DIGITAL INVESTIGATION OF LAMNIFORM SHARK VERTEBRAE FROM THE SIBILLINI MTS. (NORTHERN APENNINES, ITALY)

Authors

DOI:

https://doi.org/10.54103/2039-4942/22571

Keywords:

Early Miocene; Bisciaro; Dead-fall; CT-SCAN; 3D model.

Abstract

During the sampling of a stratigraphic section along the shore of the Fiastra Lake (Carg Project - Sheet 313 “Camerino” of the Geological Map of Italy at 1:50 000 scale), a small rock boulder with partially exposed bony material was discovered at the base of a small cliff at the northern termination of the Sibillini Mts. In this area, the classical facies of Umbria-Marche stratigraphic succession are well exposed. The Oligocene-Miocene portion of the succession is represented by the ~200 m-thick Scaglia Cinerea Formation, passing upwards to the ~100 m-thick Bisciaro Formation. The microfossil assemblage has allowed the specimen to be constrained to the lower Burdigalian. The skeletal remains were examined using a CT-SCAN, a non-invasive method that has proven to be highly performing. The analysis revealed some articulated vertebrae, deformed by lithostatic compaction, which are attributed to a shark of the Order Lamniformes. Subsequently, the vertebrae were digitally isolated, extracted from the surrounding matrix, and rendered into three-dimensional prints. Through digital retro-deformation, the body length of the lamniform shark was estimated to be approximately 4 metres. Further considerations on the vertebrae allowed us to infer that the studied shark had similarities to either Isurus oxyrinchus Rafinesque, 1810 or Carcharodon carcharias Linnaeus, 1758. The development of a dead-fall microbial community likely facilitated the preservation of the vertebrae. The studied specimen represents the first occurrence of a lamniform shark in the Lower Miocene of the Umbria-Marche Domain and represents one of the very rare recorded occurrences of lamniforms from the Lower Miocene of Italy.

References

Agassiz L. (1843) - Recherches sur les poissons fossiles. Imprimerie de Petitpierre, Neuchatel et Soleure, 1420 pp.

Allison P.A., Smith C.R., Kukert H., Deming J.W. & Bennett B.A. (1991) - Deep-water taphonomy of vertebrate carcasses: a whale skeleton in the bathyal Santa Catalina Basin. Paleobiology, 17: 78-89.

Amalfitano J., Dalla Vecchia F.M., Giusberti L., Fornaciari E., Luciani V. & Roghi G. (2017) - Direct evidence of trophic interaction between a large lamniform shark, Cretodus sp., and a marine turtle from the Cretaceous of northeastern Italy. Palaeogeography, Palaeoclimatology, Palaeoecology, 469: 104-121.

Amalfitano J., Giusberti L., Fornaciari E., Dalla Vecchia F. M., Luciani V., Kriwet J. & Carnevale G. (2019) - Large deadfalls of the "ginsu" shark Cretoxyrhina mantelli (Agassiz, 1835) (Neoselachii, Lamniformes) from the Upper Cretaceous of northeastern Italy. Cretaceous Research, 98: 250-275.

Amalfitano J., Dalla Vecchia F.M., Carnevale G., Fornaciari E., Roghi G. & Giusberti L. (2022) - Morphology and paleobiology of the Late Cretaceous large-sized shark Cretodus crassidens (Dixon, 1850) (Neoselachii; Lamniformes). Journal of Paleontology, 96: 1166-1188.

Antunes M.T., Jonet S. & Nascimento A. (1981) - Vertébrés (Crocodiliens, Poissons) du Miocène marin de l’Algarve occidentale. Ciencias da Terra (UNL), 6: 9-38.

Applegate S.P. (1967) - A survey of shark hard parts. In: Gilbert P. W., Matthewson R. F. & Rail D. P. (Eds) - Sharks, skates and rays: 37-67. John Hopkins Press, Baltimore, MD.

Baier J., Schmitt K.H. & Mick R. (2004) - Notizen zur untermiozänen Hai- und Rochenfauna der Erminger Turritellenplatte (Mittlere Schwäbische Alb, SW-Deutschland). Jahresberichte und Mitteilungen des Oberrheinischen Geologischen Vereins, 86: 361-371.

Barthelt D., Fejfar O., Pfeil F.H. & Unger E. (1991) - Notizen zu einem Profil der Selachier-Fundstelle Walbertsweiler im Bereich der miozänen Oberen Meeresmolasse Süddeutschlands. Münchner geowissenschaftliche Abhandlungen (A: Geologie und Paläontologie), 19: 195-208.

Belben R.A., Underwood C.J., Johanson Z. & Twitchett R.J. (2017) - Ecological impact of the end-Cretaceous extinction on lamniform sharks. PLoS One, 12(6): e0178294.

Beldean C., Filipescu S. & Balc R. (2010) - An Early Miocene biserial foraminiferal event in the Transylvanian Basin (Romania). Geologica Carpathica, 61(3): 227-234.

Berg L. S. (1958) - System der Rezenten und Fossilen Fischartigen und Fische. Deutscher Verlag für Wissenschaft, Berlin, 310 pp.

Bernoulli D. (1967) - Probleme der Sedimentation im Jura Westgriechenlands und des zentralen Apennin. Verhandlungen der Naturforschenden Gesellschaft in Basel, 78: 35-54.

Blanco-Piñón A., Shimada K. & González-Barba G. (2005) - Lamnoid vertebrae from the Agua Nueva Formation (Upper Cretaceous: lower Turonian), northeastern Mexico. Revista mexicana de ciencias geológicas, 22(1): 19-23.

Boessenecker R.W., Perry F.A. & Schmitt J.G. (2014) - Comparative taphonomy, taphofacies, and bonebeds of the Mio-Pliocene Purisima Formation, Central California: strong physical control on marine vertebrate preservation in shallow marine settings. PLoS One, 9(3): e91419.

Bolliger T., Kindlimann R. & Wegmüller U. (1995) - Die marinen Sedimente (jüngere OMM, St. Galler-Formation) am Südwestrand der Hörnlischüttung (Ostschweiz) und die palökologische Interpretation ihres Fossilinhaltes. Eclogae Geologicae Helvetiae, 88(3): 885-909.

Bonaparte C.L. (1838) - Selachorum tabula analytica. Nuovi Annali delle Scienze Naturali, 1: 195-214, Bologna.

Bramlette M.N. & Wilcoxon J.A. (1967) - Middle Tertiary calcareous nannoplankton of the Cipero section, Trinidad. W.I. Tulane Studies in Geology and Paleontology, 5: 93-131.

Brzobohatý R. & Schultz O. (1971) - Die Fischfauna der Eggenburger Schichtengruppe. In: Steininger F. et al. (Eds) - M1 Eggenburgien. Die Eggenburger Schichtengruppe und ihr Stratotypus. Chronostratigraphie und Neostratotypen Miozän der zentralen Paratethys: 719-759. Bratislava: Vydavateľstvo Slovenskej Akadémie Vied.

Buchholtz E.A. & Schur S.A. (2004) - Vertebral osteology in Delphinidae (cetacea). Zoological Journal of the Linnean Society, 140(3): 383-401.

Bukry D. (1971) - Cenozoic calcareous nannofossils from the Pacific Ocean. San Diego Society of Natural History Transactions, 16: 303-327.

Cailliet G.M., Natanson L.J., Welden B.A. & Ebert D.A. (1985) - Preliminary studies on the age and growth of the white shark, Carcharodon carcharias, using vertebral bands. Memoirs of the Southern California Academy of Science, 9: 49-60.

Calamita F. & Deiana G. (1988) - The arcuate shape of the Umbria-Marche-Sabina Apennines (central Italy). Tectonophysics, 146: 139-147.

Calamita F., Pace P. & Satolli S. (2012) - Coexistence of fault-propagation and fault-bend folding in curve-shaped foreland fold-and-thrust belts: examples from the Northern Apennines (Italy). Terra Nova, 24: 396-406.

Cappetta H. (1970) - Les sélaciens du Miocène de la région de Montpellier. Palaeovertebrata, Memoire extraordinaire: 1-139.

Cappetta H. (1973) - Les sélaciens du Burdigalien de lespignan (Hérault). Geobios, 6(3): 211-223.

Cappetta H. (1975) - Les Sélaciens miocènes du Midi de la France, répartition stratigraphique et bathymétrique. In 3 ème reunion annuelle des Sciences de la Terre, p. 90. Montpellier.

Cappetta H. (1987) - Handbook of Paleoichthyology. In: Chondrichthyes II Mesozoic and Cenozoic Elasmobranchii, 3B. Gustav Fischer Verlag, Stuttgart, p. 193.

Cappetta H. (2012) - Chondrichthyes - Mesozoic and Cenozoic Elasmobranchii: Teeth. Handbook of Paleoichthyology. Verlag F. Pfeil, Munich, p. 512.

Carlander K.D. (1969) - An operational-functional classification of fishery management techniques: with 1 table in the text. Internationale Vereinigung für theoretische und angewandte Limnologie: Verhandlungen, 17(2): 636-640.

Carminati E. & Doglioni C. (2012) - Alps vs. Apennines: The paradigm of a tectonically asymmetric Earth”. Earth-Science Reviews, 112: 67-96.

Carrillo-Briceño J.D., Parra J.D. & Luque J. (2019) - A new lamniform shark Protolamna ricaurtei sp. nov. from the Lower Cretaceous of Colombia. Cretaceous Research, 95: 336-340.

Centamore E., Chiocchini M., Deiana G., Micarelli A. & Pieruccini U. (1971) - Contributo alla conoscenza del Giurassico dell’Appennino umbro-marchigiano. Studi Geologici Camerti, 1: 7-89.

Cipriani A. (2016) - Geology of the Mt. Cosce sector (Narni Ridge, Central Apennines, Italy). Journal of Maps, 12: 328-340.

Citton P., Fabbi S., Cipriani A., Jansen M. & Romano M. (2019) - Hybodont dentition from the Upper Jurassic of Monte Nerone Pelagic Carbonate Platform (Umbria-Marche Apennine, Italy) and its ecological implications. Geological Journal, 54(1): 278-290.

Citton P., Romano M., Fabbi S. & Cipriani A. (2021) - Jurassic neoselachian sharks from the Mt Nerone Pelagic Carbonate Platform (Umbria-Marche Apennine, Italy): a further constrain for the palaeoecology related to PCP systems. Historical Biology, 33(9): 1383-1394.

Coccioni R. & Montanari A. (1994) - L’orizzonte del Bisciaro. In: Ciarapica G. & Passeri L. (Eds.) - Appennino Umbro-Marchigiano, Guide Geologiche Regionali, 7: 36-38, BE-MA Ed., Lodi (MI).

Compagno L.J.V. (1973) - Interrelationships of living elasmobranchs. Interrelationships of fishes: 15-61.

Compagno L.J.V. (1977) - Phyletic relationships of living sharks and rays. American zoologist, 17(2): 303-

Compagno L.J.V. (1999) - Chapter 3: endoskeleton. In: Hamlett W.C. (Ed.) - Sharks, Skates, and Rays: the Biology of Elasmobranch Fishes. Johns Hopkins University Press, Baltimore: 69-92.

Cook T.D., Newbrey M.G., Murray A.M., Wilson M.V., Shimada K., Takeuchi G.T. & Stewart J.D. (2011) - A partial skeleton of the Late Cretaceous lamniform shark, Archaeolamna kopingensis, from the Pierre Shale of western Kansas, USA. Journal of Vertebrate Paleontology, 31(1): 8-21.

Cook T.D., Wilson M.V. & Newbrey M.G. (2010) - The first record of the large Cretaceous lamniform shark, Cardabiodon ricki, from North America and a new empirical test for its presumed antitropical distribution. Journal of Vertebrate Paleontology, 30(3): 643-649.

Cosentino D., Cipollari P., Marsili P. & Scrocca D. (2010) - Geology of the central Apennines: a regional review. Journal of the Virtual Explorer, 36: 1-37.

Curzi M., Cipriani A., Aldega L., Billi A., Carminati E., Van der Lelij R., Vignaroli G. & Viola G. (2024) - Architecture and permeability structure of the Sibillini Mts. Thrust and influence upon recent, extension-related seismicity in the central Apennines (Italy) through fault-valve behavior. GSA Bulletin, 136(1-2): 3-26.

D’Erasmo G. (1924) - Ittioliti Miocenici di Rosignano-Piemonte e di Vignale. Memorie descrittive della Carta Geologica d’Italia, 9: 1-37.

Danise S., Twitchett R.J. & Matts K. (2014) - Ecological succession of a Jurassic shallow-water ichthyosaur fall. Nature Communications, 5(1): 4789.

Davis J. W. (1890) - On the fossil fish of the Cretaceous Formations of Scandinavia. Scientific Transactions of the Royal Dublin Society, 4: 363-434.

De Carvalho M.R. (1996) - Higher-level elasmobranch phylogeny, basal squaleans, and paraphyly. Interrelationships of Fishes, 3: 35-62.

De Schutter P.J. & Everaert S. (2020) - A megamouth shark (Lamniformes: Megachasmidae) in the Burdigalian of Belgium. Geologica Belgica, 23: 157-165.

Dean M.N. & Summers A.P. (2006) - Mineralized cartilage in the skeleton of Chondrichthyan fishes. Zoology, 109(2): 164-168.

Deming J., Reysenbach A.L., Macko S.A. & Smith C.R. (1997) - The microbial diversity at a whale fall on the seafloor: bone-colonizing mats and animal-associated symbionts. Microscopy Research and Technique, 37: 162-170.

Di Santo V., Goerig E., Wainwright D.K., Akanyeti O., Liao J.C., Castro-Santos T. & Lauder G.V. (2021) - Convergence of undulatory swimming kinematics across a diversity of fishes. Proceedings of the National Academy of Sciences, 118(49): e2113206118.

Di Stefano A., Baldassini N., Raffi I., Fornaciari E., Incarbona A., Negri A., Bonomo S., Villa G., Di Stefano E. & Rio D. (2023) - Neogene-Quaternary Mediterranean calcareous nannofossil biozonation and biochronology: A review. Stratigraphy, 20(4): 259-302.

Dick D.G. (2015) - An ichthyosaur carcass-fall community from the Posidonia Shale (Toarcian) of Germany. Palaios, 30(5): 353-361.

Doglioni C. (1995) - Geological remarks on the relationships between extension and convergent geodynamic settings. Tectonophysics, 252: 253-267.

Doglioni C., Carminati E. & Cuffaro M. (2006) - Simple Kinematics of Subduction Zones. International Geology Review, 48: 479-493.

Ebersole J.A. & Ehret D.J. (2018) - A new species of Cretalamna sensu stricto (Lamniformes, Otodontidae) from the Late Cretaceous (Santonian-Campanian) of Alabama, USA. PeerJ, 6: e4229.

Fabbi S. (2015) - Geology and Jurassic paleogeography of the Mt. Primo-Mt. Castel Santa Maria ridge and neighbouring areas (Northern Apennines, Italy). Journal of Maps, 11: 645-663.

Fabbi S. & Santantonio M. (2012) - Footwall progradation in syn-rift carbonate platform-slope systems (Early Jurassic, Northern Apennines, Italy). Sedimentary Geology, 281: 21-34.

Fischli H. (1930) - Die Fossilien der Quarzsande von Benken (Kt. Zürich). Mitteilungen der Naturwissenschaftlichen Gesellschaft in Winterthur, 17(18): 131-167.

Flugel E. (2010) - Microfacies of Carbonate Rocks. Springer, 929 pp.

Fornaciari E. & Rio D. (1996) - Latest Oligocene to early Middle Miocene quantitative calcareous nannofossil biostratigraphy in the Mediterranean region. Micropaleontology, 42(1): 1-37.

Frederickson J.A., Schaefer S.N. & Doucette-Frederickson J.A. (2015) - A gigantic shark from the lower Cretaceous Duck Creek Formation of Texas. PLoS One, 10(6): e0127162.

Goedert J., Amiot R., Arnaud-Godet F., Cuny G., Fourel F., Hernandez J.A., Pedreira-Segade U. & Lécuyer C. (2017) - Miocene (Burdigalian) seawater and air temperatures estimated from the geochemistry of fossil remains from the Aquitaine Basin, France. Palaeogeography, Palaeoclimatology, Palaeoecology, 481: 14-28.

Goedert J.L., Squires R.L. & Barnes L.G. (1995) - Paleoecology of whale-fall habitats from deep-water Oligocene rocks, Olympic Peninsula, Washington State. Palaeogeography, Palaeoclimatology, Palaeoecology, 118: 151-158.

Goodrich E.S. (1930) - Studies on the Structure and Development of Vertebrates. Macmillan, London, 837 pp.

Gottfried M.D., Compagno L.J.V. & Bowman, S.C. (1996) - Size and skeletal anatomy of the giant “megatooth” shark Carcharodon megalodon. In: Klimley A.P. & Ainley D.G. (Eds.) - Great White Sharks: The biology of Carcharodon carcharias. San Diego, Academic Press: 55-66.

Guerrera F., Martín-Martín M., Raffaelli G. & Tramontana M. (2015) - The Early Miocene “Bisciaro volcaniclastic event” (northern Apennines, Italy): a key study for the geodynamic evolution of the central-western Mediterranean. International Journal of Earth Sciences, 104: 1083-1106.

Guinot G. & Carrillo-Briceño J.D. (2018) - Lamniform sharks from the Cenomanian (upper Cretaceous) of Venezuela. Cretaceous Research, 82: 1-20.

Guinot G. & Cavin L. (2016) - ‘Fish’ (Actinopterygii and Elasmobranchii) diversification patterns through deep time. Biological Reviews, 91(4): 950-981.

Guinot G., Adnet S. & Cappetta H. (2012) - An analytical approach for estimating fossil record and diversification events in sharks, skates and rays. PloS One, 7(9): e44632.

Hamm S.A. & Shimada K. (2002) - Associated tooth set of the Late Cretaceous lamniform shark, Scapanorhynchus raphiodon (Mitsukurinidae), from the Niobrara Chalk of western Kansas. Transactions of the Kansas Academy of Science, 105(1): 18-26.

Haq B.U. (1966) - Electron microscope studies on some upper Eocene calcareous nannoplankton from Syria. Stockholm Contributions in Geology, 15: 23-37.

Higgs N.D., Gates A.R. & Jones D.O. (2014) - Fish food in the deep sea: revisiting the role of large food-falls. PloS One, 9(5): e96016.

Hogler J.A. (1992) - Taphonomy and paleoecology of Shonisaurus popularis (Reptilia: Ichthyosauria). Palaios, 7: 108-117.

Holec P., Hornacek M. & Sykora M. (1995) - Lower Miocene shark (Chondrichthyes, Elasmobranchii) and whale faunas (Mammalia, Cetacea) near Mucin, southern Slovakia. Geologicke Prace, 100: 37-52.

Höltke O. (2014) - Haifischbecken Ursendorf. Fossilien, 2014(2): 20-25.

Huxley T.H. (1880) - On the application of the laws of evolution to the arrangement of the Vertebrata, and more particularly of the Mammalia. Proceedings of the Scientific Meetings of the Zoological Society of London, 1880: 649-62.

Ingle D.N., Natanson L.J. & Porter M.E. (2018) - Mechanical behavior of shark vertebral centra at biologically relevant strains. Journal of Experimental Biology, 221(24): jeb188318.

Jambura P.L., Kindlimann R., López-Romero F., Marramà G., Pfaff C., Stumpf S., Türtscher J., Underwood C.J., Ward D.J. & Kriwet J. (2019) - Micro-computed tomography imaging reveals the development of a unique tooth mineralization pattern in mackerel sharks (Chondrichthyes; Lamniformes) in deep time. Scientific Reports, 9(1): 9652.

Joleaud L. (1907-1912) - Géologie et Paléontologie de la Plaine du Comtat et de ses abords. Description des terrains Néogènes. Mémoires de l’Academie Vaucluse, 1-2: 1-285.

Jost J., Kempf O. & Kälin D. (2016) - Stratigraphy and palaeoecology of the Upper Marine Molasse (OMM) of the central Swiss Plateau. Swiss Journal of Geosciences, 109(2): 149-169.

Kaim A., Kobayashi Y., Echizenya H., Jenkins R.G. & Tanabe K. (2008) - Chemosynthesis-based associations on Cretaceous plesiosaurid carcasses. Acta Palaeontologica Polonica, 53(1): 97-104.

Koch A. (1904) - Fossile Haifischzähne und Säugetierreste von Felsöesztergály, im Komitate Nógrád. Földtani Közlöny, 34: 260-273.

Kocsis L. (2007) - Central paratethyan shark fauna (Ipolytarnóc, Hungary). Geologica Carpathica, 58(1): 27-40.

Kordos L. & Solt P. (1982) - A magyarországi miocén tengeri gerinces faunaszintek vázlata. A Magyar Állami Földtani Intézet Évi Jelentése, 1984: 347-351.

Kriwet J. (2006) - Biology and dental morphology of Priscusurus adruptodontus, gen. et sp. nov. (Chondrichthyes, Lamniformes) from the Albian (Early Cretaceous) of Peru. Journal of Vertebrate Paleontology, 26(3): 538-543.

Kriwet J. & Benton M.J. (2004) - Neoselachian (Chondrichthyes, Elasmobranchii) diversity across the Cretaceous-Tertiary boundary. Palaeogeography, Palaeoclimatology, Palaeoecology, 214(3): 181-194.

Kriwet J., Klug S., Canudo J.I. & Cuenca-Bescos G. (2008) - A new early Cretaceous lamniform shark (Chondrichthyes, Neoselachii). Zoological Journal of the Linnean Society, 154(2): 278-290.

Kriwet J., Mewis H. & Hampe O. (2015) - A partial skeleton of a new lamniform mackerel shark from the Miocene of Europe. Acta Palaeontologica Polonica, 60 (4): 857-875.

Leriche M. (1927) - Les Poissons de la Molasse suisse. Prémiere fascicule. Mémoires de la Société paléontologique Suisse, 46: 1-55.

Linnaeus C. (1758) - Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Holmiae, Imprensis Laurentii Salvii. 2 vol.

Lutzeier H. (1922) - Beiträge zur Kenntnis der Meeresmolasse in der Ulmer Gegend. Neues Jahrbuch für Mineralogie Geologie und Paläontologie, 1922: 117-180.

Maisch IV H., Becker M. & Chamberlain Jr J. (2015) - Chondrichthyans from a lag deposit between the Shark River Formation (Middle Eocene) and Kirkwood Formation (Early Miocene), Monmouth County, New Jersey. Paludicola, 10: 149-183.

Marsili S., Carnevale G., Danese E., Bianucci G. & Landini W. (2007) - Early Miocene vertebrates from Montagna della Maiella, Italy. Annales de Paléontologie, 93(1): 27-66.

Martill D. M. (1987) - A taphonomic and diagenetic case study of a partially articulated ichthyosaur. Palaeontology, 30(3): 543-555.

Martin A.P. & Naylor G.J.P. (1997) - Independent origins of filterfeeding in megamouth and basking sharks (Order Lamniformes) inferred from phylogenetic analysis of cytochrome b gene sequences. In: Yano K., Morrissey J.F., Yabumoto Y. & Nakaya K. (Eds) - Biology of the megamouth shark. Tokai University Press, Kanagawa: 39-50.

Mas G. & Antunes M.T. (2008) - Presència de Tomistoma cf. lusitanica (Vianna i Moraes, 1945) (Reptilia: Crocodylia) al Burdigalià inferior de Mallorca (Illes Balears, Mediterrània occidental). Implicacions paleoambientals. Bolletí de la Societat d’Història Natural de les Balears, 51: 131-145.

Maxwell E.E., Cooper S.L., Mujal E., Miedema F., Serafini G. & Schweigert G. (2022) - Evaluating the Existence of Vertebrate Deadfall Communities from the Early Jurassic Posidonienschiefer Formation. Geosciences, 12(4): 158.

Milli S., Moscatelli M., Stanzione O. & Falcini F. (2007) - Sedimentology and physical stratigraphy of the Messinian turbidite deposits of the Laga Basin (central Apennines, Italy). Bollettino della Società Geologica Italiana, 126: 255-281.

Morse P.E., Stock M.K., James K.C., Natanson L.J. & Stock S.R. (2022) - Shark centra microanatomy and mineral density variation studied with laboratory microComputed Tomography. Journal of Structural Biology, 214(1): 107831.

Moss M.L. (1977) - Skeletal tissues in sharks. American Zoologist, 17: 335-342.

Natanson L.J. (2001) - Preliminary investigations into the age and growth of the shortfin mako, Isurus oxyrinchus, white shark, Carcharodon carcharias, and thresher shark, Alopias vulpinus, in the Western North Atlantic Ocean. International Commission for the Conservation of Atlantic Tunas Working Document, Standing Committee on Research and Statistics, 01/66.

Naylor G.J., Caira J.N., Jensen K., Rosana K.A., Straube N. & Lakner C. (2012) - Elasmobranch phylogeny: a mitochondrial estimate based on 595 species. The biology of sharks and their relatives, 31-56.

Naylor G.J.P., Martin A.P., Mattison E. & Brown W.M. (1997) - The inter-relationships of lamniform sharks: testing phylogenetic hypotheses with sequence data. In: Kocher T.D. & Stepien C. (Eds.) - Molecular systematics of fishes. Academic Press, New York: 199-218.

Nazarkin M.V. & Malyshkina T.P. (2012) - The first reliable record of selachians from the Neogene deposits of Sakhalin Island. Zoosystematica Rossica, 21(1): 180-184.

Newbrey M.G., Siversson M., Cook T.D., Fotheringham A.M. & Sanchez R.L. (2015) - Vertebral morphology, dentition, age, growth, and ecology of the large lamniform shark Cardabiodon ricki. Acta Palaeontologica Polonica, 60(4): 877-897.

Noffke N. (2021) - Microbially Induced Sedimentary Structures. Encyclopedia of Astrobiology. https://doi.org/10.1007/978-3-642-27833-4_1004-5

Paparella I., Maxwell E.E., Cipriani A., Roncacè S. & Caldwell M.W. (2017) - The first ophthalmosaurid ichthyosaur from the Upper Jurassic of the Umbrian-Marchean Apennines (Marche, Central Italy). Geological Magazine, 154(4): 837-858.

Parotto M. & Praturlon A. (2004) - The Southern Apennine Arc. In: Crescenti U., D’Offizi S., Merlino S. & Sacchi L. (Eds.) - Geology of Italy. Special Volume of the Italian Geological Society for the IGC 32 Florence-2004: 33-58.

Passeri L. & Venturi F. (2005) - Timing and causes of drowning of the Calcare Massiccio platform in Northern Apennines. Bollettino della Società Geologica Italiana, 124: 247-258.

Patacca E., Sartori R. & Scandone P. (1992) - Tyrrhenian basin and Apenninic arcs: Kinematic relations since late Tortonian times. Memorie della Società Geologica Italiana, 45: 425-451.

Petti F. & Falorni P. (2007) - Bisciaro. Carta Geologica d’Italia 1:50.000 - Catalogo delle Formazioni (Unità tradizionali). I Quaderni, serie III, del SGI, 7 (VII): 240-247.

Pierantoni P.P., Deiana G. & Galdenzi S. (2013) - Stratigraphic and structural features of the Sibillini Mountains (Umbria-Marche Apennines, Italy). Italian Journal of Geosciences, 132: 497-520.

Pierantoni P.P., Deiana G., Romano A., Paltrinieri W., Borracini F. & Mazzoli S. (2005) - Structural geometries along the thrust zone of the Umbria-Marche-Sabina mountain front. Bollettino della Società Geologica Italiana, 124: 395-411.

Pollerspöck J. & Beaury B. (2014) - Eine Elasmobranchierfauna (Elasmobranchii, Neoselachii) aus der Oberen Meeresmolasse (Ottnangium, Unteres Miozän) des Heigelsberger Grabens bei Teisendorf, Oberbayern. Zitteliana, A54: 23-37.

Probst J. (1879) - Beiträge zur Kenntnis der fossilen Fische aus der Molasse von Baltringen IV. Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg, 35: 127-191.

Rafinesque C.S. (1810) - Caratteri di alcuni nuovi generi e nuove specie di animali e piante della sicilia, con varie osservazioni sopra i medesimi. Palermo, 105 pp.

Reinecke T., Louwye S., Havekost U. & Moths H. (2011) - The elasmobranch fauna of the late Burdigalian, Miocene, at Werder-Uesen, Lower Saxony, Germany, and its relationship with early Miocene faunas on the North Atlantic, Central Paratethys and Mediterranean. Vol. 20, Belgium: Palaeo Publishing and Library.

Reinecke T., Von Der Hocht F. & Dufraing L. (2015) - Fossil basking shark of the genus Keasius (Lamniforme, Cetorhinidae) from the boreal North Sea Basin and Upper Rhine Graben: evolution of dental characteristics from the Oligocene to late Middle Miocene and description of two new species. Palaeontos, 28: 39-98.

Reisdorf A.G., Bux R., Wyler D., Benecke M., Klug C., Maisch M.W., Fornaro P. & Wetzel A. (2012) - Float, explode or sink: postmortem fate of lung-breathing marine vertebrates. Palaeobiodiversity and Palaeoenvironments, 92: 67-81.

Ridewood W.G. (1921) - On the calcification of the vertebral centra in sharks and rays. Philosophical Transactions of the Royal Society of London. Series B, Containing Papers of a Biological Character, 210(372-381): 311-407.

Romano M., Cipriani A., Fabbi S. & Citton P. (2019a) - First remains of neoginglymodian actinopterygians from the Jurassic of Monte Nerone area (Umbria-Marche Apennine, Italy). Italian Journal of Geosciences, 138(1): 88-102.

Romano M., Citton P., Cipriani A., Fabbi S., Spanò F., Marramà G. & Carnevale G. (2021) - Lamniform vertebrae from the Aptian-Albian Marne a Fucoidi of Umbria-Marche Domain (central Italy). Cretaceous Research, 125: 104832.

Romano M., Manni R., Venditti E., Nicosia U. & Cipriani A. (2019b) - First occurrence of a Tylosaurinae mosasaur from the Turonian of the Central Apennines, Italy. Cretaceous Research, 96: 196-209.

Royden L., Patacca E. & Scandone P. (1987) - Segmentation and configuration of subducted lithosphere in Italy: An important control on thrust-belt and foredeep-basin evolution. Geology, 15(8): 714-717.

Sach V.J. & Heizmann E.P.J. (2001) - Stratigraphy and mammal faunas of the Brackwassermolasse in the surroundings of Ulm (Southwest Germany). Stuttgarter Beiträge zur Naturkunde, Serie B, 310: 1-95.

Santantonio M. & Carminati E. (2011) - The Jurassic rifting evolution of the Apennines and Southern Alps (Italy): Parallels and differences. Geological Society of America Bulletin, 123: 468-484.

Santantonio M., Cipriani A., Fabbi S. & Meister C. (2022) - Constraining the slip rate of Jurassic rift faults through the drowning history of a carbonate platform. Terra Nova, 34(2): 137-145.

Schäfer W. (1972) - Ecology and Paleoecology of Marine Environments. University of Chicago Press, Chicago, Illinois.

Schmeisser McKean R.L. & Gillette D.D. (2015) - Taphonomy of large marine vertebrates in the Upper Cretaceous (Cenomanian-Turonian) Tropic Shale of southern Utah. Cretaceous Research, 56: 278-292.

Schmitz L., Thies D. & Kriwet J. (2010) - Two new lamniform sharks (Leptostyrax stychi sp. nov. and Protolamna sarstedtensis sp. nov.) from the Early Cretaceous of NW Germany. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 257: 283-296.

Scholz H. & Bienert R. (1992) - Bausteine und Fossilien aus der Oberen Meeresmolasse bei Kempten. Berichte des Naturwissenschaftlichen Vereins für Schwaben, 96(2): 2-12.

Schultz O. (2013) - Pisces. Catalogus Fossilium Austriae, 3: 1-576.

Serafini G., Amalfitano J., Danise S., Maxwell E.E., Rondelli R. & Papazzoni C.A. (2023) - Not entirely Ichthyosaur: a mysterious Lamniform and Ichthyopterygian-fall association from the abyssal Upper Cretaceous of the Northern Apennines (Italy). Palaios, 38(8): 331-344.

Serafini G., Danise S., Maxwell E. E., Martire L., Amalfitano J., Cobianchi M., Thun Hohenstein U. & Giusberti L. (2024) - Of his bones are crinoid made: taphonomy and deadfall ecology of marine reptiles from a pelagic setting (Middle-Upper Jurassic of Northeastern Italy). Rivista Italiana di Paleontologia e Stratigrafia, 130(1): 97-128.

Servizio Geologico D’Italia (2009) - Carta Geologica d’Italia alla scala 1:50.000, F. 280 “Fossombrone”.

Servizio Geologico d’Italia (2016) - Carta Geologica d’Italia alla scala 1:50.000, F. 302 “Tolentino”.

Shimada K. (1997a) - Gigantic lamnoid shark vertebra from the Lower Cretaceous Kiowa Shale of Kansas. Journal of Paleontology, 71(3): 522-524.

Shimada K. (1997b) - Skeletal anatomy of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli from the Niobrara Chalk in Kansas. Journal of Vertebrate Paleontology, 17(4): 642-652.

Shimada K. (2005) - Phylogeny of lamniform sharks (Chondrichthyes: Elasmobranchii) and the contribution of dental characters to lamniform systematics. Palaeontological Research, 9(1): 55-72.

Shimada K. (2007) - Skeletal and dental anatomy of lamniform shark, Cretalamna appendiculata, from Upper Cretaceous Niobrara Chalk of Kansas. Journal of Vertebrate Paleontology, 27(3): 584-602.

Shimada K. (2008) - Ontogenetic parameters and life history strategies of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli, based on vertebral growth increments. Journal of Vertebrate Paleontology, 28(1): 21-33.

Shimada K. & Cicimurri D. J. (2005) - Skeletal anatomy of the Late Cretaceous shark, Squalicorax (Neoselachii: Anacoracidae). Palaeontologische Zeitschrift, 79: 241-261.

Shimada K., Chandler R. E., Lam O. L. T., Tanaka T. & Ward D. J. (2017) - A new elusive otodontid shark (Lamniformes: Otodontidae) from the lower Miocene, and comments on the taxonomy of otodontid genera, including the ‘megatoothed’clade. Historical Biology, 29(5): 704-714.

Shimada K., Cumbaa S. L. & Van Rooyen D. (2006) - Caudal fin skeleton of the Late Cretaceous shark, Cretoxyrhina mantelli (Lamniformes: Cretoxyrhinidae) from the Niobrara Chalk of Kansas. Bulletin of the New Mexico Museum of Natural History, 35: 185-192.

Shirai S. (1996) - Phylogenetic interrelationships of neoselachians (Chondrichthyes: Euselachii). Interrelationships of fishes, 2: 9-34.

Siverson M. (1999) - A new large lamniform shark from the uppermost Gearle Siltstone (Cenomanian, Late Cretaceous) of Western Australia. Transactions of the Royal Society of Edinburgh: Earth Sciences, 90: 49-65.

Smart C.W. & Thomas E. (2006) - The enigma of early Miocene biserial planktic foraminifera. Geology, 34: 1041-1044.

Smith C.R. & Baco A.R. (2003) - Ecology of Whale Falls at the Deep-Sea Floor. Oceanography and Marine Biology: an Annual Review, 41: 311-354.

Sorenson L., Santini F. & Alfaro M.E. (2014) - The effect of habitat on modern shark diversification. Journal of Evolutionary Biology, 27(8): 1536-1548.

Stendardi F., Capotorti F., Fabbi S., Ricci V., Silvestri S. & Bigi S. (2020) - Geological mapping of the Mt. Vettoretto - Capodacqua area (Central Apennines, Italy) and cross-cutting relationship between Mt. Sibillini thrust and M. Vettore normal faults system. Geological Fieldtrips and Maps, 12(2.2): 22 pp. https://doi.org/10.3301/GFT.2020.04

Uchida S., Yasuzumi F., Toda M. & Okura N. (1987) - On the observations of reproduction in Carcharodon carcharias and Isurus oxyrinchus. Reports of Japanese Group for Elasmobranch Studies, 24: 5-6.

Underwood C.J. (2006) - Diversification of the Neoselachii (Chondrichthyes) during the Jurassic and cretaceous. Paleobiology, 32(2): 215-235.

Underwood C.J. & Gunter G. (2012) - The shark Carcharhinus sp. from the middle Eocene of Jamaica and the Eocene record of Carcharhinus. Caribbean Journal of Earth Science, 44: 25-30.

Vardabasso S. (1922) - Ittiofauna delle Arenarie Mioceniche di Belluno. Memorie dell’Istituto Geologico della Reale Università di Padova, 6: 23 pp.

Vicens D. & Rodríguez-Perea A. (2003) - Vertebrats fòssils (Pisces i Reptilia) del Burdigalià de cala Sant Vicenç (Pollença, Mallorca). Vertebrate fossils (Pisces and Reptilia) of the Burdigalian deposits from Cala Sant Vicenç (Pollença, Mallorca). Bolletí de la Societat d’Història Natural de les Balears, 46: 117-130.

Von Ihering H. (1927) - Die miocäne Selachierfauna von Schwaben und ihre Beziehungen zu anderen Tertiärfaunen. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilage, 57: 466-503.

Vullo R., Guinot G. & Barbe G. (2016) - The first articulated specimen of the Cretaceous mackerel shark Haimirichia amonensis gen. nov. (Haimirichiidae fam. nov.) reveals a novel ecomorphological adaptation within the Lamniformes (Elasmobranchii). Journal of Systematic Palaeontology, 14(12): 1003-1024.

Wallich G. C. (1877) - Observations on the coccosphere. Annals and Magazine of Natural History, 19: 342-350.

Welton B.J. (2015) - A new species of late early Miocene Cetorhinus (Lamniformes: Cetorhinidae) from the Astoria Formation of Oregon, and coeval Cetorhinus from Washington and California. Natural History Museum of Los Angeles County Contributions in Science, 523: 67-89.

White E.G. (1937) - Interrelationships of the elasmobranchs with a key to the order Galea. Bulletin of the American Museum of Natural History, 74: 25-138.

Whitenack L.B. & Gottfried M.D. (2010) - A morphometric approach for addressing tooth-based species delimitation in fossil mako sharks, Isurus (Elasmobranchii: Lamniformes). Journal of Vertebrate Paleontology, 30(1): 17-25.

Wintner S. P. & Cliff G. (1999) - Age and growth determination of the white shark, Carcharodon carcharias, from the east coast of South Africa. Fisheries Bulletin, 97: 153-169.

Zbyszewski G. (1949) - Les vertébrés du Burdigalien supérieur de Lisbonne. Lissabon: Services géologiques du Portugal, 77 pp.

Downloads

Published

2024-06-24

Issue

Section

Articles
Received 2024-02-22
Accepted 2024-06-05
Published 2024-06-24