The Early Pliocene re-colonisation of the deep Mediterranean Sea by benthic foraminifera and their pulsed Late Pliocene–Middle Pleistocene decline

Abstract

Ninety-five species and 19 genera of cosmopolitan, deep-sea benthic foraminifera belonging to the families Pleurostomellidae, Stilostomellidae and Nodosariidae, became extinct during the Late Pliocene–Middle Pleistocene. Only 50% of these (44 species) were present in the Pliocene or Pleistocene of the deep Mediterranean Sea (ODP Sites 654, 966, 967, 975, 976), being those which had successfully migrated in via the Strait of Gibraltar from the deep Atlantic following the annihilation of the Mediterranean deep-sea fauna during the Late Miocene Messinian Crisis. Most colonisation occurred within the first 0.8 myrs (5.3–4.5 Ma) after re-establishment of the Mediterranean–Atlantic link, with possibly a second lesser period of immigration in the Late Pliocene (3.4–3.0 Ma). We infer that colonisations may have been fortuitous and few in number, as some common members of the group in the Atlantic never succeeded in establishing in the Mediterranean Sea. There is no evidence of any new immigration events during the Pleistocene, implying that the present anti-estuarine circulation may have been in place throughout this period. Our studies suggest that these deep-water, low-oxygen-tolerant foraminifera survived the many periods of deep-water sapropel formation in the Pliocene–Early Pleistocene, possibly in somewhat shallower (~ 500 m) refuges with dysoxic, rather than anoxic conditions. The Pliocene–Pleistocene stratigraphic record of this group of elongate, cylindrical benthic foraminifera with constricted and specialised apertures is similar in the west and east Mediterranean basins. The group declined in abundance (flux) and diversity in two pulses, during the Late Pliocene (3.1–2.7 Ma) and the late Early Pleistocene (1.3–1.0 Ma) in concert with global, southern-sourced, deep-water sites (AABW, CPDW) and earlier than the single decline (1.0–0.6 Ma) in global, intermediate water sites (uNADW, AAIW). All species, with one possible exception, disappeared earlier in the Mediterranean than globally. The highest occurrence of any species of this group in Mediterranean sites was 0.8–0.43 Ma, comparable with 0.7–0.2 Ma outside with the youngest survivors being in abyssal, deep-water. Thus, despite the unusual oceanographic conditions and isolation, the deep Mediterranean Sea was in this case neither the centre for the evolution of new species nor a refuge where species survived after they had disappeared elsewhere.