Drilling predation on spatangoid echinoids from the Miocene of Sardinia: a taphonomic and paleoecological perspective

Predation is of great importance in understanding the diversification of spatangoid echinoids as well as their preservation in the fossil record. Herein, the spatangoid-dominated Miocene carbonate succession of Punta Foghe (Montiferru area, central-western Sardinia, Italy) is studied with the aim of reconstructing paleoenvironmental conditions and investigating abiotic and biotic factors, including predation, that influence the paleoecology and preservation potential of spatangoid echinoids. The echinoid fauna is dominated by the spatangoid Agassizia and, to a lesser extent, the clypeasteroid Clypeaster. The fauna also includes both regular (undetermined diadematoids and camarodonts) and irregular (Plagiobrissus, Echinocyamus, and Koehleraster) forms. Echinoids and associated fossil content, along with taphonomic and sedimentological signatures, indicate an inner sublittoral environment with moderate energy conditions affected by high-energy events. Agassizia remains, which consist mainly of complete and well-preserved tests, commonly bear subcircular drill holes that are interpreted as the result of cassid gastropod predation and investigated with respect to size selectivity and stereotypy of attack sites. Potential biases related to drilling predation and biostratinomy on the preservation potential of spatangoid tests are discussed. Agassizia lacks any morphological adaptation to minimize high predation risk, including defensive spines and the ability to burrow deeply into the sediment. An antipredator behavior possibly relied on a gregarious life-history strategy, reducing the frequency of attacks on specific individuals by cassid predators.