Article: Feeding habits in trilobites
We briefly review the various types of feeding habits in marine arthropods, and suggest that the trilobites adopted a range of different feeding strategies. We show that much of the variety of trilobite exoskeletal development, particularly in the cephalon, can be explained as a response to the adoption of specific feeding modes. We regard the primitive mode as having been predatory/scavenging, both from morphological grounds and by out-group comparison, but this habit had a long subsequent history in the group. Predators/scavengers included those trilobites with rigidly braced and attached conterminant or impendent hypostomes, which often developed posterior forks or rasps used by the animals for manipulating prey after it had been grasped by the ‘gnathobases’. Advanced predators often acquired expanded anterior glabellar lobes which are associated with the ingestion of bulky food; concomitantly, the largest trilobites of all had predatory morphology. Associated trace fossils are of the Rusophycus type in which impressions of limb bases and rarely the hypostome can be seen. Detritivors were derived from predators by detachment of the hypostome from the doublure in natant mode; it is significant that the hypostome of such feeders exhibited little change thereafter. The typical detritivor morphology is of the ‘generalized’ ptychopariid type, common in outer shelf habitats, with rectangular or tapering glabellas and small to moderate overall size. It is suggested that in some species the hypostome may have functioned as a ‘scoop’ directly to aid ingestion of sediment. Trace fossils of Cruziana semiplicata type have been associated with sediment ploughing in this feeding mode. Filter feeders evolved a vaulted cephalic chamber of trinucleimorph type, and elevated thoraces, often flanked by extended genal spines. Where it is known, the hypostome is curved up inside the cephalic chamber, within which sediment stirred into suspension by the limbs was sorted for edible particles. Filter feeding trilobites are typically small, and are uncommon outside muddy habitats. Bean-like Rusophycus are the associated trace fossils. In trinucleids ingress of the feeding current was alongside the thorax and out through the fringe pits. The combination of different feeding modes with adaptation for different prey and/or particle sizes goes some way to account for the variety of trilobites cohabiting in a single site (alpha diversity). We do not claim that the model accounts for all morphological variation displayed by the group.