Alleged Lessepsian foraminifera prove native and suggest Pleistocene range expansions into the Mediterranean Sea

Biogeographical patterns are increasingly modified by the human-driven translocation of species, a process that accelerated several centuries ago. Observational datasets, however, rarely range back more than a few decades, implying that a large part of invasion histories went unobserved. Small-sized organisms, like benthic foraminifera, are more likely to have been reported only recently due to their lower detectability compared to larger-sized organisms. Recently detected native species of tropical affinity may have thus been mistaken for non-indigenous species due to the lack of evidence of their occurrence in pre-invasion records. To uncover the unobserved past of the Lessepsian invasion-the entrance of tropical species into the Mediterranean through the Suez Canal-we collected sediment cores on the southern Israeli shelf. We deployed state-of-the-art radiocarbon techniques to date 7 individual foraminiferal tests belonging to 5 alleged non-indigenous species and show that they are centuries to millennia old, thus native. Two additional species previously considered non-indigenous occurred in centennial to millennia-old sediments, suggesting their native status. The evidence of multiple tropical foraminiferal species supposed to be non-indigenous but proved native in the eastern Mediterranean suggests either survival in refugia during the Messinian Salinity Crisis (5.96-5.33 million years) or, more likely, dispersal from the tropical Atlantic and Indo-Pacific during the Pleistocene. In the interglacials of this epoch, higher sea levels may have allowed biological connectivity between the Mediterranean and the Red Sea for shallow-water species, showing that the Isthmus of Suez was possibly a more biologically porous barrier than previously considered.