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Temporal and Latitudinal Trends in the Biodiversity of European Atlantic Cenozoic Gastropod (Mollusca) Faunas

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Temporal and Latitudinal Trends in the Biodiversity of European Atlantic Cenozoic Gastropod (Mollusca) Faunas Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 14 Temporal and latitudinal trends in the biodiversity of European Atlantic Cenozoic gastropod (Mollusca) faunas. A base for the history of biogeographic provinces Pierre LOZOUET 1 Abstract: A general overview of the biodiversity of the marine fauna during the Tertiary period is de- veloped in the context of the biogeographical evolution of the European realm. This study combines a reappraisal of the literature with a unique first-hand source of data on the richest marine group (the gastropods) from over a 25 million year period (Early Oligocene to Late Miocene). In total the French deposits have yielded more than 10,000 species from the Eocene to the Upper Miocene. Evidence of significant bias in the fossil record is pointed out for the Palaeocene (Danian) and Upper Eocene (Pria- bonian). For the period considered (Palaeocene to Late Miocene, ca 50 million years) the second highest diversity is reported in the Late Oligocene. This study reveals also the importance of pre-Mioce- ne extinctions of genera. The Oligocene was when the latitudinal differentiation of the faunas was grea- test. The local generic gastropod richness ranges from 59 (Rupelian of Belgium) to 494 (Chattian of Aquitaine). A clear trend towards homogenization appears in the Late Oligocene which leads to the de- velopment of a vast biogeographical region named Euro-West Africa. This faunal evolutionary pattern has never been demonstrated before and is a novel feature of the biogeography of the Eastern-Atlantic region. Key Words: Biogeography; Tethys; extinction; France; Paleogene; Miocene; molluscs. Citation: LOZOUET P. (2014).- Temporal and latitudinal trends in the biodiversity of European Atlantic Cenozoic gastropod (Mollusca) faunas. A base for the history of biogeographic provinces.- Carnets de Géologie [Notebooks on Geology], Brest, vol. 14, n° 14, p. 273-314. Résumé : Évolution temporelle et latitudinale de la biodiversité des faunes de gastéropodes (Mollusca) du Cénozoique atlantique européen. Une base pour l'histoire des provinces bio- géographiques.- Une revue globale de la biodiversité des faunes marines du tertiaire est proposée dans le cadre de l'évolution biogéographique du domaine européen. Cette étude est fondée sur une re- vue critique de la littérature et une source unique de données sur le groupe marin le plus riche (les gastéropodes) examinés durant une période de 25 millions d'années (Oligocène inférieur à Miocène su- périeur). Au total, les dépôts français renferment plus de 10 000 espèces dans l'intervalle Éocène à Miocène supérieur. Des biais importants dans les archives fossiles sont soulignés pour le Paléocène (Danien) et l'Éocène supérieur (Priabonien). La richesse générique locale s'étire de 59 genres (Rupélien de Belgique) à 494 genres (Chattien d'Aquitaine). Pour la période considérée (Paleocène à Moicène supérieur, environ 50 Ma), la seconde plus grande diversité (richesse spécifique) est relevée pour l'Oli- gocène supérieur. L'Oligocène inférieur est la période durant laquelle la différentiation latitudinale des faunes est la plus grande. Une nouvelle tendance apparaît à l'Oligocène supérieur qui conduit à la for- mation d'une vaste région biogéographique nommée Euro-Ouest Africaine. Cette évolution faunistique n'avait jamais été démontrée avant et apparaît comme un nouveau schéma dans la biogéographie des faunes Est-Atlantiques. Mots-clefs : Biogéographie ; Téthys ; extinction ; France ; Paléogene ; Miocène ; mollusques. Introduction they provide one of the prime sources of infor- mation on the history of the biosphere. Para- Located where the North Sea and tropical doxically we do not know how many species of Tethyan faunas converge, the French Atlantic molluscs occurred in the Cenozoic of the French Cenozoic domain outranks all others in Europe Atlantic. Therefore, I propose estimates based with its particularly rich molluscan faunas. The on two approaches. The first is for Eocene de- preservation of fossil molluscs is also exceptio- posits and is founded principally on revised nal, being the finest in the World for the Eocene bibliographic data but includes new data and (the famous Lutetian malacofauna of the Paris personal research. The second, a more detailed Basin), the best in the Tethyan realm for the approach, is for the Oligocene and Miocene Lower Oligocene (Rupelian = Stampian stage) deposits, with a new generic and species and the Upper Oligocene (Chattian stage), and synthesis based on a huge quantity of partly one of the best for the Lower and Middle Mio- unpublished material collected in the Aquitaine, cene. Loire and Paris basins (LOZOUET, 1997). Finally, Because molluscs dominate Cenozoic marine a comparison of families' specific richness fossils (KOWALEWSKI et al., 2002) and outnumber should provide insight into the global evolution all other macro-invertebrate taxonomic groups, of the malacofauna. 1 Muséum national d'Histoire naturelle, 55, rue Buffon, CP51, 75231 Paris cedex 05 (France) [email protected] Published online in final form (pdf) on October 14, 2014 [Editor: Bruno GRANIER; language editor: Stephen CAREY] 273 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 14 The paleobiogeography (provincialism) of cludes the Paris Basin, the Aquitaine Basin and French Atlantic Cenozoic molluscan faunas was the Loire, Brittany, and Cotentin (Normandy) in based on qualitative observations (CAVELIER, France and the southern North Sea Basin (Bel- 1979; BRÉBION, 1974, 1988), using restricted gium, Denmark and Germany), the Late Burdi- sources of data. This new review is the first galian/Middle Miocene of the Netherlands, and general overview on the basis of a larger col- the Upper Oligocene of Hungary (Paratethys ex- lection of data personally verified (Appendix tension) (Fig. 1). Table 3) using a multivariate analysis at the For the comparison of global species generic level. Several questions may then be richness we used data from the Palaeocene and addressed: How much have the faunas changed Paleogene (Eocene and Oligocene) of the Paris after the eastward closure of the Tethys Sea? Basin, Miocene of the Loire Basin and Oligocene Was the latitudinal gradient of specific richness and Miocene of the Aquitaine Basin (Fig. 1). On reinforced after the closure? Did the faunas the other hand the paleobiogeographic recon- affected by long-term global cooling evolve structions for the Oligocene and the Miocene towards more marked provincialism? are based on the generic distribution and are limited to malacofaunas from the Lower Study area, geological and Oligocene of Belgian and Germany, the Upper palaeobiogeographic setting Oligocene of Belgian, Germany and Hungary The geographical area taken into account in- and the Middle Miocene of the Netherlands. Figure 1: Locations of the faunas analyzed in the Cenozoic of the Eastern Atlantic coast basins and its North Sea ex- tension. 274 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 14 Global palaeobiogeography Celtic province. In contrast, the Mesogean realm (for example CAVELIER, 1979) is strictly During the Palaeogene, the composition of equivalent to Tethyan realm. In fact Mesogean marine molluscan faunas of the Atlantic coasts is routinely used in French geology as equiva- is determined mainly by the influences of the lent to Mediterranean or Tethyan. Nordic realm (warm temperate waters) and the Tethyan realm (tropical and subtropical). The Material and methods presence or absence of an isthmus between the North Sea Basin and the Paris Basin plays a The main taxonomic dataset used is the spe- major role in local biogeographic changes. The cies-level data derived from material collected Middle Lutetian uplift of the Artois axis interrup- by the author. For basins for which I have little ted communication between the London-Belgian of my own data, the data result from a critical basins and the Atlantic. The Artois isthmus re- compilation from the literature (Appendix S1). opened only much later, after the Miocene, Over the course of 25 years I (alone or with when exchange was again possible via the En- collaborators) have described about 300 new glish Channel. Meanwhile, we called the sudden species, but 500-1000 or more, principally of invasion of the Paris Basin of molluscs of sou- small size, are still undescribed. In the thern Tethyan origin a "Mesogean bubble" (LO- database these species are labelled as "sp.". ZOUET, 2012). The best example is provided by From a taxonomic point of view these "sp." can the upper part of the Paris Basin Stampian (Lo- be considered as "morphospecies" or wer Oligocene, Rupelian Stage) where a signifi- "Operational Taxonomic Units" (OTUs). These cant part of the Pierrefitte Sands fauna, before- informal units are here taken to represent hand known only in the latitude of the Aquitaine specific-level taxa, equivalent to formal Basin, appeared suddenly (ca. 30 Ma). "species". During sampling, special attention was given to fragile gastropod specimens and, Much later, in the latest Oligocene (Chattian) in particular, to shells with well preserved to Middle Miocene (Langhian and Badenian protoconchs, because this part of the shell ages), the progressive closure of the eastern contains diagnostic characters. Identification of Tethyan seaway and its brief re-opening were almost all molluscs was done or checked by the the main palaeogeographical events. According author. to PILLER et al. (2007) this intermittent connec- tion was definitively sealed during the middle The terms "outcrops" or "sites" are used for Badenian (approx. 14-15 Ma). sections a few metres in extent. For example, the famous outcrop of Saucats, "Le Peloua" The severe cooling during middle Miocene (Burdigalian of Aquitaine; National Geological time (Serravallian age) is the first reliable evi- Reserve of Saucats-La Brède), is the outcrop of dence of a global change in the climate, sug- a single layer about 40 cm thick. Generally, gested to be an incipient glaciation. The "green- sampling was conducted in just one homoge- house period" was definitely over (PROTHERO, neous layer (the richest), which rarely exceeds 1994). However, before this time, a significant 2 m in thickness (1 m is the mean value).
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