An asynchronous Mesozoic marine revolution: the Cenozoic intensification of predation on echinoids

Predation traces found on fossilized prey remains can be used to quantify the evolutionary history of biotic interactions. Fossil mollusc shells bearing these types of traces provided key evidence for the rise of predation during the Mesozoic marine revolution (MMR), an event thought to have reorganized global marine ecosystems. However, predation pressure on prey groups other than molluscs has not been explored adequately. Consequently, the ubiquity, tempo and synchronicity of the MMR cannot be thoroughly assessed. Here, we expand the evolutionary record of biotic interactions by compiling and analysing a new comprehensively collected database on drilling predation in Meso-Cenozoic echinoids. Trends in drilling frequency reveal an Eocene rise in drilling predation that postdated echinoid infaunalization and the rise in mollusc-targeted drilling (an iconic MMR event) by approximately 100 Myr. The temporal lag between echinoid infaunalization and the rise in drilling frequencies suggests that the Eocene upsurge in predation did not elicit a coevolutionary or escalatory response. This is consistent with rarity of fossil samples that record high frequency of drilling predation and scarcity of fossil prey recording failed predation events. These results suggest that predation intensification associated with the MMR was asynchronous across marine invertebrate taxa and represented a long and complex process that consisted of multiple uncoordinated steps probably with variable coevolutionary responses.

Experimental neoichnology of post-autotomy arm movements of sea lilies and possible evidence of thrashing behaviour in Triassic holocrinids

Echinoderms exhibit remarkable powers of autotomy. For instance, crinoids can shed arm and stalk portions when attacked by predators. In some species, it has been reported that the autotomized arms display vigorous movements, which are thought to divert the attention of predators. This phenomenon, however, has not been well explored. Here we present results of experiments using the shallowest water species of living stalked crinoid (Metacrinus rotundus) collected at 140 m depth. A wide range of movements of detached arms, from sluggish writhing to violent flicks, was observed. Interestingly, autotomized arms produce distinct traces on the sediment surface. They are composed of straight or arched grooves usually arranged in radiating groups and shallow furrows. Similar traces were found associated with detached arms of the oldest (Early Triassic) stem-group isocrinid (Holocrinus). This finding may suggest that the origins of autotomy-related thrashing behaviour in crinoids could be traced back to at least the Early Triassic, underscoring the magnitude of anti-predatory traits that occurred during the Mesozoic Marine Revolution. A new ethological category, autotomichnia, is proposed for the traces produced by thrashing movements of shed appendages.