Shell ornamentation as a likely exaptation: evidence from predatory drilling on Cenozoic bivalves

Predation is an important process in modern oceans and in the evolutionary history of marine ecosystems. Consequently, it has been hypothesized that shelled prey modified their ornamentation in response to predation. However, bivalve ornamentation has also been argued to be important in maintaining a stable life position in the sediment and in burrowing. To test whether concentric ribs were effective against drilling by carnivorous gastropods, we examined drill hole position and completeness for four Cenozoic bivalve species that differ in rib strength (Astarte radiata, A. goldfussi, Lirophora glyptocyma, and L. latilirata). The percentage of drill holes located between the ribs increases with increasing rib strength, whereas the percentage of drill holes on top of ribs decreases. This result suggests that gastropods select the drill hole site more effectively as rib strength increases, thereby saving time and energy, and that natural selection favors gastropods that select drill hole s...