Graptolites nearly became extinct in the latest Wenlock in all preserved stratigraphic sequences of this age. Graptolite mortalities occurred along the western coast of Laurentia and at sites that surrounded the ProtoTethys. Graptolite mass mortalities took place among deep-water, open ocean dwelling organisms. After the mass mortalities, only the Pristiograptus dubius group and retiolids, surface or near-surface dwellers, survived. For a period of time, little speciation or diversification occurred. The base of the Ludlow is marked by diversification, with appearances of S. colonus, M. nilssoni, and other groups which occur in near surface waters. None of the extensive plate movements postulated for the Silurian readily explain the mass extinctions that occurred. During the Silurian, global temperatures were warmer than present and atmospheric oxygen concentrations were lower, creating extensive oceanic anoxia. Below the oxygenated surface layers of the ocean, was an anoxic, non-sulfidic zone (i.e. nitrate-reducing) above a sulfidic zone. Graptolites lived over a range of depth from the oxygenated zone to either near or in the nitrate-reducing zones. As the oxygen concentration declined through the Silurian, the depth of the oxic zone would have become shoaler with expanding anoxia. Late Wenlock graptolites that were unable to migrate to shallower depths, living in borderline oxygen conditions, could have been killed, resulting in the mortalities of the late Wenlock. Only those graptolites that were surface dwellers survived, adapted and reradiated.