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Within 520,000 yr of the C-T boundary, nontropical late Cenomanian biotas experienced 45-75~o species extinction, depending on the group, through a series of discrete, ecologically graded, short-term events, or steps, beginning with subtropical and warm temperate stenotopic biotas, and terminating with more broadly adapted cool temperate biotas. These extinction events were closely linked or coeval with abrupt, large-scale perturbations in the ocean-climate system, as evidenced by major fluctuations in trace elements (including Ir)
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