Predators and Predation in Paleozoic Marine Environments

The Paleozoic body fossil record of potential benthic predators includes nautiloid and ammonoid cephalopods, phyllocarids, decapods, and several lineages of gnathostomes. The latter group, in particular, radiated rapidly during the Devonian. In the pelagic realm, predator-prey interactions involving cephalopods and some nektonic arthropods probably appeared in the Ordovician. Again, evidence indicates intensification of pelagic predation, much of it by arthrodires and sharks on other fishes, during the Devonian radiation of gnathostomes. Trace fossils provide direct evidence of predatory attack from the Ediacarian and Early Cambrian onward, but with a substantial increase in the Siluro-Devonian. Brachiopod and molluscan shells and trilobite exoskeletons show evidence of healed bite marks and peeling from the Cambrian onward, but with an increased frequency in the Devonian. Predatory drill holes with stereotypical position and prey-species preference are found in brachiopods (Cambrian onward) and mollusks (Ordovician onward); boreholes also show increased frequency in the middle Paleozoic. Certain of these boreholes are tentatively attributable to platyceratid gastropods. Hard-shelled benthic organisms with thicker, more spinose skeletons may have had a selective advantage as durophagous predators increased. Brachiopods, gastropods, trilobites, and crinoids show an abrupt increase in spinosity beginning in the Siluro-Devonian. But spinosity decreases after the early Carboniferous. Late Paleozoic benthos may have taken refuge in smaller size and resistant, thick-walled skeletons, as well as endobenthic and cementing modes of life. Conversely, in the pelagic realm, external armor was reduced, while more efficient, fast-swimming modes of life (e.g., in sharks) increased in the post-Devonian.