HEMIASTERID ECHINOIDS (ECHINODERMATA, SPATANGOIDA) FROM THE CRETACEOUS TETHYS TO THE PRESENT-DAY MEDITERRANEAN

Hemiasterid echinoids form one of the oldest and most varied spatangoid groups. Current data on stratigraphical and geographical distribution patterns show that their biogeographical history is closely related to Mediterranean-Tethyan structural and climatic changes. The group first appeared in the Aptian in the central Tethys (southwestern Europe) and developed and diversified remarkably in the Late Cretaceous through the subgenera Hemiaster (Hemiaster), H. (Bolbaster) and H. (Leymeriaster). Marked morphological changes arose in the early Paleogene when new Tertiary forms H. (Trachyaster), Opissaster and Ditremaster, originating in the Caribbean and Madagascan regions, respectively, replaced H. (Bolbaster) which partly survived and H. (Hemiaster) and H. (Leymeriaster) which disappeared entirely. These new forms achieved their greatest diversity during the Eocene and thrived in the Mediterranean basin until it was sealed off from the Indo-Paacific in the Middle Miocene. Then Late Neogene climatic and structural events reduced hemiasterids to only a few Mediterranean species. The survivors of modern groups persisted in refuge areas of the western Mediterranean until middle Messinian times before being reduced to only one species, the last fossil one, during the Pliocene. Then, the Cenozoic groups H. (Trachyaster) and Opissaster definitively disappeared. Outside the Mediterranean basin primitive H. (Bolbaster) and modern Ditremaster, which colonized deep-water Atlantic environments from the Late Cretaceous and Middle Miocene respectively, survived the Messinian and Plio-Pleistocene climatic crises. They flourished in the northern Atlantic where their present-day descendants Ditremaster (Sarsiaster) griegii and H. (Bolbaster) expergitus still survive. H. (Bolbaster) expergitus spread worldwide and colonized the western Mediterranean in the Plio-Pleistocene via the Straits of Gibraltar.