Skip to main content Accessibility help

Biogeographic patterns and Plio-Pleistocene extinction of Bivalvia in the Mediterranean and southern North Sea

  • Sergio Raffi (a1), Steven M. Stanley (a2) and Raffaella Marasti (a1)


An evaluation of the history of polysyringian species (filibranchs and eulamellibranchs) reveals that the huge early Pliocene bivalve fauna of the Mediterranean Basin (MB) and North Sea Basin (NSB) suffered heavy extinction during late Pliocene and early Pleistocene time. This is evidenced by low survivorship of early Pliocene species in the Recent and by a decline in species richness of the two basins from 323 known early Pliocene species to 198 living species.

Several kinds of evidence indicate that cooling rather than the areal effect of eustatic sea-level lowering was the primary cause of the excessive extinction: (1) heavy Plio-Pleistocene extinction of mollusks was not global but concentrated around the margins of the northern Atlantic—an ocean fringed by polar ice caps; (2) taxa of tropical affinities were most severely affected; (3) heavy extinction occurred in the MB in areas not marked by facies change; (4) in the MB the onset of extinctions coincided with the onset elsewhere, but because of tectonic activity, water depths in the MB were not under tight eustatic control; (5) 14 species present in both the MB and NSB during early Pliocene time are now restricted to waters south of the NSB; and (6) the majority of species common to the two basins during the early Pliocene (eurythermal species) have survived to the present.

Molluscan data support palynological evidence that the climate in the MB was warmer and less seasonal in early Pliocene time than today, when latitudinal temperature gradients are steeper. Molluscan evidence indicates that the North Sea is exceptional in being less seasonal (though cooler) today than in early Pliocene time, and we attribute this anomaly to the local effects of the Gulf Stream, which was strengthened in mid-Pliocene time by the uplift of the Isthmus of Panama.

The heavy extinction in the MB and NSB about 3.2–3.0 ma ago approximately coincided with the earliest deposition of glacial tills in Iceland and with isotopic shifts in the tests of planktonic foraminifers preserved in deep-sea cores. Additional heavy extinction probably coincided with a pulse of severe cooling in late Pliocene time, 2.5–2.4 ma ago. Heavy extinction of mollusks in the MB and NSB continued into the early Pleistocene but not into the middle and late Pleistocene, apparently because by this time it was primarily only eurythermal species that survived. Today the molluscan faunas of the MB and NSB are unusually eurythermal; few species are restricted to a single biogeographic province.



Hide All
Adam, W. and Knudsen, J. 1955. Note sur quelques espèces de mollusques marine nouveaux ou peu connus de l'Afrique occidentale. Bull. Inst. Roy. Sci. Nat. Belgique. 31(61):125.
Bassindale, R. 1961. On the marine fauna of Ghana. Proc. Zool. Soc. Lond. 137:481510.
Bellardi, L. 1872–1890. I Molluschi dei terreni terziari del Piemonte e della Liguria. Vols. 1–6. Mem. R. Acc. Sci. Torino. 27, 29, 34, 37, 38, 40.
Blackwelder, B. W. 1981. Late Cenozoic stages and molluscan zones of the U.S. Middle Atlantic coastal plain. Paleontol. Soc. Mem. 12.
Bouchet, P. 1982. Les Terebridae (Mollusca, Gastropoda) de l'Atlantique oriental. Boll. Malacol. 18(9–12):185216.
Briggs, J. 1974. Marine Zoogeography. 475 pp. McGraw-Hill Book Co.; New York.
Buchanan, J. B. 1954. Marine molluscs of the Gold Coast. J. W. Afr. Sci. Ass. 1:3045.
Buchanan, J. B. and Anderson, M. M. 1955. Additional records to the marine molluscan fauna of the Gold Coast. J. W. Afr. Sci. Ass. 1:5761.
Coomans, H. E. 1962. The marine mollusk fauna of the Virginian area as a basis for defining Zoogeographical provinces. Beaufortia. 9:83104.
Cornet, R. and Marche-Marchad, I. 1951. Mollusques. In: Inventaire de la faune marine de Roscoff. Tr. St. Biol. Roscoff. Suppl. 5:180.
Dautzenberg, Ph. 1898. Coup d'oeil sur la faune de Départment de la Loire-Inférieure. Mollusques. Extract from “Nantes et la Loire-Inférieure.” 23 pp. Fr. Ass. Adv. Sci.; Nantes.
Dautzenberg, Ph. 1912. Mission Gruvel sur la côte occidentale d'Afrique (1909–1910). Mollusques marins. Ann. Inst. Océanogr. 5(3):1111.
Dautzenberg, Ph. 1913. Atlas de poche des coquilles des côtes de France. 152 pp. P. Klincksieck; Paris.
Dautzenberg, Ph. 1927. Mollusques provenant des campagnes scientifiques du Prince Albert Ier de Monaco dans l'Océan Atlantique et dans le Golfe de Rés. Camp. Sci. 72:1400.
Dautzenberg, Ph. and Durouchoux, P. 1913. Les mollusques de la Baie de Saint-Malo. Feuille Jeun. Natur. Suppl. 522:164.
Dautzenberg, Ph. and Fischer, H. 1906. Mollusques provenant des dragages effectués à l'ouest de l'Afrique pendant les campagnes scientifiques de S.A.S. le Prince de Monaco. Rés. Camp. Sci. 32:1125.
Dautzenberg, P. and Fischer, H. 1912. Mollusques provenant des campagnes de l'Hirondelle et de la Princesse-Alice dans les Mers due Nord. Rés. Camp. Sci. 37:1629.
De Stefani, C. and Pantanelli, D. 1878. Molluschi pliocenici dei dintorni di Siena. Bull. Soc. Malac. It. 4:5215.
Dollfus, G. F. and Berkeley Cotter, J. C. 1909. Mollusques tertiaires du Portugal. Le Pliocène au Nord du Tage. Ier pt. Pélécypoda. 103 pp. Comm. Serv. Géol. Port.; Lisbonne.
Dollfus, G. F., Berkeley Cotter, J. C., and Gomes, J. P. 1903–1904. Mollusques tertiaires du Portugal. Planches de Céphalopodes, Gastéropodes et Pélécypodes laissées par F. A. Pereira Da Costa. 55 pp. Comm. Serv. Géol. Port.; Lisbonne.
Ekman, S. 1953. Zoogeography of the sea. 417 pp. Sidgwick & Jackson; London.
Erünal-Erentöz, L. 1958. Mollusques du Néogène des Bassins de Karaman, Adana et Hatay (Turquie). Publ. Et. Rech. Minièr. Turquie. Sér. C. 4:1232.
Fischer, H. 1898. Liste des Mollusques marine recueillis à Guéthary et à St. Jean-de-Luz. Pp. 127136. Trans. Soc. Sci. Arcachon; Paris.
Fischer-Piette, E. 1975. Révision des Venerinae s.s. (Mollusques Lamellibranches). Mém. Mus. Nat. Hist. Nat. Sér. A Zool. 93:164.
Fischer-Piette, E. 1977. Révision des Cardiidae (Mollusques Lamellibranches). Mém. Mus. Nat. Hist. Nat. Sér. A Zool. 101:1212.
Fischer-Piette, E. and Delmas, D. 1967. Révision des Mollusques Lamellibranches du genre Dosinia Scopoli. Mém. Mus. Nat. Hist. Nat. Sér. A Zool. 47:191.
Fischer-Piette, E., Fischer, P. H., Germain, I., and Pallary, P. 1942. Les Mollusques d'Adanson. J. Conch. 85:103365.
Fischer-Piette, E. and Métivier, B. 1971. Révision des Tapetinae (Mollusques Bivalves). Mém. Mus. Nat. Hist. Nat. Sér. A Zool. 71:1106.
Fischer-Piette, E. and Vukadinovic, D. 1977. Suite des révisions des Veneridae (Moll. Lamellibr.) Chioinae, Samaranginae et complément aux Vénus. Mém. Mus. Nat. Hist. Nat. Sér. A Zool. 106:3186.
Forbes, E. 1856. Map of the distribution of the marine life. Plate no. 31. In: Johnson, Alexander K. The Physical Atlas of Natural Phenomena (new ed.). Edinburgh and London.
Fredj, G. 1974. Stockage et exploitation des données en écologie marine. C-Considérations biogéographiques sur le peuplement benthique de la Meditérranée. Mém. Inst. Océanogr. Monaco. 7:188.
Gartner, S. Jr. 1977. Calcareous nonnofossil biostratigraphy and revision zonation of the Pleistocene. Mar. Micropaleontol. 2:125.
Gladenkov, Y. B. 1981. Marine Plio-Pleistocene of Iceland and problems of its correlation. Quat. Res. 1 15:1823.
Glemarec, M. 1968. Pélécypodes marins de la côte atlantique française. 21 pp. Laboratoire de Zoologie; Brest.
Glemarec, M. 1978. Distribution bathymétrique & latitudinale des bivalves du Golfe de Gascogne. Haliotis. 9:2332.
Glibert, M. 1957–1959. Pélécypodes du Diestien, du Scaldisien et du Merxemien de la Belgique. Bull. Inst. Roy. Sci. Nat. Belgique. 32(9):140, 32(47):1–28, 34(42):1–27, 35(36):1–24.
Gubert, M. 1958. Tableau stratigraphique des mollusques du Néogène de la Belgique. Bull. Inst. Roy. Sci. Nat. Belgique. 34(32):132.
Habe, T. 1964. Shells of the western Pacific in color. 233 pp. Hoikusha Publishing Co. Ltd.; Osaka.
Hall, C. 1964. Shallow-water marine climates and molluscan provinces. Ecology. 45(2):226234.
Harmer, F. W. 1914–1925. The Pliocene Mollusca of Great Britain. Palaeontol. Soc. 67, 68, 70, 71–73, 75, 76.
Hedgpeth, J. W. 1957. Marine biogeography. Geol. Soc. Am. 67:359382.
Heering, J. 1950. Pelecypoda (and Scaphopoda) of the Pliocene and older Pleistocene deposits of the Netherlands (Peel Region). Meded. Geol. Sticht. 9:1225.
Hidalgo, J. G. 1917. Fauna malacologica de España, Portugal y las Baleares: Moluscos testaceos marinos. Trab. Mus. Nac. Cienc. Nat. Ser. Zool. 30:1752.
Holme, N. A. 1961. The bottom fauna of the English Channel. J. Mar. Biol. Ass. U.K. 41:397461.
Keigwin, L. E. 1978. Pliocene closing of the Isthmus of Panama, based on biostratigraphic evidence from nearby Pacific Ocean and Caribbean Sea cores. Geology. 6:630634.
Keigwin, L. D. and Thunnel, R. C. 1979. Middle Pliocene climatic change in the western Mediterranean from faunal and oxygen isotopic trends. Nature. 282:294296.
Kira, T. 1965. Shells of the Western Pacific in color. 224 pp. Hoikusha Publishing Co. Ltd.; Osaka.
Knudsen, J. 1956. Marine Prosobranchs of Tropical West Africa (Stenoglossa). Atlantide Report. 4:7110.
Lacourt, A. W. 1977. Contribution à la connaissance de la fauna marine de la région d'Arcachon. Zool. Meded. 52:223253.
Lande, E. 1975. A Collection of Pelecypoda from Möre and Romsdal, Western Norway. Sarsia. 59:714.
Lecointre, G. 1952. Recherches sur de Néogène et la Quaternaire marine de la côte atlantique du Maroc. Tome 2. Paléontologie. Serv. Géol. Maroc. Notes et Mém. 99:1173.
Lewy, Z. 1975. Molluscs distribution on the Atlantic continental shelf off Southern Spanish Sahara, West Africa. Meteor Forsch.-Ergebnisse. 21:5260.
MacAndrew, R. 1850. Notes on the distribution and range in depths of mollusca and other marine animals observed on the coasts of Spain, Portugal, Barbary, Malta and southern Italy in 1849. Rep. Br. Assn. Adv. Sci. 1850:264304.
MacAndrew, R. and Woodward, H. 1863. Species of Mollusca obtained in Corunna Bay. Ann. Mag. Nat. Hist. 14:232324.
Marasti, R. and Raffi, S. 1976. Osservazioni biostratigrafiche e paleoecologiche sulla malacofauna del Piacenziano di Maiatico (Parma, Emilia occidentale). Boll. Soc. Paleontol. Ital. 15:189214.
Marasti, R. and Raffi, S. 1977. Diversità tassonomica dei bivalvi pliocenici del Bacino padano: 1° I bivalvi dell'Emilia occidentale. L'Atenco Parmense. Acta Natur. 13:370.
Marasti, R. and Raffi, S. 1979. Observations on the paleoclimatic and biogeographic meaning of the Mediterranean Pliocene molluscs. State of the problem. VII. International Congress on the Mediterranean Neogene, Athens. Ann. Géol. Pays Hellén. Tome hors série 1979. 2:727734.
Marasti, R. and Raffi, S. 1980a. Extinction of polysyringian bivalves in the Mediterranean Pliocene. Pp. 107115. In: Instituto di Geologia Univ. Parma. Volume dedicato a Sergio Venzo. Grafiche STEP; Parma.
Marasti, R. and Raffi, S. 1980b. La diversità tassonomica dei bivalvi del Pliocene mediterraneo: elenco preliminare. Proposta di lavoro al Gruppo di ricerca C.N.R. “Paleobenthos.” Prospettive di ricerca. 30 pp. Riunione del 24–28 maggio 1980, Palermo.
Marche-Marchad, I. 1958. Nouveau catalogue de la Collection des Mollusques testacés marins de l'I.F.A.N. Cat. Inst. Franc. Afr. Noire. 14:164.
Mars, P. 1963. Les faunes et la stratigraphie du Quaternaire Méditerranéen. Rec. Trav. St. Mar. End. 28:6197.
McDougall, I. and Wensink, H. 1966. Paleomagnetism and geochronology of the Pliocene-Pleistocene lavas in Iceland. Earth and Planet. Sci. Letters. 1:232236.
McMillan, N. F. 1968. British shells. 196 pp. F. Warne & Co. Ltd.; London–New York.
Meco Cabrera, J. 1981. Neogastropodos fosiles de Las Canarias orientales. An. Est. Atlanticos. 27:601615.
Meco Cabrera, J. 1982. Los Bivalvos fosiles de Las Canarias orientales. An. Est. Atlanticos. 28:65125.
Monbet, Y. 1973. Contribution à la connaissance de la fauna malacologique du plateau continental au large d'Arcachon. Bull. Soc. linnéenne Borddeaux. 3(7):154156.
Mosquera, E. R. 1983. Moluscos de la Ria de Vigo. I. Gastropodos. 383 pp. Velograf, S. A. Milladoiro, s/n; Santiago de Campostela (Espñna).
Nickles, M. 1950. Mollusques testacés marins de la Côte occidentale d'Afrique. Manuels ouest-africains. 2:1269.
Nickles, M. 1955. Scaphopodes et Lamellibranches récoltés dan l'Ouest Africain. Atlantide Rep. 3:93237.
Nobre, A. 1931. Moluscos Marinhos de Portugal. 463 pp. Impresa Portuguesa; Pôrto.
Nobre, A. 1936. Moluscos Marinhos de Portugal. Vol. 2, 377 pp. Comp. Edit. Minho; Barcelos.
Nordsieck, F. 1968. Die europäischen Meeres-Gehäuseschnecken (Prosobranchia). 273 pp. G. Fischer Verlag; Stuttgart.
Nordsieck, F. 1969. Die europäischen Meeresmuscheln (Bivalvia). 256 pp. G. Fischer Verlag; Stuttgart.
Nordsieck, F. and Talavera, F. G. 1979. Moluscos marinos de Canarias y Madera (Gastropoda). 208 pp. Aula de Cultura; Tenerife.
Ockelmann, W. K. 1958. The zoology of east Greenland. Marine Lamellibranchiata. Meddelelser om Gronland. 122(4):256.
Ortea Rato, J. A. 1977. Moluscos marinos gastropodos y bivalves del litoral asturiano entre Ribadesella y Ribadeo con especial atencion a la subclase de los opistobranquios. 581 pp. Tesis Doctoral. Sec. Biol. Univers.; Orviedo.
Pasteur-Humbert, Ch. 1962a. Les mollusques marins testacés du Maroc. I. Les Gastéropodes. Trav. Inst. Sci. Chérifien. Sér. Zool. 23:1245.
Pasteur-Humbert, Ch. 1962b. Les mollusques marins testacés du Maroc. II. Les Lamellibranches et les Scaphopodes. Trav. Inst. Sci. Chérifien. Sér. Zool. 28:1184.
Peres, J. M. 1967. The Mediterranean benthos. Oceanogr. Mar. Biol. Ann. Rev. 5:449533.
Philippi, R. A. 1936. Enumeratio molluscorum Siciliae. 268 pp. Berolini.
Philippi, R. A. 1844. Enumeratio molluscorum Siciliae. Vol. 2:303 pp. Halis Saxonum.
Piani, P. 1967. Catalogo dei molluschi conchiferi viventi nel Mediterraneo. Boll. Malacologico. 16(5–6):113224.
Raffi, S. 1982. Stato delle conoscenze sulla sezione pleistocenica marina del Torrente Stirone (Parma). Pp. 141144. In: Cremonini, G. and Ricci Lucchi, F., eds. Guida alla geologica del margine appenninico-padano. Guida Geol. Reg. S.G.I.; Bologna.
Raffi, S. 1986. Are the Mediterranean Lower Pleistocene Boreal Guests only of historical importance or possibly a basic tool of paleoclimatic inference? Palaeogeogr., Palaeoclimatol., Palaeoecol. In press.
Raffi, S. and Marasti, R. 1982. The Mediterranean bioprovince from the Pliocene to the Recent: observations and hypotheses based on the evolution of the taxonomic diversity of molluscs. Pp. 151177. In: Montanaro Gallitelli, E., ed. Palaeontology, Essential of Historical Geology (Proc. Intern. Meeting, Venice). S.T.E.M. Mucchi; Modena.
Ruggieri, G. 1967. The Miocene and later evolution of the Mediterranean Sea. Pp. 283290. In: Adams, G. C. and Ager, D. V., eds. Aspects of Tethyan Biogeography. Systematics Association. Publication n.7.
Sacco, F. 1890–1904. I Molluschi dei terreni terziari del Piemonte e della Liguria. Vols. 6–30. Carlo Clausen; Torino.
Salvat, B. 1967. Mollusques des plages océaniques et semi-arbritées du Bassin d'Arcachon. Bull. Mus. Nat. Hist. Nat. Sér. 2, 39:11771191.
Saracino, C., Marano, G., Casavola, N., and Rizzi, E. 1982. Ciclo riproduttivo e biometria di Protopecten glaber (L.) (Bivalvia, Pectinidae) nel Basso Adriatico. Boll. Mus. Ist. Univ. Genova. Suppl. 50:332337.
Schopf, T. J. M., Fischer, J. B., and Smith, C. A. 1977. Is the marine latitudinal diversity gradient merely another example of the species area curve? Pp. 365386. In: Gray, J. and Boucot, A. J., eds. Historical Biogeography, Plate Tectonics, and the Changing Environment. Oregon State Univ. Press; Corvallis.
Shackleton, N. J., Backman, J., Zimmerman, H., Kent, D. V., Hall, M. A., Roberts, D. G., Schnitker, D., Baldauf, J. G., Desprairies, A., Homrighausen, R., Huddlestun, P., Keene, J. N., Kaltenback, A. J., Krumsiek, K. A. O., Morton, A. C., Murray, J. W., and Westberg-Smith, J. 1984. Oxygen isotope calibration of the onset of ice-rafting and history of glaciation in the North Atlantic region. Nature, 307:620623.
Shackleton, N. J. and Opdyke, N. D. 1976. Oxygen-isotope and paleomagnetic stratigraphy of Pacific core 28-239 Late Pliocene to Latest Pleistocene. In: Cline, R. M. and Hays, J. D., eds. Investigation of Late Quaternary Paleoceanography and Paleoclimatology. Geol. Soc. Am. Mem. 145:449464.
Sorgenfrei, T. 1958. Molluscan Assemblages from the Marine Middle Miocene of South Jutland and their Environments. Geol. Surv. Denmark. Ser. 2, 79:1503.
Stanley, S. M. 1982a. Glacial refrigeration and Neogene regional mass extinction of marine bivalves. Pp. 179191. In: Montanaro Gallitelli, E., ed. Palaeontology, Essential of Historical Geology (Proc. Intern. Meeting, Venice). S.T.E.M. Mucchi; Modena.
Stanley, S. M. 1982b. Species selection involving alternative character states: an approach to macroevolutionary analysis. Proc. N. Am. Paleontol. Conv. 2:505510.
Stanley, S. M. 1984. Marine mass extinctions: A dominant role for temperature. Pp. 69117. In: Nitecki, M. H., ed. Extinctions. Univ. Chicago Press; Chicago.
Stanley, S. M., Addicott, W. O., and Chinzei, K. 1980. Lyellian curves in paleontology: Possibilities and limitations. Geology. 8:422426.
Stanley, S. M. and Campbell, L. D. 1981. Neogene mass extinction of Western Atlantic molluscs. Nature. 293:457459.
Suc, J. P. and Zagwijn, W. H. 1983. Plio-Pleistocene correlations between the N-W Mediterranean and N-W Europe according to recent biostratigraphic and paleoclimatic data. Boreas. 12:153166.
Sverdrup, H. U., Johnson, M. W., and Fleming, R. H. 1942. The Oceans: Their physics, chemistry, and general biology. 1,087 pp. Prentice-Hall, Inc.; Englewood Cliffs, NJ.
Talavera, F. G. 1975. Moluscos de sedimentos de la plataforma continental de Mauritania. Bol. Inst. Esp. Oceanogr. 192:318.
Taylor, J. D. and Taylor, C. N. 1977. Latitudinal distribution of predatory gastropods on the eastern Atlantic shelf. J. Biogeog. 4:7381.
Tebble, N. 1966. British bivalve seashells: A handbook for identification. 212 pp. British Museum (Natural History); London.
Thunnell, R. C. and Williams, D. F. 1983. The stepwise development of Pliocene-Pleistocene paleoclimatic and paleooceanographic conditions in the Mediterranean: oxygen isotopic studies of DSDP Sites 125 and 132. Utrecht Micropaleontol. Bull. 30:111127.
Vail, P. R., Mitchum, R. M., and Thompson, S. 1977. Global cycles of relative changes of sea level. Am. Assoc. Petrol. Geol. Mem. 26:8397.
Van Straaten, L. M. J. U. 1956. Composition of shell beds formed in tidal flat environment in the Netherlands and in the Bay of Arcachon (France), Geol. Mijnb. 18:209226.
Van Urk, R. M. 1964. The genus Ensis in Europe. Basteria. 28:1344.
Van Urk, R. M. 1971. Fossil Ensis species in the Netherlands. Basteria. 35:176.
Van Urk, R. M. 1980. Fossil and recent Lutraria (Mollusca, Bivalvia) in Europe, with descriptions of four new species. Meded. Werkgr. Tert. Kwart. Geol. 17(5):235266.
Vermeij, G. J. 1980. Molluscan extinction at the Panama isthmus: Predation or food? Geol. Soc. Am. Abst. 12:541.
Vermeij, G. J. and Petuch, E. J.Differential extinction in tropical American molluscs: endemism, architecture, and the Panama land bridge. Malacologia. In press.
Wise, K. P. and Schopf, T. J. 1981. Was marine faunal diversity in the Pleistocene affected by changes in sea level? Paleobiology. 7:394399.
Wood, S. V. 1848–1882. A monograph of the Crag Mollusca or descriptions of shells from the Middle and Upper Tertiaries of the East of England. Palaeontol. Soc. 1, 5, 7, 10, 25, 27, 36.
Woodward, S. P. 1866. A manual of the Mollusca: A treatise on recent fossil shells. 542 pp. 2d ed.Virtue Brothers; London.
Zagwijn, W. H. 1974. The Plio-Pleistocene boundary in western and southern Europe. Boreas. 3:7597.


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed