During the Devonian (416-359 Mya) the combined landmass of Baltica and Avalonia rotated around Laurentia, which remained static near the Equator. The Laurentian warm, shallow seas and on shelves a diverse assemblage of benthos evolved, including the largest trilobites exceeding . The Old Red Sandstone Continent stretched across northern Laurentia and into Avalonia and Baltica but for most of the Devonian a narrow seaway formed a barrier where the North Atlantic would later open. Tetrapods evolved from fish in the Late Devonian, with the oldest known fossils from Greenland. Low sea-levels during the Early Devonian produced natural barriers in Laurussia which resulted in provincialism within the benthic fauna. In Laurentia the Transcontinental Arch divided brachiopods into two provinces, with one of them confined to a large embayment west of the Appalachians. By the Middle Devonian, these two provinces had been united into one and the closure of the Rheic Ocean finally united faunas across Laurussia. High plankton productivity from the Devonian-Carboniferous boundary resulted in anoxic events that left black shales in the basins of Laurentia.