The plankton and the benthos: origins and early history of an evolving relationship

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Abstract
Modern marine plankton communities include a broad diversity of metazoans that are suspension-feeding or micropredatory as adults. Many benthic marine species have larval stages that reside, and often feed, in the plankton for brief to very long periods of time, and most marine benthic communities include large numbers of suspension-feeders. This has not always been the case. Cambrian benthic communities included relatively few suspension-feeders. Similarly, there were few metazoan clades represented in the plankton, either as adult suspension-feeders or as larvae. Review of the fossil record suggests that the diversification of the plankton and suspension-feeding marine animals began in the Late Cambrian and continued into the Ordovician. These changes were accompanied by, and probably influenced, concurrent major changes in the marine realm, including an increase in tiering within benthic communities, the replacement of the Cambrian fauna by the Paleozoic fauna, and a general taxonomic diversification. The ultimate cause of these changes is uncertain, but it appears likely that the plankton was and is a refuge from predation and bioturbation for adults and larvae alike. The expansion in plankton biomass thus provided increased ecological opportunities for suspension-feeders in the plankton and benthos.