Secondary Evolutionary Escalation between Brachiopods and Enemies of Other Prey

The fossil record of predation indicates that attacks on Paleozoic brachiopods were very rare, especially compared to those on post-Paleozoic mollusks, yet stratigraphically and geographically widespread. Drilling frequencies were very low in the early Paleozoic (≪1%) and went up slightly in the mid-to-late Paleozoic. Present-day brachiopods revealed frequencies only slightly higher. The persistent rarity of drilling suggests that brachiopods were the secondary casualties of mistaken or opportunistic attacks by the enemies of other taxa. Such sporadic attacks became slightly more frequent as trophic systems escalated and predators diversified. Some evolutionarily persistent biotic interactions may be incidental rather than coevolutionary or escalatory in nature.