Inferring internal anatomy from the trilobite exoskeleton: the relationship between frontal auxiliary impressions and the digestive system

The digestive system of trilobites is rarely preserved. As a result, many aspects of its organization remain unknown. Fortunately, the exoskeleton sometimes preserves evidence of
soft-tissue attachment sites that can be used to infer internal anatomy. Among them are the frontal auxiliary impressions (FAIs), probable soft-tissue insertion sites located on the fronto-median glabellar lobe of some trilobites. FAIs are herein described in the Carboniferous trilobite Phillipsia belgica Osmo´ lska 1970 – representing the only known example of such structures in the Proetida and their youngest occurrence. A taphonomic scenario is proposed to explain their variable preservation. Although particularly common in the Phacopina, FAIs or FAI-like structures are also found in several orders that differ greatly. Comparisons with modern analogues suggest that FAIs might represent attachment sites for extrinsic muscles associated with a differentiated crop within the foregut. A review of purported remains of the trilobite digestive system indicates that it usually consisted of a tube-like tract flanked by a variable number of metamerically
paired diverticulae. Its anterior portion is not particularly individualized, except in a few specimens that might hint at the presence of a crop. This differentiation of a crop might have constituted a secondarily evolution of the foregut in trilobites, occurring independently in different clades. Accompanied by a strengthening of associated extrinsic muscles, this modification of the foregut might explain the presence of more conspicuous muscle insertion sites on the glabella. Study of FAIs might therefore provide new data on the anatomy of the foregut in trilobites and evidence of diverse feeding habits.