Low drilling frequency in Norian benthic assemblages from the southern Italian Alps and the role of specialized durophages during the Late Triassic

Abstract

Drillholes represent one of the clearest lines of evidence for predation of benthic invertebrates in the fossil record and are frequently used as a primary proxy for predation intensity in the fossil record. Drillholes are abundant in the late Cretaceous and Cenozoic, but their occurrence is patchy in older deposits of the Mesozoic. The inconsistent record of drillholes in pre-Cretaceous deposits of Mesozoic age are problematic for interpretations of predation-prey dynamics and adaptive radiations, and the role of taphonomy or diagenesis have not been resolved. Here we present drilling percentages for assemblages of well-preserved shelly benthic invertebrates (mainly comprised of bivalves and rare gastropods) from the upper Norian (Upper Triassic) in northern Italy in order to compare these values with reported drilling percentages from the Carnian San Cassiano Formation, a rare Triassic sedimentary unit that has yielded many drilled fossils. The Norian fossil deposits reported here are comparable to those of the San Cassiano in terms of depositional environment, preservation, and region, and can be reasonably compared to the drilling percentage of fossils from the San Cassiano. The sampled deposits are collected from marly limestone horizons in the Argillite di Riva di Solto in the Southern Italian Alps, deposited in the Lombardian Basin, and which are interbedded with shale units containing well-preserved fish and arthropod fossils, enabling a correlation between paleoecological structure of the shelly benthos and the demersal-pelagic predator diversity. Over four hundred bivalve fossils yielded a drilling percentage of 0.24% (1/406), which is typical for fossil assemblages of this age, but the single occurrence of a drillhole in this study is in marked contrast to the many drilled specimens reported from the San Cassiano Formation deposit in Italy. The drilled specimen (with complete drillhole) was an infaunal bivalve and no incomplete drillholes were observed in other specimens. Thus, drilling percentages for the Triassic are consistently low, but present, suggesting that drilling predation was an ecologically minimal influence to benthic communities and unlikely to have driven the significant ecological changes observed in benthic communities during the Late Triassic. Although drilling predation occurred during the Late Triassic, we present an updated database of specialized durophagous predators (including fishes, sharks, and reptiles) that are likely to have been more ecologically influential on benthic communities during the Norian Stage, fishes in particular.