First report on swimming trace fossils of fish from the Upper Permian and Lower Triassic of the Dolomites (Italy)

• Autorzy: Ronchi A. , Santi G. , Marchetti L. , Bernardi M. , Gianolla P.

Identyfikatory

DOI

10.14241/asgp.2018.015

• Języki publikacji: EN

Abstrakty

EN

In the Upper Permian continental to marginal-marine succession of the Southern Alps (Dolomites, north Italy), the ichnological record consists of diverse vertebrate footprints and non-diverse invertebrate trace fossils, mainly occurring in the “Bletterbach ichnoassociation” of the Val Gardena Sandstone Formation. After the Permian-Triassic Boundary event, vertebrate ichnoassociations are scarce until the Middle Triassic (Anisian), whereas the uppermost Permian-Lower Triassic Werfen Formation preserves a rich invertebrate trace-fossil record. To date, fish body and trace fossils (Undichna) are very rare in the pre- and post-extinction deposits of the Dolomites; only Undichna gosiutensis Gibert, 2001 was identified in the “Voltago Conglomerate” (Middle Anisian), whereas some unidentified fossil fish casts were found in the Permian Val Gardena Sandstone and some fish remains in the overlying Werfen Formation. Recently, for the first time, fish trails have been discovered in the Val Gardena Sandstone (Lopingian) and in the Werfen Formation (Campil member, Early Triassic, Smithian). Val Gardena Sandstone yielded Undichna cf. quina Trewin, 2000 and U. bina Anderson, 1976 and these represent the oldest fish trails found in the Southern Alps so far. Conversely, the specimens found in the Werfen Formation can be assigned to Undichna cf. britannica Higgs, 1988. They represent the oldest Mesozoic record of fish trace fossils in northern Italy and one of the few records of Undichna from marine environments. These trace fossils are consistent with the fossil association found in the two formations and reflect fish swimming activity in different environments: in very shallow, calm, brackish distal-floodplain to marginal-marine environments in the Late Permian, in association with abundant and diverse tetrapod tracks, and non-diverse invertebrate trace fossils, and in inter- to subtidal calm, shallow, marine environments in the Early Triassic, together with abundant, but not diverse invertebrate trace fossils.

Słowa kluczowe

EN

Trace fossils; Undichna; Permian; Triassic; northern Italy

• Wydawca: Polskie Towarzystwo Geologiczne
• Czasopismo: Annales Societatis Geologorum Poloniae
• Rocznik: 2018
• Tom: Vol. 88, No. 2
• Strony: 111--125
• Opis fizyczny: Bibliogr. 84 poz., rys., wykr.

Twórcy

autor

Ronchi A.

• Dipartimento di Scienze della Terra e dell’Ambiente, University of Pavia, Italy

autor

Santi G.

• Dipartimento di Scienze della Terra e dell’Ambiente, University of Pavia, Italy

autor

Marchetti L.

• Urweltmuseum GEOSKOP/Burg Lichtenberg (Pfalz), Thallichtenberg, Germany

autor

Bernardi M.

• MUSE – Museo delle Scienze, Trento, Italy

autor

Gianolla P.

• Dipartimento di Fisica e Scienze della Terra, University of Ferrara, Italy

Bibliografia

• 1. Accordi, B., 1955. Archaeolopidotus leonardii n. gen., n. sp. e altri pesci Permo-werfeniani delle Dolomiti. Memorie Istituto Geologico Mineralogico Universita di Padova, 19: 1-29. Accordi, B., 1956. A new genus of Lower Triassic holostean fish from the Dolomite Alps. Journal of Paleontology, 30: 345-351.
• 2. Anderson, A. M., 1970. An analysis of supposed fish trails from interglacial sediments in the Dwyka series near Vryheid, Natal. In: Haughton, S. H. (ed.), Proceedings Papers 2nd I.U.G.S. Gondwana Symposium, South Africa, C.S.I.R., Pretoria, pp. 637-647.
• 3. Anderson, A. M., 1976. Fish trails from the Early Permian of South Africa. Palaeontology, 19: 397-409.
• 4. Avanzini, M., Bernardi, M. & Nicosia, U., 2011. The Permo-Triassic tetrapod faunal diversity in the Italian Southern Alps. In: Ahmad Dar, I. & Ahmad Dar, M. (eds), Earth and Environmental Sciences, In Tech, Rijeka, pp. 591-608.
• 5. Avanzini, M. & Mietto, P., 2008. Lower and Middle Triassic footprints-based biochronology in the Italian Southern Alps. Oryctos, 8: 3-13.
• 6. Baucon, A. & De Carvalho, C. N., 2016. Stars of the aftermath: Asteriacites beds from the Lower Triassic of the Carnic Alps (Werfen Formation, Sauris di sopra), Italy. Palaios, 31: 161-176.
• 7. Bauer, K., Kustatscher, E., Butzmann, R., Fischer, T. C., van Konijnenburg-van Cittert, J. H. A. & Krings, M., 2014. Gynkgophytes from the Upper Permian of the Bletterbach Gorge. Rivista Italiana di Paleontologia e Stratigrafia, 120: 271-279.
• 8. Bernardi, M., Klein, H., Petti, F. M. & Ezcurra, M. D., 2015. The origin and early radiation of Archosauriforms: Integrating the skeletal and footprint record. PLoS ONE, 10 (6): e0128449.
• 9. Bernardi, M., Petti, F. M., Kustatscher, E., Franz, M., Hart- kopf-Fröder, C., Labandeira, C. C., Wappler, T., van Konijnenburg-van Cittert, J. H. A., Peecook, B. R. & Angielczyk, K. D., 2017. Late Permian (Lopingian) terrestrial ecosystems: A global comparison with new data from the low latitude Bletterbach Biota. Earth-Science Reviews, 175: 18-43.
• 10. Blieck, A., Conti, M. A., Dalla Vecchia, F. M., Fluegel, H. W., Gand, G., Hubmann, B., Lelievre, H., Mariotti, N., Nicosia, U., Poplin, C., Schneider, J. W. & Werneburg, R., 1997. Palaeozoic vertebrates of the Alps: a review. Bulletin de la Societé Géologique de France, 168: 343-350.
• 11. Bordy, E. M., Linkermann, S. & Prevec, R., 2011. Palaeoecological aspects of some invertebrates trace fossils from the midto Upper Permian Middleton Formation (Adelaide Subgroup, Beaufort Group, Karoo Supergroup), Eastern Cape, South Africa. Journal of African Earth Sciences, 61: 238-244.
• 12. Brandner, R., Horacek, M., Keim, L. & Scholger, R., 2009. The Pufels/Bulla road section: deciphering environmental changes across the Permian-Triassic boundary to the Olenekian by integrated litho-, magneto- and isotope stratigraphy. A field trip guide. GeoAlp, 6 S: 116-132.
• 13. Broglio Loriga, C., Goczan, F., Haas, J., Lenner, K., Neri, C., Oravecz Sheffer, A., Posenato, R., Szabo, I. & Toth Mak, A., 1990. The Lower Triassic sequences of the Dolomites (Italy) and Transdanubian Mid-Mountains (Hungary) and their correlation. Memorie Scienze Geologiche Universita Padova, 42: 41-103.
• 14. Broglio Loriga, C., Masetti, D. & Neri, C., 1983. La Formazione di Werfen (Scitico) delle Dolomiti occidentali. Rivista Italiana di Paleontologia e Stratigrafia, 88: 501-598.
• 15. Buatois, L. & Mángano, G., 1993. Trace fossils from a Carboniferous turbiditic lake: implications for the recognition of additional nonmarine ichnofacies. Ichnos, 2: 237-258.
• 16. Buatois, L. & Mángano, G., 1994. Pistas de peces en el Carbonifero de la Cuenca Paganzo: Su significado estratigráfico y paleoambiental. Ameghiniana, Revista de la Associación Paleontologica Argentina, 31: 33-40.
• 17. Buatois, L. & Mángano, G., 2011. Ichnology: Organism-Substrate Interactions in Space and Time. Cambridge University Press, Cambridge, 358 pp.
• 18. Buatois, L. A., Netto, R. G. & Mángano, G., 2010. Ichnology of late Paleozoic postglacial transgressive deposits in Gondwana: Reconstructing salinity conditions in coastal ecosystem affected by strong meltwater discharge. Geological Society of America, Special Paper, 468: 149-173.
• 19. Cameron, B., Wood, D. & Van Dommelen, R., 1998. Vertebrate and invertebrate trace fossils from the Horton Bluff Formation (Lower Carboniferous) of Nova Scotia. Mining Matters for Nova Scotia, 98: 6.
• 20. Cardonatto, M. C. & Melchor, R. N., 2014. Biomechanical analysis of fish swimming trace fossils (Undichna): preservation and mode of locomotion. In: Hembree, D. I., Platt, B. F. & Smith, J. J. (eds), Experimental approaches to understanding fossil organism. Topics in Geobiology, 41. Springer Science+Business Media, Dordreicht, pp. 265-303.
• 21. Cassinis, G., Cortesogno, L., Gaggero, L., Massari, F., Neri, C., Nicosia, U. & Pittau, P. (eds), 1999. Stratigraphy and facies of the Permian deposits between Eastern Lombardy and the Western Dolomites. Field Trip Guidebook. International Field Conference on "The Continental Permian of the Southern Alps and Sardinia (Italy). Regional Reports and General Correlations”. Earth Science Department, Pavia University, Italy, 157 pp.
• 22. Cassinis, G., Massari, F., Neri, C. & Venturini, C., 1988. The continental Permian in the Southern Alps (Italy). A review. Zeitschrift fur Geologische Wissenschaften, 16: 1117-1126.
• 23. Cassinis, G., Perotti, C. R. & Ronchi, A., 2012. Permian continental basins in the Southern Alps (Italy) and peri-Mediterranean correlations. International Journal of Earth Sciences, 101: 129-157.
• 24. Ceoloni, P., Conti, M. A., Mariotti, N. & Nicosia, U., 1986. New Late Permian tetrapod footprints from Southern Alps. Memorie della Societa Geologica Italiana, 34: 45-65.
• 25. Citton, P., Nicolosi, I., Carluccio, R. & Nicosia, U., 2016. Unveiling trampling history through trackway interferences and track preservational features: a case study from the Bletterbach gorge (Redagno, Western Dolomites, Italy). Palaeontologia Electronica, 19.2.20A: 1-20
• 26. Conti, M. A., Leonardi, G., Mariotti, N. & Nicosia, U., 1975. Tetrapod footprints, fishes and molluscs from the Middle Permian of the Dolomites (N. Italy). Memorie Geopaleontologiche, Universita di Ferrara, 3: 141-150.
• 27. Conti, M. A., Leonardi, G., Mariotti, N. & Nicosia, U., 1977. Tetrapod footprints of the “Val Gardena Sandstone” (North Italy). Their paleontological, stratigraphic and paleoenvironmental meaning. Palaeontographia Italica, 70, Nuova Serie, 40: 1-91.
• 28. Conti, M. A., Leonardi, G., Mietto, P. & Nicosia, U., 2000. Orme di tetrapodi non dinosauriani del Paleozoico e Mesozoico in Italia. In: Leonardi, G. & Mietto, P. (eds), Dinosauri in Italia. Le orme giurassiche dei Lavini di Marco (Trentino) e gli altri resti fossili italiani. Accademia Editoriale, Pisa, pp. 297-320.
• 29. Conti, M. A., Mariotti, N., Nicosia, U. & Pittau, P., 1997. Selected bioevents succession in the continental Permian of the Southern Alps (Italy): improvements of intrabasinal and interregional correlations. In: Dickins, M., Zunyi, Y., Yhongfu, Y., Lucas, S. G. & Acharyya, S. J. J. (eds), Late Palaeozoic and Early Mesozoic Circum-Pacific Events and Their Global Correlation. Cambridge University Press, Cambridge, pp. 51-65.
• 30. Costeur, L. & Ezquerra, R., 2009. Early Cretaceous fish trails from La Roja, Spain. Cretaceous Research, 30: 1027-1030.
• 31. Dalla Vecchia, F. M., 1988. First record of a petalodont (Petalodus ohioensis Safford, 1853) from the Alps. Gortania-Atti del Museo Friulano di Storia Naturale, 9: 47-56.
• 32. Feng, X., Chen, Z.-Q., Benton, M. J., Wu, S., Bottjer, D. J. & Thompson, J. R., 2017. A diverse trackway-dominated marine ichnoassemblage from the Lower Triassic in the northern Paleotethys: Ichnology and implications for biotic recovery. Palaeogeography, Palaeoclimatology, Palaeoecology, https://doi.org/10.1016/j.palaeo.2017.11.059.
• 33. Fillmore, D. L., Lucas, S. G. & Simpson, E. L., 2011. The fish swimming trace Undichna from the Mississippian Mauch Chunk Formation, Eastern Pennsylvania. Ichnos, 18: 27-34.
• 34. Fliri, F., Bortenschlager, S., Felber, H., Heissel, W. & Resch, W. 1970. Der Bänderton von Baumkirchen (Inntal, Tirol) eine neue Schlüsselstelle zur Kenntnis der Würm-vereisung der Alpen. Zeitschrift für Gletscherkunde und Glazialgeographie, 6: 5-35.
• 35. Gianolla, P., De Zanche, V. & Mietto, P., 1998. Triassic sequence stratigraphy in the Southern Alps (Northern Italy): definition of sequences and basin evolution. In: De Graciansky, P.-C., Hardenbol, J., Jacquin, T. & Vail, P. R. (eds), Mesozoic and Cenozoic Sequence Stratigraphy of European Basins. SEPM Special Publications, 60: 719-747.
• 36. Gianolla, P. & Jacquin, T., 1998. Triassic sequence stratigraphic framework ofwestern European basins. In: De Graciansky, P. C., Hardenbol, J., Jacquin, T. & Vail, P. R. (eds), Mesozoic and Cenozoic Sequence Stratigraphy of European Basins. SEPM Special Publication, 60: 643-650.
• 37. Gibert, J. M., 2001. Undichna gosiutensis, isp. nov.: a new fish trace fossil from the Jurassic of Utah. Ichnos, 8: 15-22.
• 38. Gibert, J. M., Buatois, L. A., Fregenal-Martinez, M. A., Mánga- no, M. G., Ortega, F., Poyato-Ariza, F. J. & Wenz, S., 1999. The fish trace fossil Undichna from the Cretaceous of Spain. Palaeontology, 42: 409-427.
• 39. Italian IGCP 203 Group (eds), 1986. Field Guide-Book of Permian and Permian-Triassic Boundary in the South-Alpine Segment of the Western Tethys. Project n. 203 SGI Brescia, July 1986. Tipolitografica Commerciale Pavese, 180 pp.
• 40. Higgs, R., 1988. Fish trails in the Upper Carboniferous of SouthWest England. Palaeontology, 31: 255-272.
• 41. Hofmann, R., Goudemand, N., Wasmer, M., Bucher, H., & Hautmann, M., 2011. New trace fossil evidence for an early recovery signal in the aftermath of the end-Permian mass extinction. Palaeogeography, Palaeoclimatology, Palaeoecology, 310: 216-226.
• 42. Hofmann, R., Hautmann, M. & Bucher, H., 2015. Recovery dynamics of benthic marine communities from the Lower Triassic Werfen Formation, northern Italy. Lethaia, 48: 474-496.
• 43. Knaust, D., 2013. The ichnogenus Rhizocorallium: Classification, trace makers, palaeoenvironments and evolution. Earth-Science Reviews, 126: 1-47.
• 44. Kustatscher, E., Bauer, K., Butzmann, R., Fischer, T. C., Meller, B., van Konijnenburg-van Cittert, J. H. A. & Kerp, H., 2014. Sphenophytes, pteridosperms and possible cycads from the Wuchiapingian (Lopingian, Permian) of Bletterbach (Dolomites, Northern Italy). Review of Palaeobotany and Palynology, 208: 65-82.
• 45. Kustatscher, E., van Konijnenburg-van Cittert, J. H. A., Bauer, K., Butzmann, R., Meller, B. & Fischer, T. C., 2012. A new flora from the Upper Permian of Bletterbach (Dolomites, N-Italy). Review of Palaeobotany and Palynology, 182: 1-13.
• 46. Kustatscher, E., Bernardi, M., Petti, F. M., Franz, M., van Konijnenburg-van Cittert, J. & Kerp, H., 2017. Sea-level changes in the Lopingian (late Permian) of the northwestern Tethys and their effects on the terrestrial palaeoenvironments, biota and fossil preservation. Global and Planetary Change, 148: 166-180.
• 47. Labandeira, C. C., Kustatscher, E. & Wappler, T., 2016. Floral Assemblages and Patterns of Insect Herbivory during the Permian to Triassic of Northeastern Italy. PLoS ONE 11(11): e0165205.
• 48. Leonardi, G., 1987. Glossary and Manual of Tetrapod Footprint Palaeoichnology. Departamento Nacional Produçáo Mineral, Brasilia, 117 pp.
• 49. Leonardi, P., 1951. Orme di tetrapodi nelle Arenarie di Val Gardena (Permiano medio-inferiore) dell’Alto Adige Sud-orientale. Memorie Istituto di Geologia Universita di Padova, 16: 1-15.
• 50. Leonardi, P., 1960. Campagne geo-paleontologiche 1957-1959 nella regione Dolomitica e nei colli Berici. La Ricerca Scientifica, 30: 2062-2093.
• 51. Leonardi, P., 1967. Werfeniano (Trias inferiore). In: Leonardi, P. (ed.), Le Dolomiti: geologia dei monti tra l’Isarco e il Piave, 1: 107-123.
• 52. Leonardi, P., Conti, M. A., Leonardi, G., Mariotti, N. & Nicosia, U., 1975. Pachypes dolomiticus n. gen. n. sp.; Pareiasaur footprint from the “Val Gardena Sandstone” (Middle Permian) in the western Dolomites (N. Italy). Atti Accademia Nazionale dei Lincei Rendiconti Classe Scienze Fisiche, Matematiche, Naturali, Serie 8, 57: 221-232.
• 53. Lucas, S. G., Lerner, A. J., Bruner, M. & Shipman, P., 2004. Middle Pennsylvanian ichnofauna from Eastern Oklahoma, USA. Ichnos, 11: 45-55.
• 54. Lu, Z., Hou, J., Chen, B., Hao, C. & Li, X., 2004. Genetic interpretation of fish swimming trails and calculation of fish body length in the Late Triassic Hengsham, Shaanxi Province, China. Science in China, Serie D, Earth Sciences, 47: 272-279.
• 55. Lu, T., Wang, Z., Yang, X. & Zhang, X., 2012. First record of Lower Triassic Undichna spp. fish swimming traces from Emei, Sichuan Province, China. Chinese Science Bulletin, 57: 1320-1324.
• 56. Marchetti, L., Belvedere, M. & Mietto, P., 2017a. Lopingian tetrapod footprints from the Venetian Prealps, Italy: New discoveries in a largely incomplete panorama. Acta Palaeontologica Polonica, 62: 801-817.
• 57. Marchetti, L., Mujal, E., & Bernardi, M., 2017b. An unusual Amphisauropus trackway and its implication for understanding seymouriamorph locomotion. Lethaia, 50: 162-174.
• 58. Marchetti, L., Voigt, S. & Klein, H., 2017c. Revision of Late Permian tetrapod tracks from the Dolomites (Trentino-Alto Adige, Italy). Historical Biology, doi: 10.1080/08912963.2017.1391806.
• 59. Massari, F., Conti., M. A., Fontana, D., Helmold, K., Mariotti, N., Neri, C., Nicosia, U., Ori, G. G., Pasini, M. & Pittau, P., 1988. The Val Gardena Sandstone and the Bellerophon Formation in the Bletterbach gorge (Alto Adige, Italy): biostratigraphy and sedimentology. Memorie Scienze Geologiche Universita Padova, 40: 229-273.
• 60. Massari, F. & Neri, C., 1997. The infill of a supradetachment (?) basin: the continental to shallow-marine Upper Permian succession of Dolomites and Carnia (Italy). Sedimentary Geology, 110: 181-221.
• 61. Massari, F., Neri, C., Pittau, P., Fontana, D. & Stefani, C., 1994. Sedimentology, palynostratigraphy and sequence stratigraphy of a continental to shallow-marine riftrelated succession: Upper Permian of the eastern Southern Alps (Italy). Memorie Scienze Geologiche Universita Padova, 46: 119-243.
• 62. Mietto, P., 1986. Orme di tetrapodi nella Formazione di Werfen del Recoarese. Rivista Italiana di Paleontologia e Stratigrafia, 92: 321-326.
• 63. Minter, N. J. & Braddy, S. J., 2006. The fish and amphibian swimming traces Undichna and Lunichnium, with examples from the Lower Permian of New Mexico, USA. Palaeontology, 49: 1123-1142.
• 64. Morrissey, L. B., Braddy, S. J., Bennett, J. P., Marriott, S. B. & Tarrant, P. R., 2004. Fish trails from the Lower Old Red Sandstone of Tredomen Quarry Powys, Southeast Wales. Geological Journal, 39: 337-358.
• 65. Neri, C., 2007. Formazione di Werfen: In: Cita, M. B., Abbate, E., Aldighieri, B., Balini, M., Conti, M. A., Falorni, P., Germani, D., Groppelli, G., Manetti, P. & Petti, F. M. (eds), Carta Geologica d’Italia -1:50.000, Catalogo delle Formazioni, Unita tradizionali (1). Quaderni serie III, 7, Fascicolo VI. APAT, Dipartimento Difesa del Suolo, Servizio Geologico d’Italia. Roma, pp. 83-86.
• 66. Neri, C., Gianolla, P., Furlanis, S., Caputo, R. & Bosellini, A., 2007. Note Illustrative della Carta Geologica d’Italia alla scala 1:50.000, Foglio 029 Cortina d’Ampezzo. Roma: Systemcart, pp. 1-200.
• 67. Porada, H. & Bouougri, E., 2007. Wrinkle structures - a critical review. Earth Science Review, 81: 199-215.
• 68. Richthofen, F., von, 1860. Geognostische Beschreibung der Umgegend von Predazzo, S. Cassian und der Seisser Alpe in Südtirol. Perthes, Gotha, 327 pp.
• 69. Seilacher, A., 1967. Bathymetry of trace fossils. Marine Geology, 5: 413-428.
• 70. Sirna, G., Dalla Vecchia, F. M., Muscio, G. & Piccoli, G., 1994. Catalogue of Paleozoic and Mesozoic vertebrates and vertebrate localities of the Tre Venezie area (North Eastern Italy). Memorie di Scienze Geologiche di Padova, 46: 255-281.
• 71. Soler-Gijón, R. & Moratalla, J. J., 2001. Fish and tetrapod trace fossils from the Upper Carboniferous of Puertollano, Spain. Palaeogeography, Palaeoclimatology, Palaeoecology, 171: 1-28.
• 72. Stefani, M., Furin, S. & Gianolla, P., 2010. The changing climate framework and depositional dynamics of Triassic carbonate platforms from the Dolomites, Palaeogeography, Palaeoclimatology, Palaeoecology, 290: 43-57.
• 73. Todesco, R. & Avanzini, M., 2008. First record of the fish trace fossil Undichna from the Middle Triassic of Italy. Studi Trentini di Scienze Naturali, Acta Geologica, 83: 253-257.
• 74. Trewin, N. H., 2000.The ichnogenus Undichna, with examples from the Permian of the Falkland Islands. Palaeontology, 43: 979-997.
• 75. Turek, V., 1989. Fish and amphibian trace fossils from Westphalian sediments of Bohemia. Palaeontology, 32: 623-643.
• 76. Twitchett, R. J. A., 2000. High resolution biostratigraphy for the Lower Triassic of northern Italy, Palaeontological Association Newsletters, 43: 19-22.
• 77. Twitchett, R. J. & Wignall, P. B., 1996. Trace fossils and the aftermath of the Permo-Triassic mass extinction: evidence from northern Italy. Palaeogeography, Palaeoclimatology, Palaeoecology, 124: 137-151.
• 78. Valentini, M., Conti, M. A. & Mariotti, N., 2007. Lacertoid footprints of the Upper Permian Arenaria di Val Gardena formation (Northern Italy). Ichnos, 14: 193-218.
• 79. Valentini, M., Nicosia, U. & Conti, M. A. 2009. A re-evaluation of Pachypes, a pareiasaurian track from the Late Permian. Neues Jahrbuch fur Geologie und Paläontologie, 251: 71-94.
• 80. Venturini, C., 1990. Geologia delle Alpi Carniche centro-orientali. Edizione Museo Friulano di Storia Naturale, Comune di Udine, 36, 217 pp.
• 81. Voigt, S., Berman, D. S., & Henrici, A. C., 2007. First well-established track-trackmaker association of Paleozoic tetrapods based on Ichniotherium trackways and diadectid skeletons from the Lower Permian of Germany. Journal of Vertebrate Paleontology, 27: 553-570.
• 82. Wisshak, K., Volohonsky, E. & Blomeier, D., 2004. Acanthodian fish trace fossils from the Early Devonian of Spitsbergen. Acta Palaeontologica Polonica, 49: 629-634.
• 83. Zanche, V., Gianolla, P., Mietto, P., Siorpaes, C. & Vail, P., 1993. Triassic sequence stratigraphy in the Dolomites (Italy). Memorie di Scienze Geologiche, 45: 1-27
• 84. Zonneveld, J.-P., 2011. Suspending the rules: unraveling the ichnological signature of the Lower Triassic post-extinction recovery interval. Palaios, 26: 677-681.

Uwagi

Opracowanie rekordu w ramach umowy 509/P-DUN/2018 ze środków MNiSW przeznaczonych na działalność upowszechniającą naukę (2019).