian Sea side. Doglioni, 2012; Cosentino, Cipollari, Marsili, & The allochthonous units, forming the orogenic struc- Scrocca, 2010; Vitale & Ciarcia, 2013 and reference ture, can be grouped in three main tectonic complexes: therein) with the E-migration of the thrust front, as (i) Ligurian Accretionary Complex (LAC), (ii) Apen- consequence of the downgoing slab-retreat (Malin- nine Platform (AP) units and (iii) Lagonegro- verno & Ryan, 1986). The subduction started in the Basin (LMB) units. In the study area, the tectonic pile /Eocene time (Rossetti et al., 2001) with is unconformably covered by Mio-Pliocene wedge- different paroxysmal tectonic stages such as the Oligo- top basin deposits and Quaternary post-orogenic sedi- cene–Langhian and upper Serravallian–Tortonian fast ments and volcanics. LAC occupies the highest tectonic thrust front migrations that caused the opening of positions, covering the AP units, in turn overthrusting Ligurian–Provençal and Tyrrhenian back-arc basins, the LMB units. The foreland is represented by the Apu- respectively (Dewey, Helman, Turco, Hutton, & lian Platform not cropping out in the Campania region. Knott, 1989; Faccenna, Becker, Lucente, Jolivet, & Ros- As it is clear from many wells and seismic surveys car- setti, 2001; Milia & Torrente, 2014; Milia, Valente, et ried out for the oil exploration (ViDEPI project) or for al., 2017). In the Aquitanian–Burdigalian interval, the scientific purposes (CROP04 project), the LMB units orogenic pulses formed an incipient orogenic prism form tectonic duplexes and imbricated slices over- made up of oceanic to transitional successions (LAC; thrusting the buried Apulian carbonates. Ciarcia et al., 2012; Vitale & Ciarcia, 2013). Sub- Southern Apennines result from the subduction of sequently the thrust front eastward migrated, involving the Neo-Tethys oceanic lithosphere beneath the younger and younger foredeep basin deposits covering Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

STOP 1 - Pietraroja overview

In this first stop, in the cemetery area, we will have the opportunity to have an overview of the area while enjoying a panoramic view of the Meso-Cenozoic sediments that will be the subject of the following stops. Inside the cemetery we will observe M.te Cigno, the Valley, the sediments of Pietraroia Formation, and the ruins of the ancient village of Pietraroja ; outside the cemetery, we will enjoy a spectacular view of the Fossil-Lagerstätte of “Le Cavere”.

STOP 2 - The Fossil-Lagerstätte of “Le Cavere” in the village of Pietraroja

Cristiano Dal Sasso & Antonello Bartiromo

Location and historical background Pietraroja sits near the top of a 970 m high carbonate relief which rises almost vertically from the plain adjacent to the eastern margin of the Matese Mountains (central southern Italian Apennines), some 70 km northeast of (Figs. 1-2). The Lower Cretaceous levels, assigned to the Lower Albian on the basis of foraminiferal biozonology (Bravi & Garassino, 1998; Carannante et al., 2006), crop out at the “Le Cavere” locality, just above the village. The limestones of Pietraroja have been known for their beautifully preserved since the nineteenth century. The palaeon- tological richness of the outcrop was mentioned for the first time by Breislak (1798), however, Niccolò Braucci da Caivano in his manuscript, studied by D’Erasmo (1941), probably referred to Pietraroja locality mentioning fossil fish from Matese Mountains. Some limestone beds from this site were also used for lithography (Bassani, 1892). . Costa (1851, 1853-1864, 1865, 1866) descri- bed the “Calcari ad Ittioliti di Pietraroja” in a series of superbly illustrated volumes (Fig. 3). The term was slightly modified into “calcari selciferi ed ittiolitiferi di Pietraroja” (Catenacci & Manfredini, 1963), which has been used frequently to indicate that geo- logical formation. A formal name has not been validated, yet (Petti, pers. comm., 2010). Here we use the term Plattenkalk sensu Carannante et al. (2006). The age of these limestones, considered to be , was a subject of debate up to the end of nineteenth century, when Bassani (1885) re-examined the fossil fishes collected by Costa and confirmed their Cretaceous affinity. The deposit was then studied by D’Erasmo (1914, 1915) and D’Argenio (1963); in more recent times, research has been carried out by the Università di Napoli “Federico II”, which in 1982, in cooperation with the Museo di Scienze Naturali di Torino, conducted excavations (Bravi, 1987, 1988, 1994). After the Scipionyx fossil came to light (Dal Sasso & Signore, 1998), a small quarry was opened in 2001 by the MSNM (Dal Sasso et al., 2014). Today, the fossiliferous outcrop is protected by a fence.

Lithology, stratigraphy and sedimentology The Cretaceous succession of Pietraroja, which is about 340 m thick, includes two well-stratified Plat- tenkalk horizons (Fig. 4). According to Carannante et al. (2006), each Plattenkalk presents distinct fining and thinning upward trends with a detritic basal la- yer. The lower Plattenkalk is averagely coarser and is characterised by the occurrence of frequent bentho- nic foraminifera-rich grainstones and packstones: it is relatively poor in macrofossils and, to date, no vertebrate remains have been reported from this interval. Above this is a second Plattenkalk horizon, with a depth of 8-9 m. The thickness of the upper Plattenkalk increases to the southwest, reaching a maximum of some 15 m at the Le Cavere outcrop, which is the source of the major fossil finds. The up- per Plattenkalk is averagely finer when compared with the lower one, as it is mostly composed of fine laminated mudstones. In the middle and upper part of this interval there is a prevalence of mudstones Fig. 2 - The north-western slope of the “Civita di Pietraroja” and the outcrop with thin strata and lenses of packstones and wa- of “Le Cavere” (centre right). Part of the fossiliferous site was fenced off after ckestones rich in spicules of siliceous sponges. The the discovery of Scipionyx (after Dal Sasso & Maganuco, 2011)

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Fig. 3 - Lithography illustrating crassirostris and other fossil fishes unearthed from the Pietraroja Plattenkalk in the nine- teenth century (after Costa, 1853-1864) trend of the layers is about N-S leading eastward, with an average dip around 20°. Carannante et al. (2006) dated the Pietraroja Plattenkalk as Lower Albian on the basis of its microfossil assemblage, which in- cludes Bacinella irregularis, Glomospira sp., Cuneolina aff.pavonia, Praechrysalidina infracretacea, Nummuloculina sp., Thaumato- porella sp., Debarina sp., Ovalveolina reichelii, Sabaudia minuta, orbitolinids, miliolids and textularids. This fits well with previous biostratigraphic data provided by Bravi & De Castro (1995) and Bravi & Garassino (1998). Stratigraphic position of Scipionyx - according to the person who found the fossil (Todesco, pers. comm., 1993), Scipionyx was collected at a point of the Le Cavere quarry that, based on the present-day topography, falls into the lower-right quarter of the fenced-off area of the outcrop. More precisely, the discovery site is located to the right of the entrance gate and some 7-9 m from the paved road (once a gravel one). Todesco also has stated (pers. comm., 2010) that the layer from which Scipionyx came from outcropped some 50-80 cm higher than the present stepping (bottom) layer, but in any case at an elevation below the level of the road. The position of this layer, as well as the contents of the slab, are consistent with the lower-middle series of the upper Plattenkalk. According to Dal Sasso & Maganuco (2011), both the texture and chemical composition of the sediment em- bedding Scipionyx fit well with the description of the “spicule-rich packstone/wackestone microfacies” (Carannante et al., 2006). This microfacies “largely dominate[s] the lower-middle portions of the upper Plattenkalk”, providing good sedimentological evidence that Scipionyx comes from one of those layers.

Depositional environment Fossil-rich Plattenkalks have been documented from shallow lagoonal to relatively deep basinal settings, so there is still great uncertainty on the depositional environment that created the Pietraroja Plattenkalk. On this point, three distinct models have been proposed.

Lagoon model - According to some authors (D’Argenio, 1963; Bravi & Garassino, 1998), the Pietraroja Plattenkalk was laid-down close to a coastal area, in a very shallow lagoonal environment that was frequently isolated from the open sea but subject to tidal influence and occasional storms. Apparent cyclicity in the lithotype distribution, inferred from the graded horizons in most layers of the upper Plattenkalk, led to hypothesise that sediment deposition was linked to the rhythm of tides and storms which reached the lagoons and poured the finer sediments of the surrounding above-sea-level areas into them.

Slope/shallow basin model - Catenacci & Manfredini (1963) believe that deposition of the Pietraroja Plattenkalk took place in a coastal strip which ran between the margin of the platform and the deepest sediments of the opposite basin (Molise-Sannitica depression). The transitional nature of this environment is inferred by sedimentological and stratigraphic observations, inclu- ding a lateral “flute-beak” transition from reefoidal to plattenkalk deposits. Freels (1975) estimated the Pietraroja basin to have been about 60 m deep and 1 km wide.

Submarine channel model - A third model was proposed more recently by Carannante et al. (2006). According to it, the Pie- 11 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Fig. 4 - A) lithostratigraphic column of the Cretaceous succession of Pietraroja, according to Carannante et al. (2006); B) stratigraphic sequence of the north-eastern side of the “Civita di Pietraroja”, according to Bravi & Garassino (1998). Plattenkalk I and II in B roughly correspond to lower and upper Plattenkalk in A. Abbreviations: U) unconformity; E) erosional surfaces. Microfossils in green, macro- fossils in brown (after Dal Sasso & Maganuco, 2011)

12 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019 traroja Plattenkalk sequences would not be shallow lagoonal deposits or intra-platform basin-fill, but deposits of a submarine channel. In particular, the lower Plattenkalk and the fine-grained, fossil-rich upper Plattenkalk are interpreted as representing, respectively, channel-fill and abandon deposits, which followed previous coarse-grained channel-fill sequences as a response to the demise of the channel as a sedimentary conduit. According to this model, Scipionyx was buried in a single, rapid event by a turbidite. As Bartiromo (2013, p. 75) claimed, the shallow water interpretation can not be excluded: plant taphonomy and the analyses of paleozoological data contradict the submarine channel model. The evidence that phosphorus is limited to the body of Scipio- nyx (i.e., not present in the sediment) has lead Dal Sasso & Maganuco (2011) to prefer the idea of a depositional environment with relatively shallow, open marine waters, rather than a Solnhofen-like lagoon, where phosphates deriving from organic activity would have more easily accumulated.

Palaeogeography Palaeogeographical and palaeobotanical (Bartiromo, 2008, 2013) data related to the Albian age support the hypothesis that Scipionyx and the other terrestrial fauna of Pietraroja inhabited temporarily isolated lands that rose up in the Cretaceous Tethys during the Middle-Upper Aptian tectonic phases. Those lands were part of the Adria Plate and of the Periadriatic Domain (Zap- paterra, 1990), which is considered either an independent microplate or an African Promontory (Cillari et al., 2009). In any case, the Periadriatic Domain was a complex puzzle of small units traditionally described as an archipelago of carbonate platforms that was well-separated from both Gondwana and Laurasia. Within this scenario, the carbonate platform bearing the present-day Pietraroja – the Apenninic (or Laziale-Abruzzese-Campa- na) Platform – is assumed to have been separated in Aptian-Albian times from the adjacent Apulian Platform by the Lagonegro- Molise Basin. The Apenninic Platform, not exceeding 250x200 Km in size (Nicosia et al., 2007), can thus be imagined as a small island no larger than the present Sardinia. The evidence on subaerial emergences, and the present knowledge of dinosaur occurrence (tracksites) in the Italian Periadriatic platforms is surprising and in definite contrast with the idea of a continuous marine environment (Nicosia et al., 2007; Sacchi et al., 2009). The temporary land connections that permitted the dispersal of terrestrial vertebrates within the area are considered by Sacchi et al. (2009) as filtering-bridges.

The fossil assemblage The biodiversity of the site, revealed by the high number of taxa found there (e.g., Fig. 7), indicates that the depositional basin was surrounded by a variety of natural settings. The presence of terrestrial reptiles, along with terrestrial plants and fish genera found also in ancient freshwater settings (e.g., Lepidotes, Pleuropholis), confirms that dry land with sur- face freshwater was not far. Palaeogeographical data related to the Albian age support the hypothesis that Scipionyx and the other terrestrial fauna of Pietraroja inhabited temporarily isolated lands bordered by large, shallow lagoons and biogenic margins and ramps. Below is a concise list of the flora and fauna so far described.

Microfossils - The site contains a significant quantity of radiolarians and spicules from sili- ceal sponges. Benthic macroforaminifers are also found in the lower and upper Pietraroja Plattenkalks. These include orbitolinids, alveolinids, miliolids and agglutinated foraminifers. Some layers are rich in calcareous algae (Dasycladaceae).

Plants - As for plant fossils of “Le Cavere”, Costa (1865) mentioned “Quattro specie innomina- te” (Four unnamed ). Some plant fossils collected from this site have been assigned to Zamites and Brachyphyllum (Bravi & Garassino, 1998) while Bartiromo et al. (2006a) do- cumented the presence of a conifer, Frenelopsis. In a further study, Bartiromo et al. (2006b) discovered the impression of a matoniaceous fern assigned to Phlebopteris. More recently, Bartiromo (2008, 2013) studied plant macrofossils (Fig. 5) with epidermal fe- atures (Fig. 6) from the Plattenkalk 2 as exposed in “Le Cavere” quarry. Plant remains include sterile foliage bearing shoots and reproductive structures of gymnosperms and possible angiosperm leaves occurring in a controversial marine depositional environment (see the considerations above). The following taxa were recorded: Brachyphyllum sp., Frenelopsis sp., Cheirolepidiaceae gen. et sp. indet. and Nageiopsis?. Bituminous strata of the lower Plattenkalk 2 are full of plant debris composed mainly of isolated leaves and sterile axes. In the Pietraroja Fossil-Lagerstätte, the occurrence of large detached branches is likely the Fig. 5 - Le Cavere. Frenelopsis sp. result of physiological, e.g., drought periods (Gomez et al., 2002a) or water stress at the on- Three-dimensional preservation (im- set of unfavourable seasons (Alvin, 1983), or mechanical (e.g., storms) damage. In the Al- pression) of a large branched shoot bian locality of Pietraroja, Frenelopsis is the dominant, best preserved and well articulated showing one-order branch. Speci- plant. Taphonomic evidence from Frenelopsis from the of Italy (Gomez et al., men M20563: Museo di Paleontolo- 2002a), suggest that large branches could not have endured long time under high-energy gia, Università Federico II, Napoli 13 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

transport without the shoots becoming fragmented. Frenelopsis from Pietra- roja were most likely transported a relatively short distance, and certainly grew near the site of deposition. However, the general conditions of preser- vation of plant fossils from Pietraroja could be also due i) to a long-lasting flotation before burial, and/or ii) to a subaerial exposure in the leaf litter (Gomez et al., 2002b) or, in general, iii) may be evidence of prolonged expo- sure to oxic environment before deposition in an anoxic environment. The different states of preservation of the studied plant fossils would seemingly suggest different source areas. Probably onlyFrenelopsis grew close to the hypothesized lagoon, with organic debris supplied to the sedimentary basin from the surrounding emerged land. Therefore, taphonomic considerations suggest parautochtonous deposition for Frenelopsis while the other plant re- mains are allochthonous. Among the plants studied, the cheirolepidiaceous Fig. 6 - Le Cavere. Frenelopsis sp. Inner view of cuti- conifers have ecological significance owing to their (debated) xeromorphic cle showing two rows of stomatal complexes, the foliar features. These adaptations suggest a warm and dry, or possibly coastal hypodermal cells (arrow) undercovering ordinary palaeoenvironment. epidermal cells and dorsal plates (dotted arrow) of Further research on this topic are being carried out and they will expand the guard cells. Specimen M 20913: Museo di Paleonto- phytotaxonomic knowledge of the emergent lands within the Early Creta- logia, UMuseo di Paleontologia, Università Federico II, Napoli ceous Apenninic Carbonate Platform.

Invertebrates - The invertebrate records are abundant and include Nerineidae gastropods, undetermined bivalves, and Hopli- taceae ammonites of the Trochleiceras. Echinoderms are rare (Fig. 7A) and represented only by small-sized Asteroidea and Ophiuroidea. Interestingly, three brand new taxa of decapod crustaceans were described by Bravi & Garassino (1998): the penaeid Micropenaeus tenuirostris, the caridean Parvocaris samnitica (Fig. 7B) and the thalassinid Huxleycaris beneventana. Pos- sible palaemonid and astacid crayfish are also present.

Fishes - Fishes are the most abundant and diversified fossil vertebrates at Pietraroja (for an almost complete list, see Bravi, 1999). Most coprolites, which abound in some layers of the upper Plattenkalk, can also be referred to fish. The Osteichthyes are repre- sented by basal neopterygians (Holostei) and more derived neopterygians (Teleostei). The former include three extinct suprage- neric taxa: the (Fig. 7C), with the genera Paleobalistum (D’Erasmo, 1914, 1915) and Ocloedus (Poyato-Ariza & Wenz, 2002), have a disk-shaped, laterally flattened body, well-adapted for swimming in narrow reefs, and typical dome-shaped crushing teeth, which allowed them to feed on hard-shelled invertebrates; the Macrosemiiformes are represented by Notagogus pentlandi, a species with a peculiar dorsal fin composed from two closely spaced lobes; the Semionotiformes include the genus Lepidotes, a freshwater taxon covered by very robust, diamond-shaped ganoid scales. At least six suprageneric teleostean taxa are represented at Pietraroja: the , such as Belonostomus crassirostris (Costa, 1853-1864), were slender-bo- died pelagic predators equipped with a long rostrum and pointed teeth; the , open-water pre- dators of even larger size, are represen- ted by the genera Chirocentrites and (Signore et al., 2005, 2006). Other predatory teleosteans include the elopomorph Anaethalion and the possibly endemic pholidophoriform Pleuropholis decastroi, which is regar- ded a freshwater-brackish form. Ano- ther species known only in this locality is the ionoscopiform Ionoscopus petra- roiae. The Clupeomorpha are represen- ted by two genera: Diplomystus, a form with a very convex abdomen – related to present-day sardines – and Clupa- vus, which is one of the smallest and most abundant fishes in the Pietraroja outcrop. The Chondrichthyes are repre- sented by a fossil guitarfish,Rhinobatus obtusatus (Costa, 1853-1864): the only specimen so far recovered is remarka- Fig. 7 - A snapshot of the fossil biodiversity at Pietraroja: A) an indeterminate sea star; B) a bly articulated and well-preserved, skin decapod crustacean (Parvocaris samnitica); C) a pycnodontid fish Ocloedus( costai); D) a gui- remains included (Fig. 7D). tarfish Rhinobatus( obtusatus). Not to scale. (After Dal Sasso & Maganuco, 2011) 14 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Tetrapods - are represented by a single specimen, the batrachian megacephalus (McGowan, 2002), belonging to the - tidae – an extinct group distantly related to extant salamanders. Lepidosauromorph reptiles are repre- sented by sphenodontian and squamate lizards; the former include two different taxa, represented by single specimens: Derasmosaurus pietrarojae (Fig. 8) (Barbera & Macuglia, 1988), and a still unnamed animal which in turn preserves within its abdominal cavity the remains of a small lizard, named Eichstaet- tisaurus gouldi (Evans et al., 2004). Two other Squa- mata are known from the Pietraroja outcrop: Cho- metokadmon fitzingeri (Costa, 1853-1864), similar in body shape to modern scincids, is in fact related to the Anguimorpha; and Costasaurus rusconii (Estes, 1983), a small-bodied, short-limbed animal thought originally to be an . A fourth lizard taxon awaits description. Archosauromorph reptiles are re- presented by the theropod dinosaur Scipionyx sam- niticus (Dal Sasso & Signore, 1998) and by at least Fig. 8 - The holotype of Derasmosaurus pietraroiae (MPN 541), the best preser- two semi-complete, articulated Crocodylomorpha ved sphenodontian reptile from Pietraroja. Scale bar = 10 mm. (After Dal Sasso that pertain to a single hylaeochampsid eusuchian & Maganuco, 2011) taxon, recently named Pietraroiasuchus ormezzanoi (Buscalioni et al., 2012).

STOP 3 – Pietraroja PaleoLab

The Paleo-Lab is a Museum-Laboratory inaugurated in 2005 under the aegis of the , the Ministry for Cul- tural Heritage, the Superintendence for Archaeological Heritage of - and Benevento, the Campania Region and the Municipality of Pietraroja. The Paleo-Lab - dedicated primarily to a school audience - is desi- gned to allow visitors travelling through the geological time, brin- ging them to know the main features of Pietraroja’s past. The tour starts with a “geological lift” thanks to which the visitor travels, in a few minutes, through the geological time, up to the Cretaceous. In continuing the tour, the exhibits and the scenography reconstruct the ancient conditions of the area, allowing to know the ancient Pietraroja and the organisms that populated it. A subsequent room allows the visitor to retrace (and discover) the geological events that led to the Pietraroja Cretaceous lagoon becoming today a por- tion of the Matese Apennines. The last rooms of the Paleo-Lab are dedicated to the evolutionary history of living forms on Earth. The- re is also a space dedicated to the educational laboratory, equip- ped with materials that illustrate the tools used in research labora- tories by geologists and paleontologists. The building adjacent to the Paleo-Lab is being renovated, and will be available to the Ente Geopaleontologico di Pietraroja for the set up of a research center equipped with library, educational and re- search laboratories.

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STOP 4 - Cava Canale

Filomena Ornella Amore

In this locality, in a disused quarry, we will observe the same “Cal- cari ittiolitici” outcropping at “Le Cavere”, but fractured and with traces of bio-ero- sion. These “Calcari ittiolitici” are overlaid by the Miocene sediments of the Cusano (Fig. 8) and Longano formations (Selli, 1957); in front of the quarry, on the opposite side of the road, the sediments of Pietraroja Formation (Selli, 1957).

Cusano Formation The Cusano Fm consists of calcarenites and calcirudites, known as “Calcari a Brio- zoi e Lithothamni” (Bryozoans and Lithothamnium Limestones) and characterized by thick massive banks generally without stratification (Fig. 9). The fossil content is mainly constituted by coralline algae (Archeolithothamnium sp., Lithophyllum sp.), benthic foraminifera (Miogypsinidae, Amphisteginidae, Textularidae, Milioli- dae), encrusting bryozoans (Cyclostomata - Crisiidae, Tubuliporidae; Cheilostoma- ta - in particular Porella cervicornis), serpulids, large lamellibranchs (Pettinids and Ostreids; Fig.10a), echinoderms (Echinolampas sp., Scutella sp.) and arthropods (Balanus sp.). There are also local enrichments of Teleostei and Selacei fish teeth (Fig.10c). Encrusting coralline algae can form large rhodoliths: roundish calcareous bodies, with a nucleus of a different nature, around which algal colonies have de- veloped (Fig. 10d). The sediments were deposited in a neritic marine environment of an open carbonate platform at depths of between about 50 and 200 m and with Fig. 8 – Cava Canale The Early Cretaceous “Calcari sea-water temperatures similar to those of the current temperate zones. The Cusa- ittiolitici” overlaid by the Miocene sediments of no Fm sediments (Fig. 10b), transgressive on Cretaceous sediments are regarded the Cusano Formation as Burdigalian – Langhian in age, basing on the occurrence; in the basal interval, of Miogypsina spp. and of Pecten pseudobeudanti (Barbera et al., 1978; Carannante & Simone, 1996; Ruggiero et al., 2005).

Longano Formation The Longano Fm (Fig. 11) was established by Selli in 1957 on sediments outcrop- ping near Longano (Isernia). It represents the continuation upwards of the Briozoi and Litotamni limestones of the Cusano Fm. These are marly limestone, marl and clayey marl rich in planktonic foraminifers including Orbulina spp., and are also called “Calcari and Marne ad Orbulina”.

This Formation is Serravallian-Tortonian in age, due to the occurrence of Orbulina universa from the bottom and Globorotalia acostaensis and Globigerinoides obli- quus extremus in the upper part (Ciampo et al., 1983; Sgrosso, 1998; Lirer et al., 2005). The sediments of Longano Formation are indicative of open and deep sea environmental condition. At the transition between the two formations it is pos- sible to observe an interval, about 150 cm thick, very rich in glauconitic grains. Fig. 9 – The Miocene succession of the Matese These grains have fecal origin (fecal pellets), or derive from the precipitation of Massif phosphates in micro-environments such as chambers of foraminifera, sporangic

Fig. 10 - From left: i) Civita di Pietraroja. Limestones of the Cusano Fm overlaying the Early Cretaceous ichnolitolytic limestones showing deep bioerosions; ii) Cava Canale. Pecten in the Cusano Formation; iii) Cava Canale fish teeth in the Cusano Formation; iV) Cava Canale Rhodoliths in the Cusano Formation

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Fig. 11 - Longano Formation, Regia Piana area Fig. 12 - Regia Piana, Longano Formation (lower part) containing an echinid cavities of melobesian algae, in conditions of poor or absent sedimentation. In the basal part of the “Marne a Orbulina”, there are elements of Cusano Fm, with numerous fossils, including echinids (Fig.12), followed by sediments containing irregular echinids, brachiopods and fish teeth; upwards the marls become calcilutitic bearing numerous limivorous traces. The sediments of this formation testify the drowning of the platform starting from the Serravallian, owing to important tectonic events (Amore et al., 2001).

Pietraroia Flysch In front of the quarry, on the opposite side of the road, the se- diments of Pietraroja Formation (Selli, 1957) outcrops (Fig.13). These deposits, consisting of well-stratified clays and sandstones with evident turbiditic structures of deep basin, are Tortonian in age on the basis of occurrence of Globigerinoides obliquus extre- mus - planktonic foraminifera- and of Discoaster surculus - cal- careous nannoplankton (Ciampo et al., 1987; Lirer et al., 2005). They represent the evolution of marly limestone and marl of the Longano Fm and the onset of the terrigenous sedimentation in the basin, which is related to the kinematic evolution of the area. These synorogenic deposits, which can reach as many as hun- dreds of meters in thickness, are called flysch and show typical turbidite sedimentary structures; they consist predominantly of silicoclastic material, deriving from the erosion of the foredeep, as well as older sedimentary, igneous and metamorphic rocks. (Ciampo et al., 1987). Fig. 13 – Cava Canale opposite side of the road sediments of the Pietraroia Formation

STOP 4 - Regia Piana bauxite mines

Antonello Bartiromo

Regia Piana (the area we will visit) and Bocca della Selva (about 4 km West of Regia P.), in the municipality of (Mate- se Mountains), are two areas of geological interest for the presence of a spectacular outcrop of sediments of Longano Formation and of bauxite (Figs. 14, 15), a rock that represents the raw materials for the extraction of metallic aluminium. Almost all of the aluminum that has ever been produced has been extracted from bauxite. Bauxites developed from the weathe- ring of alumosilicate-rich parent rocks. These residual deposits are mainly formed under humid tropical to sub-tropical climates, with rainfalls in excess of 1.2 m and annual mean temperatures higher than 22 °C (Bárdossy & Aleva, 1990). Aluminum in bauxi- tes is known to be precipitated in the form of gibbsite [Al(OH)3] or amorphous Al-hydroxides. Iron is separated from aluminum and is frequently concentrated as hematite and goethite. The bauxite deposits from Matese Mountains form flat, contiguous lenses of a few meters in thickness over shallow karst to- pography. These bauxite deposits are at present uneconomic due to their small dimensions and scattered distribution. Bauxite from Matese Mountains has been mined from 1901 -when Michele Monti investigated the bauxite possibilities and obtain license to extract bauxite from Regia Piana locality- through 1963. In particular, the quarrying of bauxites from the Cusano Mutri deposit was done in 1919-1921 by Società Anonima Monte Mutri and then from 1939 by the Società Anonima Montecatini, till 1963 (Del Prete et al., 2002).The mining of bauxite in Matese Mountains had a slow start in the first ten years after the ore was discovered. The bauxite extracted from the locality of Regia Piana had a relatively great commercial importance at local level.

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We will visit 1) the tunnel of extraction located at 50 m from the paved road and 2) the mining area in which a cableway was made for transportation of bauxite ore toward the villa- ge of Cusano Mutri.

The tunnel of extraction are composed of the sub- horizontal gallery with an average width of 1,6 meters and an average height of 2 meters. They have a length of 10 meters till more than 1500 meters. Today only few mines are open for explo- ration because some others are obstructed by lan- dslides. The tunnels are no desert but represent an important shelter for the local fauna. Bauxite deposits, occur along a Cretaceous strati- graphic gap in the Mesozoic carbonate platforms of Central and : in the Abruzzi-Moli- se, and in Campania districts. In the Cam- pania region (southern Italy) two small Cretaceous bauxite districts exist: in the Matese Mountains (Regia Piana and Bocca della Selva localities) and in the province (San Felice, Dragoni and Fig. 14 - Simplified geological map of the southeastern Matese Mountains. 1. Maiorano localities). The bauxites of the Matese Shallow-water platform carbonates (Upper Jurassic-Lower Cretaceous); 2. Pie- Mts. and the Caserta province are incorporated traroja “Plattenkalk” (Lower Albian); 3. Bauxite deposits (Late Albian-Turonian.); 4. Shallow-water platform carbonates (Turonian-Senonian) locally passing to in a thick Meso-Cenozoic carbonate succession, pelagic mudstones and marls (); 5. Open shelf-ramp carbonates (Cu- which builds the main part of the Matese and of sano Formation, Burdigalian-Langhian) and overlying slope marls and mudsto- the Mt. Maggiore mountain chains (Carannante et nes (Longano Formation, Serravallian-Lower Tortonian); 6. Foredeep, siliciclastic al., 2009). turbidites and pelites (Pietraroja flysch, Tortonian-Messinian?); 7. Basinal cherty During the late Aptian-Coniacian (early Senonian) carbonates (Albian-Eocene); 8. Quaternary deposits; 9. Faults. The location of the Apennine platform, as the Apulian platform Bocca della Selva and Regia Piana bauxitic levels are showed in the map. Simpli- were punctuated by repeated and long lasting fied geological map after sheet number 162, , of Geological Map of emersions, locally testified by bauxite deposits Italy, 1:100.000 scale. Servizio Geologico d’Italia, Roma, 1970, and several other (D’Argenio & Mindszenty, 1995). The varied time- sources. From Bartiromo (2013) span of the “mid-Cretaceous” stratigraphic gaps supports the hypothesis of a complex tectonically-controlled paleo-topography and of a differential evolution of the related subdomains (Carannante et al., 1994). D’Argenio & Mindszenty (1995) hypothesized that a lithospheric bulge, induced by the early phases of orogenic collision, had been responsible for a long-lived Cretaceous exposure of some sectors of the southern Apennines shelf domain (as in Apulia). More recently, Schettino & Turco (2011), propose instead the existence of a Cretaceous to Cenozoic E-W, left-lateral, strike slip fault crossing southern Italy, represen- ting the boundary between Adria s.s. and Apulia plates. This fault, operating in a transpressive stress regime, should have induced the uplift of the carbo- nate platforms and the subaerial exposure. Either way, the subaerial exposu- res resulted in extensive karstification and, generally, in bauxite deposition (Carannante et al., 1987, 1994). The bauxite bodies are unconformably cove- red by Upper to Coniacian carbonate sediments. Below the un- conformity, the limestone is extensively karstified and preserves a complex diagenetic record consisting of multiple events of dissolution, cementation, and internal sedimentation (D’Argenio et al., 1986). In the localities of Regia Piana and Bocca della Selva the stratigraphic gap extends from Middle-Up- per Albian to Turonian-Lower Coniacian (Carannante et al., 2009; Catenacci et al., 1963; Crescenti & Vighi, 1970). As to the source material of the bauxites (Fig. 15), most authors (Bárdossy et al., 1977; D’Argenio et al., 1986; D’Argenio & Mindszenty, 1995) support the idea of possible windblown fine pyroclastics of acid to intermediate compo- sition, and exclude any contribution from the insoluble residues of the host carbonates. Though several intense palaeoweathering periods have been Fig. 15 - Bauxite from Regia Piana recorded in the European continent from Upper Paleozoic to Mesozoic, the Cretaceous-Eocene interval is considered the most favourable for bauxitiza- 18 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019 tion. Many of the circum-Mediterranean bauxites, including those located in the Italian peninsula are Cretaceous in age, all formed in karstic environments (karst bauxites) of exhumed carbonates, which behaved as both a physical and chemical trap (Mondillo et al., 2011). The textures of the bauxite ore from Regia Piana is mainly oolitic to pisolitic and generally “impure”, due to the occurrence of Al(- Fe) hydroxides together with almost ubiquitous clays in the matrix. Abundant angular fragments of hematite-goethite occur as mm sized clasts in the bauxite of Bocca della Selva. The texture of the bauxite at Regia Piana is predominantly ooids-supported (Fig. 16 Ooids-supported bauxite.) and it is possible to find locally graded-bedded levels. The cores of the ooids generally consist of older, detrital bauxite grains (bauxite pebbles), or of hematite-goethite fragments (Fig. 17 - Composite oolite with a hematite- goethite fragment at the core). There are differences in the mineral distribution between the ooids and the groundmass. Gene- rally, the Al-hydroxides boehmite and the Fe-mineral hematite seem to be more abundant in the ooids, whereas the clay mineral kaolinite is (generally secondarily) enriched in the groundmass. The presence of bauxite grains/pebbles and hematite-goethite angular fragments occurring among the ooids suggests that at least part of the material occurring in the actual deposits derives from the erosion of older bauxites, formed elsewhere on the exposed carbonate platform. A multiple reworking of the original lateritic soils seems to have occurred, with transport and widespread deposition of the bauxite material on a karst landscape; therefore in the deposits of Matese Mts. it has never been possible to find a typical lateritic bauxite profile in situ on an alumosilicate protore. Probably, an in situ weathering and diagenetic leaching, should have take place after the transport and subsequent reworking of the original lateritic material. Compared to the deposits of Caserta province and the Matese Mountains, the Regia Piana deposit has the highest alumina con- tent. As for the Matese Mountains, the Al2O3 content varies between values of 46.03-48.57 wt.% at the Bocca della Selva deposit and of 67.52 wt.% at the Regia Piana deposit. Fe2O3 content is higher at Bocca della Selva (35.39-38.95 wt.%), whereas is very low in the samples from Regia Piana (about 10 wt.%). In respect to the districts of Caserta and Matese Mountains, the Regia Piana bauxite has the highest Al2O3 and TiO2 values and the lowest Fe2O3 and Cr2O3 values. SiO2 and TiO2 have average values of 5.50 wt.% and 2.68 wt.%, respectively. Most lateritic bauxites can be directly related, through their textures and composition, to the underlying source rocks (e.g., Bardossy and Aleva, 1990), but this is rarely the case for bauxites developed above carbonate sequences. Despite the well- constrained stratigraphic framework for bauxite formation, the origin of the Cretaceous karst bauxites in the Campania district, as of others in Southern Italy, remains controversial. The dissolution of the host carbonate to leave an insoluble residue is not considered a viable mechanism for bauxite formation, because the host carbonates are too pure (Bárdossy et al., 1977; D’Arge- nio et al., 1986; D’Argenio & Mindszenty, 1995). As Mondillo et al. (2011) claimed, the Mesozoic paleogeography and the Bahamian type character of the Apennine carbonate platform, do not support the idea that the parental source materials for the Campania bauxite were transported on the platform through a hydrographic network, as stated by previous authors (Bárdossy et al., 1977; D’Argenio & Mindszenty, 1995). Even if the origin of the Cretaceous karst bauxites in the Campania district remains controversial, there is a strong possibility that bauxites in the Southern Apennines may have formed from windblown pyrocla- stics (D’Argenio & Mindszenty, 1995) that covered the platform carbonates as a thin blanket, and were then subjected to lateritization and local remobilization, as in the case of the Jamaican bauxites (Comer, 1974). At this stage it is not possible to clearly distinguish if the source material derives from a contemporary volcanism or from the erosion of an exposed continental terrain (or rather from both). In addition, in the Apennine carbonate platform volcanic input played an important role as is also hi- ghlighted for plant-bearing fossil-lagerstätten (Bartiromo, 2013; Graziano et al., 2016; Bartiromo et al., in press). It is worth to note that the so-called “livello ad Orbitoline” from Mt. Tobenna and the deposits of Bauxite in Campania have the same detrital mi- neral association (zircon, monazite, rutile, ilmenite). These similarities should suggest Fig. 16 - Regia Piana. Ooids-supported bau- a possible common source material for the two sedimentary lithologies, deposited in xite. From Mondillo et al. (2011) a continental environment in the bauxite case, and in a marine environment in the “livello a Orbitoline” case. From the “Regie Piane” it is also possible to glimpse the location of two recently disco- vered Cretaceous (Aptian) fossil sites (Bartiromo et al., 2012; Graziano et al., 2016) no- teworthy for their palaeontological content. The late Aptian plant assemblage (Barti- romo et al., 2012) collected near the Monte Cigno mountain (Cusano Mutri) consists mainly of impressions and compressions of sterile foliage shoots and reproductive structures of conifers. The genera Cupressinocladus, Pagiophyllum, Araucarites and Frenelopsis have been found. On the basis of macroscopical and cuticular characters as seen under light and scanning electron microscopes a new cheirolepidiacean spe- cies, Frenelopsis cusanensis, was described. It is worth to note also the presence of an Fig. 17 - Regia Piana. Composite oolite with aquatic angiosperm living and reproducing below the surface of the water, Montse- a hematite-goethite fragment at the core. chia vidalii (Fig. 18). The species is for the first time recorded outside of . Mont- From Mondillo et al. (2011) sechia, open the possibility that aquatic plants were locally common at a very early 19 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

stage of angiosperm evolution and that aquatic habitats may have played a major role in the diversification of some early angiosperm lineages. (Gomez et al. 2015). Xeromorphic features displayed by almost all taxa suggest semi-arid or arid con- ditions in a subtropical or tropical climate. The floral assemblage displays a strong affinity with the Euro-Sinian Province of the Northern Hemisphere, which is con- firmed by the absence of typical Gondwanan representatives. This new late Ap- tian fossil site contains also vertebrates and a plethora of invertebrates. The second Upper Aptian fossil plant-bearing locality (Graziano et al., 2016) inclu- des several decimeter-to-meter scale lacustrine intervals straddling a meter-scale plant-rich Plattenkalk and is located in the area named Civitella Licinio (Cusano Mutri). The monogeneric parautochthonous plant remains (Frenelopsis sp.) were deposited in a supratidal-to-paralic coastal mudflat close to a restricted, shallow- marine lagoon, at the verge of an arid–semiarid climatic phase. SEM and EDS analyses documented common to abundant windblown volcanic particles (glass shards and sanidine crystals) throughout the Frenelopsis-rich Plattenkalk (~118.3 to ~118.2 My).

Fig. 18 - Montsechia vidalii showing the typical “bouquet”. From Bartiromo et al. (2012)

References Alvin K.L. (1983). Reconstruction of a Lower Cretaceous conifer. Botanical Journal of the Linnean Society, 86: 169-176. Amato V., Aucelli P.P.C., Cesarano M., Pappone G., Rosskopf C. M. & Russo Ermolli E. (2011). The Sessano intra-montane basin: New multi-proxy data for the Quaternary evolution of the Molise sector of the Central-Southern Apennines (Italy). Geomorphology, 128: 15–31. Amore O., Anzalone E., Barbera C., Cavallo S., Conte M., Donadio C., Fiano V., B., Nazzaro R., Petti C., Raia P., Ruggiero E., Sgrosso I., Taddei A., Taddei R., Tangredi R., Varriale A., Vigliotti M. (2001). Geologia e ambiente nei Parchi Regionali del Matese e del Taburno-Camposauro (Campania). Atti V Conf. Naz. ASITA, 1: 93–98. Aucelli P., Cesarano M., Di Paola G., Filocamo F. & Rosskopf C.M. (2013) Geomorphological map of the central sector of the Matese Mountains (Southern Italy): an example of complex landscape evolution in a Mediterranean mountain environment. Journal of Maps, 9:4, 604-616. Barbera C. & Macuglia L. (1988). Revisione dei tetrapodi del Cretacico inferiore di Pietraroia (Matese orientale, Benevento) appartenenti alla collezione Costa del Museo di Paleontologia dell’Università di Napoli. Memorie della Società Geologica Italiana, 41: 567-574. Barbera C., Simone L. & Carannante G. (1978). Depositi circalittorali di piattaforma aperta nel Miocene Campano. Analisi sedimentologica e paleoecologica. Bollettino della Società Geologica Italiana, 97: 821-834 Bárdossy G. & Aleva G.J.J. (1990). Lateritic bauxites. Amsterdam, Elsevier, 624 pp. Bárdossy G., Boni M., Dall’Aglio M., D’Argenio B. & Pantò G. (1977). Bauxites of peninsular Italy. Composition, origin and geotectonic significan- ce. (Gebr. Borntraeger). Monographic series on Mineral Deposits, 15: 1–-61. Bartiromo A. (2008). Studio paleobotanico dei giacimenti del Cretacico della Campania (Italia). Aspetti tafonomici, paleoecologici, stratigrafici e sedimentologici (PhD Thesis). Università degli Studi di Napoli “Federico II” (in Italian, with English and French abstract). Bartiromo A. (2013). Plant remains from the Lower Cretaceous Fossil-Lagerstätte of Pietraroja, Benevento, southern Italy. Cretaceous Research, 46: 65-79. Bartiromo A., Barale G., Barone Lumaga M.R., Bravi S. & Barattolo F. (2012). An Early Cretaceous flora from Cusano Mutri, Benevento, southern Italy. Cretaceous Research, 33: 116-134. Bartiromo A., Barone Lumaga M. R. & Bravi S. (2006a). First finding of a fossil fern (Matoniaceae) in the paleontological site of Pietraroja (Bene- vento, Southern Italy). Bollettino della Società Paleontologica Italiana, 45 (1): 29-34. Bartiromo A., Bravi S. & Barone Lumaga M.R. (2006b). First finding of a fossil fern (Matoniaceae) in the paleontological site of Pietraroja (Bene- vento, Southern Italy). Bollettino della Società Paleontologica Italiana, 45: 29-34. Bartiromo A., Graziano R., Raspini A. & Bravi S. (in press). A new terrestrial plant-rich Fossil-Lagerstätte from the middle Cenomanian (Late Cretaceous) of the Apennine Carbonate Platform (Magliano Vetere, southern Italy): depositional and palaeoenvironmental settings. Sedi- mentary Geology. Bassani F. (1885). Risultati ottenuti dallo studio delle principali ittiofaune cretaciche. Rendiconti dell’Istituto Lombardo, Milano, 18: 513-535. Bassani F. (1892). Marmi e calcare litografico di Pietraroja (Provincia di Benevento).Rendiconti del Regio Istituto d’Incoraggiamento, fascicoli 7 e 8 e Luglio e Agosto 1892, 1-4. Bonardi G., Ciarcia S., Di Nocera S., Matano F., Sgrosso I. & Torre M. (2009). Carta delle principali unità cinematiche dell’Appennino meridionale. Nota illustrativa. Italian Journal of Geosciences, 128: 47-60. Bonardi G., Amore F.O., Ciampo G., De Capoa P., Miconnet P. & Perrone V. (1988). Il complesso Liguride Auct.: stato delle conoscenze e problemi aperti sulla sua evoluzione pre-appenninica ed i suoi rapporti con l’Arco Calabro. Memorie della Società Geologica Italiana, 41: 17-35. Bravi S. (1987). Contributo allo studio del giacimento ad ittioliti di Pietraroja (Benevento). Dipartimento di Paleontologia, Università degli Studi di Napoli “Federico II”, unpublished degree thesis. Bravi S. (1988). Contributo allo studio del giacimento ad Ittioliti di Pietraroja (Benevento). I. Pleuropholis decastroi n.sp. (Pisces, , Pholidophoriformes). Memorie della Società Geologica Italiana, 41 (I): 575-586. Bravi S. (1994). New observations on the Lower Cretaceous fish Notagogus pentlandi Agassiz (Actinopterygii, Halecostomi, Macrosemiidae). Bollettino della Società Paleontologica Italiana, 33: 51-70. Bravi S. (1999). A tentative reassessment of the fauna and flora from the Pietraroja plattenkalk (Bn). In: Third International Symposium on Li-

20 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

thographic Limestones. Renesto S. (ed.). Rivista del Museo Civico di Scienze Naturali “E. Caffi”, 20: 39–41. Bravi S. & De Castro P. (1995). The Cretaceous fossil fishes level of Capo d’Orlando, near Castellammare di Stabia (NA). Biostratigraphy and depositional environment. Memorie di Scienze Geologiche, 47: 45-72. Bravi S. & Garassino A. (1998). New biostratigraphic and paleoecological observations on the Plattenkalk of the lower Cretaceous (Albian) of Pietraroia (Benevento, S. Italy), and its decapod crustacean assemblage. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 138: 119-171. Breislak S. (1798). Topografia fisica della Campania. Firenze. Briggs D. E. G. (2003). The role of decay and mineralization in the preservation of soft-bodied fossils. Annual Review Earth and Planetary Scien- ces, 31: 275-301. Buscalioni A.D., Piras P., Vullo R., Signore M. & Barbera C. (2012). Early eusuchia crocodylomorpha from the vertebrate-rich Plattenkalk of Pie- traroia (Lower Albian, southern Apennines, Italy). Zoological Journal of the Linnean Society, 163: 199-227. Carannante G., D’Argenio B., Ferreri V. & Simone L. (1987). Cretaceous paleokarst of the Campanian Appennines from early diagenetic to late filling stage. A case history.Rendiconti della Societa Geologica Italiana, 9: 251-256. Carannante G., D’Argenio B., Dello Iacovo B., Ferreri V., Mindszenty A. & Simone L. (1988). Studi sul carsismo cretacico dell’Appennino Campano. Memorie della Società Geologica Italiana, 41: 733-759. Carannante G., Esteban M., Milliman J. & Simone L. (1988). Carbonate facies as paleolatitude indicators: problems and limitations. Sedimentary Geology, 60: 333-346. Carannante G., Pugliese A., Ruberti D., Simone L., Vigliotti M. & Vigorito M. (2009). Evoluzione Cretacica di un settore della piattaforma apula da dati di sottosuolo e di affioramento (Appennino campano-molisano).Bollettino delle Società Geologica Italiana, 128: 3-31. Carannante G., Pugliese A., Simone L. & Vigorito M. (2004). A Cretaceous tectonically controlled carbonate margin: the case history of the Ma- tese Mountains, central-southern Apennines, Italy. 23rd IAS Meeting of Sedimentology, Coimbra. Abstract Volume: 79. Carannante G., Signore M. & Vigorito M. (2006). Vertebrate-rich Plattenkalk of Pietraroia (Lower Cretaceous, Southern Apennines, Italy): a new model. Facies, 52: 555-577. Carannante G. & Simone L. (1996). Rhodolith facies in the Central-Southern Apennines Mountains (Italy). In: Models for Carbonate Stratigraphy from Miocene Reef Complexes of the Mediterranean Regions (E. Franseen et al., eds.). SEPM, Concepts in Sedimentology and Series, 5: 261-275. Catenacci V., De Castro P. & Sgrosso I. (1963). Complessi guida del Mesozoico calcareo dolomitico nella zona orientale del massicio del Matese. Memorie delle Società Geologica Italiana, 4: 1-20. Catenacci E. & Manfredini M. (1963). Osservazioni stratigrafiche sulla Civita di Pietraroia (Benevento). Bollettino della Società Geologica Italiana, 82: 65-92. Chin K., Eberth D. A., Schweitzer M. H., Rando T. A., Sloboda W. J. & Horner J. R. (2003). Remarkable preservation of undigested muscle-tissue within a late Cretaceous tyrannosaurid from Alberta, Canada. Palaios, 18 (3): 286-294. Chin K. & Bishop J. (2007). Exploited twice: bored bone in a theropod coprolite from the Jurassic Morrison Formation of Utah, USA. In: Sedi- ment-Organism Interactions: A Multifaceted Ichnology. Bromley R. G., Buatois L. A., Mángano M. G., Genise J. F. & Melchor R. N. (eds.). SEPM Special Publications, 88: 379-387. Ciampo G., Sgrosso I. & Ruggiero Taddei E. (1987). Inizio della sedimentazione terrigena nel Matese nei Monti del Casetano e nei Monti di Suio. Bollettino della Società Geologica Italiana, 106: 323-330. Comer J.B. (1974). Genesis of Jamaican bauxite. Economic Geology, 69: 1251-1264. Costa O. G. (1851). Cenni intorno alle scoperte fatte nel Regno riguardante la Paleontologia. Il Filiatre Sebezio, Napoli, 21: 40-46. Costa O. G. (1853-1864). Paleontologia del Regno di Napoli, I-III. Atti Accademia Pontaniana, 5, 7, 8. Costa O. G. (1865). Studi sopra i terreni ad Ittioliti delle Provincie napolitane diretti a stabilire l’età geologica de’ medesimi. Parte II: Calcarea stratosa di Pietraroja. Atti dell’Accademia delle Scienze Fisiche e Matematiche di Napoli, s. 2° , 2 (16): 1-33. Costa O. G. (1866). Nuove osservazioni e scoperte intorno ai fossili della calcarea ad ittioliti di Pietraroja. Atti dell’Accademia delle Scienze Fisiche e Matematiche di Napoli, s.1°, 2 (22): 1-12. Crescenti V. & Vighi L. (1964). Caratteristiche, genesi e stratigrafia dei depositi bauxitici cretacici del Gargano e delle Murge; cenni sulle argille con pisoliti bauxitiche del Salento (Puglie). Bollettino della Società Geologica Italiana, 83: 285-337. Dal Sasso C. & Signore M. (1998). Exceptional soft-tissue preservation in a theropod dinosaur from Italy. Nature, 392: 383-387. Dal Sasso C. (2001). Dinosauri italiani. Marsilio Editore, Venezia. Dal Sasso C. (2004). Dinosaurs of Italy. Indiana University Press, Bloomington. Dal Sasso C. & Maganuco S. (2011). Scipionyx samniticus (: Compsognathidae) from the Lower Cretaceous of Italy. Osteology, onto- genetic assessment, phylogeny, soft tissue anatomy, taphonomy and palaeobiology. Memorie della Società italiana di Scienze naturali e del Museo civico di Storia naturale di Milano, Vol XXXVII, Fascicolo I. Dal Sasso C., Marramà G. & Carnevale G. (2014). Vertebrates from the uppermost stratigraphic sequence of the Pietraroja Plattenkalk (Early Cretaceous, southern Italy) (poster). XII EAVP Meeting. Torino, 24-28 June 2014 – Abstract Book, p. 45. Del Prete S., Mele R., Allocca F. & Bocchino B. (2002). Le miniere di bauxite di Cusano Mutri. (Monti del Matese – Campania). Opera Ipogea, 1: 3-34. Doglioni C. (1991). A proposal for the kinematic modelling of W-dipping subductions-possible applications to the Tyrrhenian-Apennines system. Terra Nova, 3: 423-434. D’Argenio B. (1963). I calcari ad ittioliti del Cretacico inferiore del Matese. Atti della Accademia di Scienze Fisiche e Matematiche, Napoli, 4(3): 1-63. D’Argenio B. & Mindszenty A. (1995). Bauxites and related paleokarst: Tectonic and climatic event markers at regional unconformities. Eclogae Geologicae Helvetiae, 88: 453-499. D’Argenio B., Mindszenty A., Bárdossy G., Juhász E. & Boni M. (1986). Bauxites of southern Italy revisited. Rendiconti della Societa Geologica Italiana, 9: 263-268. D’Argenio B., Pescatore T. & Scandone P. (1973). Schema geologico dell’Appennino Meridionale (Campania, Lucania). Accademia Nazionale dei Lincei, Quaderno 183: 49-72. D’Erasmo G. (1941). Niccolò Braucci da Caivano (1719e1774) e della sua opera inedita dal titolo Istoria Naturale della Campania Sotterranea.

21 Paleodays 2019 - XIX Edizione delle Giornate di Paleontologia Benevento/Pietraroja (21)22-24(25) Maggio 2019

Atti della Reale Accademia delle Scienze fisiche e matematiche della Società Reale di Napoli, 2 (Serie 3a N): 2. D’Erasmo G. (1914-15). La fauna e l’età dei calcari ad ittioliti di Pietraroia (Prov. di Benevento). Palaeontographia italica, 20: 29-111. Estes R. (1983). Sauria Terrestria, Amphisbaenia. In: Handbuch der Paläoherpetologie 10A. Wellnhofer P. (ed.). Gustav Fischer Verlag. Evans S. E., Raia P. & Barbera C. (2004). New lizards and rhynchocephalians from the Lower Cretaceous of southern Italy. Acta Palaeontologica Polonica, 49 (3): 393–408. Freels D. (1975). Plattenkalk-Becken bei Pietraroia (Prov. Benevento, S-Italien) als Voraussetzung einer Fossillagerstätten-Bildung. Neues Jahr- buch für Geologie und Paläontologie Abh., 148: 320-352. Gomez B., Daviero-Gomez V., Coiffard C., Martín-Closas C. & Dilcher D.L. (2015). Montsechia, an ancient aquatic angiosperm. Proceedings of the National Academy of Sciences USA, 35: 10985-10988. Gomez B., MartíneClosas C., Barale G., De Porta N.S., Thévenard F. & Guignard G. (2002b). Frenelopsis (Coniferales: Cheirolepidiaceae) and rela- ted male organ genera from the Lower Cretaceous of Spain. Palaeontology, 45: 997-1036. Gomez B., Thévenard F., Fantin M. & Giusberti L. (2002a). Late Cretaceous plants from Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research, 23: 671-685. Graziano R., Raspini A. & Bartiromo A. (2016). Late Aptian palaeoclimatic turnovers and volcanism: Insights from a shallow-marine and conti- nental succession of the Apennine carbonate platform, southern Italy. Sedimentary Geology, 339: 188-217. Lirer F, Persico D., & Vigorito M. (2005). Calcareous plankton biostratigraphy and age of the Middle Miocene deposits of Longano Formation (Matese, southern Apennines): New biostratigraphic data. Rivista Italiana di Paleontologia e Stratigrafia, 111: 91-108. Martill D. M., Frey E., Sues H-D. & Cruickshank A. R. I. (2000). Skeletal remains of a small theropod dinosaur with associated soft structures from the Lower Cretaceous Santana Formation of northeast Brazil. Canadian Journal of Earth Sciences, 37 (6): 891-900. McGowan G. J. (2002). Albanerpetontid amphibians from the Lower Cretaceous of Spain and Italy: a description and reconsideration of their systematics. Zoological Journal of the Linnean Society of London, 135: 1-32. Mondillo N., Balassone G., Boni M. & Rollinson G. (2011). Karst bauxites in the Campania Apennines (southern Italy): a new approach. Periodico di Mineralogia, 80/3(Spec. Issue): 407-432. Mostardini F. & Merlini S. (1986). Appennino centro meridionale. Sezioni geologiche e proposta di modello strutturale. Memorie della Società Geologica Italiana, 35: 177-20. Nicosia U., Petti F. M., Perugini G., D’Orazi Porchetti S., Sacchi E., Conti M. A., Mariotti N. & Zarattini A. (2007). Dinosaur Tracks as Paleogeographic Constraints: New Scenarios for the Cretaceous Geography of the Periadriatic Region. Ichnos, 14: 69-90. Patacca E. & Scandone P. (1989). Post-Tortonian mountain building in the Apennines. The role of the passive sinking of a relic lithospheric slab. Atti dei Convegni Lincei, 80: 157-176. Patacca E. & Scandone P. (2007). Geology of the Southern Apennines. Bollettino della Società Geologica Italiana, Special Issue, 7: 75-119. Patacca E., Scandone P., Bellatalla M., Perilli N. & Santini U. (1992). La zona di giunzione tra l’Arco appenninico settentrio- nale e l’arco appenni- nico meridionale nell’Abruzzo e nel Molise. Studi Geologici Camerti, vol. spec. CROP, 11 (1991/92): 417-441. Pescatore T. (1988). La sedimentazione miocenica nell’Appennino campano-lucano. Memorie della Società Geologica Italiana, 41: 37-46. Pescatore T., Renda P., Schiattarella M. & Tramutoli M. (1999). Stratigraphic and structural relationships between Meso‐Cenozoic Lagonegro basin and coeval carbonate platforms in southern Apennines, Italy. Tectonophysics, 315: 269-286. Poyato-Ariza F. J. & Wenz S. (2002). A new insight into pycnodontiform fishes.Geodiversitas , 24(1): 139-248. Ruben J. A., Dal Sasso C., Geist N. R., Hillenius W. J., Jones T. D. & Signore M. (1999). Pulmonary function and metabolic physiology of theropod dinosaurs. Science, 283: 514-516. Ruggiero E., Amore F.O., Anzalone E., Barbera C., Cavallo S., Conte M., Fiano V., Massa B., Raia P. & Sgrosso I. (2003). I Geositi del Parco Regionale del Matese: Itinerario da Pesco Rosito a . Geologia dell’ambiente, vol. spec.:110-120. Ruggiero E., Cavallo S., Taddei A. & Taddei R. (2005). I beni geo-ambientali del parco regionale del Matese. CD CD ipertestuale. Università di Napoli Federico II. Sacchi E., Conti M. A., D’Orazi Porchetti S., Logoluso A., Nicosia U., Perugini G. & Petti F. M. (2009). Aptian dinosaur footprints from the Apulian platform (Bisceglie, Southern Italy) in the framework of periadriatic ichnosites. Palaeogeography, Palaeoclimatology, Palaeoecology, 271: 104-116. Schettino A. & Turco E. (2011). Tectonic history of the western Tethys since the Late . Geological Society of America Bulletin, 123: 89-105. Selli R. (1957). Sulla trasgressione del Miocene nell’Italia meridionale. Giornale di Geologia, 26: 1-54 Sgrosso I. (1988a). Nuovi elementi per un più articolato modello paleogeografico nell’Appennino centro-meridionale.Memorie della Società Geologica Italiana, 41: 225-252. Sgrosso I. (1998b). Possibile evoluzione cinematica miocenica nell’orogene centro-sud appenninico. Bollettino della Società Geologica Italiana, 117: 679-724. Signore M., Bucci E., Pede C. & Barbera C. (2005). A new ichthyodectid fish from the Lower Cretaceous of Pietraroja (Southern Italy). PalArch, 5: 25-29. Signore M., Pede C., Bucci E. & Barbera C. (2006). First report of the genus Cladocyclus in Lower Cretaceous of Pietraroja (Southern Italy). Bollet- della Società Paleontologica Italiana, 45: 141-146. Vigorito M., Simone L. & Carannante G. (2003). Tectonically controlled carbonate channelized slope complexes: a Cretaceous-Miocene case- history from Matese Mountains (Central-Southern Apennines, Italy). Slope 2003-Submarine Slope Systems: Processes, Products and Prediction. April 28–29, 2003, Liverpool, Abstracts: 95. Vitale S., Amore F.O, Ciarcia S., Fedele L., Grifa C., Prinzi E.P., Tavani S. & Tramparulo F.D.A. (2017). Structural, stratigraphic, and petrological clues for a Cretaceous–Paleogene abortive rift in the southern Adria domain (southern Apennines, Italy). Geological Journal 2017. https:// doi. org/10.1002/gj.2919 Vitale S. & Ciarcia S. (2018). Tectono-stratigraphic setting of the Campania region (southern Italy). Journal of Maps, 14: 9-21. Wilby P. R. & Briggs D. E. G. (1997). Taxonomic trends in the resolution of detail preserved in fossil soft tissues. Géobios, 20: 493-502. Zappaterra E. (1990). Carbonate paleogeographic sequences of the periadriatic region. Bollettino della Società Geologica Italiana, 109: 5-20.

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