Description and classification of Late Triassic Neritimorpha (Gastropoda, Mollusca) from the St Cassian Formation, Italian Alps


Authors: Bandel K

Published in: Bulletin of Geosciences, volume 82, issue 3; pages: 215 - 274; Received 27 April 2007; Accepted in revised form 30 July 2007;

Keywords: Gastropoda, Neritimorpha, Triassic, Tethys, Italy, Jordan, new taxa (fossil, living),

full text (PDF, 2.05 MB)

Export to RIS



Neritopsis represents the only surviving genus of an ancient group of the Neritimorpha that has no internal dissolution of its shell walls. While the two known living species have lecithotrophic early development without larval shell, Neritopsis aqabaensis n. sp. from the Gulf of Aqaba, Jordan, has a planktotrophic veliger. These living species of the genus differ in their protoconch shape as well as teleoconch morphology and ornament from the Triassic species that can be considered related to Neritopsis. Neritopsidae with the modern Neritopsinae is distinguished from the Triassic Cassianopsinae n. subfam. based on the genus Cassianopsis n. gen. with three species by features of their protoconch as well as the different characters of the operculum. Zardiniopsis n. gen. differs from these by higher shell shape and a smaller more complexly ornamented protoconch. Fossariopsis has a more angular shell shape. Colubrellopsinae n. subfam. with Colubrellopsis n. gen. resembles Cassianopsis n. gen. in respect to protoconch and features of the aperture of the teleoconch, but the former has rounded whorls and an ornament of axial ribs. Among the Fedaiellidae n. fam. with smooth shells two species of Fedaiella are redefined. The characters of the inner lip of their aperture connect them with the Neritopsidae, whereas the operculum in the Fedaiellidae with concentric structure on the outside distinguishes them from neritopsids.
In distinction to the groups of the Neritopsoidea members of the Dephinulopsidae have a smooth inner lip of the aperture. Here Delphinulopsinae and Platychilininae n. subfam. differ from each other in the shape and ornament of their teleoconch. Schwardtopsis n. gen. resembles a juvenile Delphinulopsis grown to a larger size. The large concave callus of the inner lip and almost open coiling defines Delphinulopsis. Rows of nodes, low initial shell and rapid growth in shell diameter to an almost limpet shape characterises Platychilina, and lamellar growth increments on an almost limpet-like shell with flat initial part is present in Marmolatella. The Palaeonaricidae n. fam. contains two species of Palaeonarica, which have a Nerita-like shell with simple aperture.
The Naticopsidae of the Carboniferous and Permian has its continuation in the Ampezzonaticopsinae n. subfam. of the Triassic, being connected to each other by the sinuous ornament of ribs on their larval shells. The genera are distinguished by teleoconch shape and differences in the ornament of their larval shell. Ampezzonaticopsis n. gen. has whorls well separated by deep sutures, Cortinaticopsis n. gen. has a simple aperture and a callus covering the umbilicus. The Hologyrinae n. subfam. with Hologyra have a ridge in the columellar furrow and an operculum that resembles that attributed to Carboniferous Naticopsis. Their protoconch has a characteristic chevron ornament on its larval whorls.


Bandel, K. 1982. Morphologie und Bildung der frühontogenetischen Gehäuse bei conchiferen Mollusken. Facies 7, 1–198.View article

Bandel, K. 1988. Repräsentieren die Euomphaloidea eine natürliche Einheit der Gastropoden? Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 67, 1–33.

Bandel, K. 1990. Shell structure of the Gastropoda excluding the Archaeogastropoda, 117–134. In Carter, J.G. (ed.) Skeletal Biomineralization: Patterns Processes and Evolutionary Trends. Volume I. Van Nostrand Reinolds, New York.

Bandel, K. 1993. Trochomorpha aus der triassischen St. Cassian Formation (Gastropoda, Dolomiten). Annales des Naturhistorischen Museums in Wien 95, 1–99.

Bandel, K. 1994. Comparison of Upper Triassic and Lower Jurassic gastropods from the Peruvian Andes (Pucará Group) and the Alps (Cassian Formation). Palaeontographica 233A, 127–160.

Bandel, K. 1997. Higher classification and pattern of evolution of the Gastropoda. A synthesis of biological and paleontological data. Courier Forschungsinstitut Senckenberg 201, 57–81.

Bandel, K. 2000. The new family Cortinellidae (Gastropoda, Mollusca) connected to a review of the evolutionary history of the subclass Neritimorpha. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 217(1), 111–129.View article

Bandel, K. 2001. The history of Theodoxus and Neritina connected with description and systematic evaluation of related Neritimorpha (Gastropoda). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 85, 65–164.

Bandel, K. 2002. Reevaluation and classification of Carboniferous and Permian Gastropoda belonging to the Caenogastropoda and their relation. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 86, 81–188.

Bandel, K. & Frýda, J. 1999. Notes on the evolution and higher classification of the subclass Neritimorpha (Gastropoda) with the description of some new taxa. Geologica et Palaeontologica 33, 219–235.

Bandel, K. & Heidelberger, E. 2001. The new family Nerrhenidae (Neritimorpha, Gastropoda) from the Givetian of Germany. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2001(12), 705–718.View article

Bandel, K. & Kiel, S. 2003. Relationships of Cretaceous Neritimorpha (Gastropoda, Mollusca), with the description of seven new species. Bulletin of the Czech Geological Survey 7, 53–65.

Biggelaar, J.A.M. van den & Haszprunar, G. 1996. Cleavage patterns and mesentoblast formation in the Gastropoda: an evolutionary perspective. Evolution 50, 1520–1540.View article

Blaschke, F. 1905. Die Gastropodenfauna der Pachycardientuffe der Seiseralpe in Südtirol nebst einem Nachtrag zur Gastropodenfauna der roten Raibler Schichten vom Schlernplateau. Beiträge zur Paläontologie und Geologie Österraich-Ungarns und des Orients 17, 161–222.

Blendinger, W. & Blendinger, E. 1989. Windward-leeward effects on Triassic carbonate bank margins in the Dolomites, northern Italy. Sedimentary Geology 64, 143–166.View article

Blodgett, R.B., Frýda, J. & Stanley, G.D. 2001. Dephinulopsidae, a new neritopsoidean family from the Upper Triassic (upper Carnian to lower Norian) of the Wallowa terrane, northeastern Oregon. Journal of Czech Geological Society 46, 307–318.

Böhm, J. 1895. Die Gastropoden des Marmolatakalkes. Palaeontographica 42, 211–308.

Boggild, O.B. 1930. The shell structure of the molluscs. Det Kongelige Danske Videnskabernes Selkabs Skrifter, Naturvidenskabelig og Mathematisk Afdeling 9, 231–326.

Bouchet, P. & Rocroi, J.-P. 2005. Classification and nomenclator of gastropod families. Malacologia 47, 1–397.

Bourne, G.C. 1908. Contributions to the morphology of the group Neritacea of aspidobranch gastropods. Part I. The Neritidae. Proceedings of the Zoological Society of London 1908, 810–887.View article

Brösamlen, R. 1909. Beitrag zur Kenntnis der Gastropoden des schwäbischen Jura. Palaeontographica 56, 177–321.

Broili, F. 1907. Die Fauna der Pachycardientuffe der Seiser Alp. Scaphopoden und Gastropoden. Palaeontographica 54, 69–138.

Cossmann, M. 1898. Referat Koken: Die Gastropoden der Trias von Hallstadt. Revue critique de paléozoologie 2, 148–150.

Cossmann, M. 1925. Essais de Paléontologie Comparée. Treizieme Livraison. 345 pp. Les Presses Universitaires de France, Paris.

Diener, C. 1926. Fossilium Catalogus I: Animalia, Pars 34 – Glossophora triadica. 242 pp. W. Junk, Berlin.

Fretter, V. & Graham, A. 1962. British prosobranch molluscs, their functional anatomy and ecology. 755 pp. Ray Society, London.

Frýda, J. 2000. Some Givetian (late Middle Devonian) gastropods from the Paffrath area (Bergisches Land, Germany). Memoirs of the Queensland Museum 45, 359–374.

Frýda, J. & Bandel, K. 1997. New Early Devonian gastropods from the Plectonotus (Boucotonotus)Palaeozygopleura Community in the Prague Basin (Bohemia). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 80, 1–57.

Frýda, J., Blodgett, R.B. & Stanley, G.D. 2003. New neritopsoidean gastropods (Neritimorpha) from the Late Triassic (Late Carnian–Early Norian) of the Wallowa Terrane, northeastern Oregon. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 87, 55–72.

Fürsich, F.T. & Wendt, J. 1977. Biostratinomy and palaeoecology of the Cassian Formation (Triassic) of the southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology 22, 257–323.View article

Golikov, A.N. & Starobogatov, Y.I. 1975. Systematics of prosobranch gastropods. Malacologia 15(1), 185–232.

Gordon, M. & Yochelson, E.L. 1982. A Naticopsis operculum found in situ (Gastropoda: Mississippian). Journal of Paleontology 56, 260–265.

Gründel, J. 1980. Zur Gastropodenfauna des Unteren Muschelkalkes im Raume Halle-Unstruttal (DDR). Freiberger Forschungshefte C348, 41–59.

Haas, O. 1953. The Late Triassic gastropods from Central Peru. Bulletin of the American Museum of Natural History 101, 9–328.

Haszprunar, G. 1988. On the origin and evolution of the major gastropod groups, with special reference to the Streptoneura. Journal of Molluscan Studies 54, 367–441.View article

Heidelberger, D. 2001. Mitteldevonische (Givetische) Gastropoden (Mollusca) aus der Lahnmulde (südliches Rheinisches Schiefergebirge). Geologische Abhandlungen Hessen 106, 1–291.

Heidelberger, D. & Bandel, K. 1999. Micromorph Gastropoda from the Middle Devonian (Givetian) limestone of the Soetenich Syncline (Eifel). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 83, 129–162.

Herholz, M. 1992. Mikromorphe Gastropoden aus dem rheinisch-westfälischen Steinkohlerevier (Oberkarbon). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1992, 242–256.View article

Holthuis, B.V. 1995. Evolution between marine and freshwater habitats: a case study of the gastropod Neritopsina. 286 pp. Doctoral dissertation, University of Washington.

Jagt, J.W.M. & Janssen, A.W. 1988. Jouannetia (Bivalvia, Pholadidae) and Neritopsis (Gastropoda, Neritopsidae), two mollusks from the Danian (Paleocene) of the Maastricht area (SE Netherlands and NE Belgium). Mededelinden Werkgroep voor Tertiare en Kwartaire Geologie 25, 163–174.

Jeffery, D.L., Hoare, R.D.,Mapes, R.H. & Brown, C.L. 1994. Gastropods (Mollusca) from the Imo Formation (Mississippian, Chesterian) of north-central Arkansas. Journal of Paleontology 68, 58–79.View article

Kaim, A. & Sztajner, P. 2005. The opercula of neritopsid gastropods and their phylogenetic importance. Journal of Molluscan Studies 71, 211–219.View article

Kano, Y., Chiba, S. & Kase, T. 2002. Major adaptive radiation in neritopsine gastropods estimated from 28S rRNA sequences and fossile records. Proceeding of the Royal Society 2699, 2457–2465.View article

Kano, Y. & Kase, T. 2000. Taxonomic revision of Pisulina (Gastropoda: Neritopsina) from submarine caves in the tropical Indo-Pacific. Paleontological Research 4, 107–129.

Kittl, E. 1892. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. Part 2. Annalen des kaiserlich-königlichen naturhistorischen Hofmuseums 7, 35–97.

Kittl, E. 1894. Die triadischen Gastropoden der Marmolata und verwandter Fundstellen in den weissen Riffkalken Südtirols. Jahrbuch der Kaiserlich-königlichen geologischen Reichsanstalt 44 (1894), 99–182.

Kittl, E. 1899. Die Gastropoden der Esinokalke, nebst einer Revision der Gastropoden der Marmolatakalke. Annalen des kaiserlich-königlichen naturhistorischen Hofmuseums 14, 1–237.

Kittl, E. 1901. Bakonyi Triasz-Gastropodák. 54 pp. Budapest.

Klipstein, A.V. 1843. Beiträge zur geologischen Kenntnis der östlichen Alpen. 311 pp. Georg Friedrich Geyer, Giessen.

Knight, J.B. 1941. Paleozoic gastropod genotypes. Bulletin of the Geological Society of America, Special Paper 32, 1–510.

Knight, J.B., Cox, L.R., Keen, A.M., Batten, R.L., Yochelson, E.L. & Robertson, R. 1960. Part I, Mollusca. Systematic descriptions, I169–I351. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology. University of Kansas Press, Lawrence.

Koninck, L. de 1853. Notes sur le genre Hypodema. Mémoires de la Societé Royale des Sciences de Liége 8, 140–144.

Koken, E. 1889. Über die Entwicklung der Gastropoden vom Cambrium bis zur Trias. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, suppl. vol. 6, 305–484.

Koken, E. 1892. Über die Gastropoden der rothen Schlernschichten nebst Bemerkungen über Verbreitung und Herkunft einige triassischer Gattungen. Neues Jahrbuch für Mineralogie 1892(2), 25–36.

Koken, E. 1897. Die Gastropoden der Trias von Hallstatt. Abhandlungen der Kaiserlich- königlichen geologischen Reichsanstalt in Wien 17(4), 1–112.

Kues, B.S. & Batten, R.L. 2001. Middle Pennsylvanian gastropods from the Flechado Formation, North-Central New Mexico. Journal of Paleontology, Memoir 54, Supplement 1(75), 1–95.View article

Laube, G.C. 1869. Die Fauna der Schichten von St. Cassian. Kaiserliche Akademie der Wissenschaften, Denkschrift 30, 1–48.

Leonardi, P. & Fiscon, F. 1959. La fauna Cassiana di Cortina d’Ampezzo. 3. Gasteropodi. Memorie degli Istituti di Geologia e Mineralogia dell’Universita di Padova 21, 1–103.

Mojsisovics, E. von 1879. Die Dolomit-Riffe von Südtirol und Venetien. Beiträge zur Bildungsgeschichte der Alpen. 552 pp. Hölder, Wien.

Münster, G.G. von 1841. Beschreibung und Abbildung der in den Kalkmergelschichten von St. Cassian gefundenen Versteinerungen, 25–152. In Wissmann, H.L. & Münster, G.G. Beiträge zur Geologie und Petrefacten-Kunde des südöstlichen Tirol’s vorzüglich der Schichten von St. Cassian 4. Buchner, Bayreuth.

Nützel, A. & Erwin, D.H. 2004. Late Triassic (Late Norian) gastropods from the Wallowa Terrane (Idaho, USA). Paläontologische Zeitschrift 78, 361–416.View article

Nützel, A. & Senowbari-Daryan, B. 1999. Gastropods from the Later Triassic (Norian-Rhaetian) Nayband Formation of central Iran. Beringeria 23, 93–132.

Ponder, W.F. & Lindberg, D. 1997. Towards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society 119, 83–265.View article

Robertson, R. 1971. Scanning electron microscopy of planktonic larval marine gastropod shells. The Veliger 14(1), 1–12.

Sasaki, T. 1998. Comparative anatomy and phylogeny of the Recent Archaeogastropoda (Mollusca: Gastropoda). The University Museum, The University of Tokyo, Bulletin 38, 1–223.

Sasaki, T. 2001. Macro- and microstructure of shell and operculum in two recent gastropod species, Nerita (Theliostyla) albicilla and Cinnalepeta pulchella (Neritopsina Neritoidea). Palaeontological Research 5, 21–31.

Scott, B.J. & Kenny, R. 1998. Superfamily Neritoidea, 694–702. In Beesley, P.L., Ross, G.J.B. & Wells, A. (eds) Mollusca, The Southern Synthesis. Fauna of Australia 5, Part B. Commonwealth Scientific and Industrial Research Organization, Canberra.

Suzuki, S., Togo, Y. & Uozumi, S. 1991. Exosceletal adaptations for life environment in neritacean gastropods, 397–401. In Suga, S. & Nakahara, H. (eds) Biological Systems. Springer Verlag, Tokyo.

Thiele, J. 1929–1931. Handbuch der systematischen Weichtierkunde. Gustav Fischer Verlag, Jena.

Tichy, G. 1980. Über die Erhaltung von Farben und Farbmustern an traissischen Gastropoden-Gehäusen (Colour retention in Triassic gastropods). Verhandlungen der Geologischen Bundesanstalt A 1980(2), 175–217.

Thompson, F.G. 1980. Proserpinoid land snails and their relationships within the Archaeogastropoda. Malacologia 20, 1–33.

Troschel, F.H. 1856. Das Gebiss der Schnecken zur Begründung einer natürlichen Classification 1. 252 pp. Nicolaische Verlagsbuchhandlung, Berlin.

Troschel, F.H. & Thiele, J. 1865–1893. Das Gebiss der Schnecken zur Begründung einer natürlichen Classification 2. 409 pp. Nicolaische Verlagsbuchhandlung, Berlin.

Urlichs, M. 1974. Zur Stratigraphie und Ammonitenfauna der Cassianer Schichten von Cassian (Dolomiten/ Italien). Schriftenreihe der Erdwissenschaftlicher Kommission 2, 207–222.View article

Urlichs, M. 1994. Trachyceras Laube, 1869 (Ammonoidea) aus dem Unterkarn (Obertrias) der Dolomiten (Italien). Stuttgarter Beiträge zur Naturkunde, Serie B 217, 1–55.

Wendt, J. 1982. The Cassian patch reefs (Lower Carnian, Southern Alps). Facies 6, 185–202.View article

Wenz, W. 1938–1944. Gastropoda, Teil I. 1639 pp. In Schindewolf, O.H. (ed.) Handbuch der Paläozoologie, Bd. 6. Gebrüder Borntraeger, Berlin.

Wilson, B.R. 1993. Australian marine shells. Prosobranch gastropods. Part 1 (Prosobranch gastropods). 408 pp. Odyssey Publishing, Kallaroo, Western Australia.

Yin, H.-F. & Yochelson, E.L. 1983. Middle Triassic Gastropoda from Qingyan, Ghizhou Province, China: Euomphalacea and Loxonematacea. Journal of Paleontology 57, 1098–1127.

Zardini, R. 1978. Fossili Cassiani (Trias Mediosuperiore) Atlanti dei Gasteropodi della Formazione di S. Cassiano Raccolti della Regione Dolomitica Atorno a Cortina d’Ampezzo. 58 pp. Cortina d’Ampezzo.

Zardini, R. 1980. Fossili Cassiani (Trias Mediosuperiore) Atlanti dei Gasteropodi della Formazione di S. Cassiano Raccolti della Regione Dolomitica Atorno a Cortina d’Ampezzo. 14 pp. Cortina d’Ampezzo.

Zardini, R. 1985. Fossili Cassiani (Trias Mediosuperiore) Atlanti dei Gasteropodi della Formazione di S. Cassiano Raccolti della Regione Dolomitica Atorno a Cortina d’Ampezzo. 16 pp. Cortina d’Ampezzo.

Zittel, K.A. 1873. Die Gastropoden der Stramberger Schichten. Paläontologische Mittheilungen aus dem Museum des königlich Bayrerischen Staates 2, 311–491.

Zittel, K.A. 1895. Grundzüge der Palaeontologie. 971 pp. R. Oldenbourg, München and Leipzig.