Paleobiogeography of scleractinian reef corals: Changing patterns during the Oligocene–Miocene climatic transition in the Mediterranean

During the Oligocene–Miocene Greenhouse-to-Icehouse climatic transition, the biogeography of reef corals or zooxanthellate-like scleractinian corals was gradually changing from a pan-tropical Tethyan Province in the Eocene to three reef-coral Provinces of the Western Atlantic–Caribbean, Indo-Pacific and Mediterranean.Our REEFCORAL database encompasses updated and homogenized data on paleoenvironmental and systematicsof scleractinian corals occurring in the Oligocene and Miocene outcrops from circum-Mediterranean regions, provided by most of relatively recently published data in the literature and by the study of published and unpublished collections of coral specimens from the same area, including the important collections housed at the MNHN (Paris) and our own collections. As there is no validated direct criterion for the identification of the coral-zooxanthellate symbiosis in the fossil record, and considering the difficulty to use the biogeochemical approaches in the context of this study, the subjectivity of the morphological criteria and the relative recent age of the fossil corals we are dealing with, a uniformitarian approach has been used for inferring the symbiotic status of scleractinian genera in REEFCORAL. Among the 158 genera included in our database, 93 can be considered as zooxanthellate and 10 have a doubtful zooxanthellate status. This relativelyexhaustive database was used to reconstruct the temporal and spatial distribution of scleractinian corals in the Mediterranean during the Oligocene–Miocene time in order to discuss the interplaying effects of the global cooling at that time, the re-organization of the Tethyan realm resulting from the African, Arabian and Eurasian plate collision and the emergence of the Alpine chains, driving the gradual northward movement of the whole region outside the tropical/subtropical belt. It is shown that the structure of the Mediterranean z-coral Oligocene–Miocene paleobiodiversity was characterized by many geographically-restricted genera with a moderate to short stratigraphical range and a few long-ranging widespread genera. A major consequence of this structure is that the extinction pattern has proceeded through the preferential extinction of rare-occurrence genera through time. The potential rapid long-distance dispersal of most coral larvae compared to the size of the Oligocene–Miocene Mediterranean, explains why no biogeographical subprovinces can be distinguished for the z-coral fauna. On a local scale,ecological processes tend to sort coral taxa by limiting z-coral development to geographically restricted and discontinuous areas. This accounts for the large amount of geographically-restricted taxa forming the Mediterranean coral fauna.The interaction of plate-tectonics, Alpine orogenesis and climate at local to subregional scales exerts strong controls over the spatio-temporal distribution of z-coral assemblages within the circum-Mediterranean realm. In particular, we suggest that the richness and composition of the Eastern Atlantic coral fauna are indirectly related to the opening and closure of the eastern seaway connection with the Indian Ocean, which controlled the E–W circulation of surface waters and hence the westwards dispersal of pelagic larvae. At the scale of the whole region, the gradual regional climatic change produced by the northwards migration of the entire area, superimposed on the global cooling, appears in large part responsible for the extinction pattern of z-corals through time in the Mediterranean biogeographical Province.