Sea urchins are a major component of recent marine communities where they exert a key role as grazers and benthic predators. However, their impact on past marine organisms, such as crinoids, is hard to infer in the fossil record. Analysis of bite mark frequencies on crinoid columnals and comprehensive genus-level diversity data provide unique insights into the importance of sea urchin predation through geologic time. These data show that over the Mesozoic, predation intensity on crinoids, as measured by bite mark frequencies on columnals, changed in step with diversity of sea urchins. Moreover, Mesozoic diversity changes in the predatory sea urchins show a positive correlation with diversity of motile crinoids and a negative correlation with diversity of sessile crinoids, consistent with a crinoid motility representing an effective escape strategy. We contend that the Mesozoic diversity history of crinoids likely represents a macroevolutionary response to changes in sea urchin predation pressure and that it may have set the stage for the recent pattern of crinoid diversity in which motile forms greatly predominate and sessile forms are restricted to deep-water refugia.echinoderms | escalation | macroecology I t has long been hypothesized that predator-prey interactions represent a significant driving force of evolutionary change in the history of life (1-4). However, not only is predation itself hard to detect in the fossil record, which makes it difficult to ascertain its intensity over geologic time, but macroevolutionary predictions of the hypothesis are far from simple (5-13). Recent sea urchins (Echinoidea), are known to play a key role in shallow sea ecosystems as grazers and benthic predators that can modify the distribution, abundance, and species composition of coral and algal reef communities (14-16); however, only few data have hinted at the importance of sea urchins to crinoids (17)(18)(19).Crinoids (Crinoidea), commonly known as sea lilies or feather stars, were one of the dominant components of many shallow-sea environments through much of geologic history and a key contributor to the sedimentary record (20). Although predation by fish on crinoids and its evolutionary consequences have received the most attention (21-27), sparse data indicated that crinoids may be the prey of benthic invertebrates (28), most notably sea urchins (17-19, 29, 30). Recently it has been shown that during the Triassic, the radiation of cidaroid sea urchins capable of handling the crinoid skeleton coincided with high frequency of bite marks on crinoids likely produced by the jaw apparatus of these sea urchins (18). Because it was also during the Triassic that various modes of active and passive motility appeared among crinoids, a group that throughout its rich pre-Triassic history was almost exclusively sessile, it was argued that crinoid motility, an effective escape strategy against benthic predation, was an evolutionary response to echinoid predation (18).The hypothesized evolutionary response of crinoids to benthic pr...