Many Late Paleozoic environments have been interpreted as marine because of the co-occurrence of supposedly exclusively marine trace fossils. Beginning in the Late Ordovician, however, nonmarine trace-fossil diversity increased throughout the Paleozoic. This diversification of nonmarine organisms and nonmarine trace fossils was especially prevalent in Devonian and later times. Diversification of freshwater organisms is indicated by the large number of freshwater fish, arthropods, annelids and molluscs that had developed by the Carboniferous. In addition to diverse freshwater assemblages, entirely terrestrial vertebrate and invertebrate ecosystems had developed by the Devonian. This rapid diversification of freshwater and terrestrial organisms is inherently linked to development and diversification of land plants and subsequent shedding of large quantities of organic detritus in nonmarine and marginal-marine areas. Nearshore marine organisms and their larvae that are able to tolerate relatively short periods of lowered salinities will follow salt-water wedges inland during times of reduced freshwater discharge. Similarly, amphidromous marine organisms will migrate periodically inland into nonmarine environments. Undoubtedly, both of these processes were active in the Paleozoic. However, both processes are restricted to stream/distributary channels, interdistributary bays, or estuaries. Therefore, the presence of diverse trace-fossil assemblages in association with floodplain deposits is interpreted to reflect true nonmarine adaptation and diversity. Conversely, diverse trace-fossil assemblages in association with stream/distributary channel deposits, interdistributary-bay deposits, or estuarine deposits may reflect migration of salt-water wedges inland, or migration of marine organisms into freshwater environments (amphidromy), or both. ?? 1989.
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The potential of paleozoic nonmarine trace fossils for paleoecological interpretations