During the Cambrian and Early Ordovician, when wide oceans separated all major continents, only pelagic marine organisms, such as plankton, could move freely across the open ocean and therefore the oceanic gaps between continents are easily detected in the fossil records of marine bottom dwellers and non-marine species. By the Late Ordovician, when continents were pushed closer together closing the oceanic gaps, benthos (brachiopods and trilobites) could spread between continents while ostracods and fishes remained isolated. As Laurussia formed during the Devonian and Pangaea formed, fish species in both Laurussia and Gondwana began to migrate between continents and before the end of the Devonian similar species were found on both side of what remained of the Variscan barrier.