Trilobite tagmosis and body patterning from morphological and developmental perspectives.

The Trilobita were characterized by a cephalic region in which the biomineralized exoskeleton showed relatively high morphological differentiation among a taxonomically stable set of well defined segments, and an ontogenetically and taxonomically dynamic trunk region in which both exoskeletal segments and ventral appendages were similar in overall form. Ventral appendages were homonomous biramous limbs throughout both the cephalon and trunk, except for the most anterior appendage pair that was antenniform, preoral, and uniramous, and a posteriormost pair of antenniform cerci, known only in one species. In some clades trunk exoskeletal segments were divided into two batches. In some, but not all, of these clades the boundary between batches coincided with the boundary between the thorax and the adult pygidium. The repeated differentiation of the trunk into two batches of segments from the homonomous trunk condition indicates an evolutionary trend in aspects of body patterning regulation that was achieved independently in several trilobite clades. The phylogenetic placement of trilobites and congruence of broad patterns of tagmosis with those seen among extant arthropods suggest that the expression domains of trilobite cephalic Hox genes may have overlapped in a manner similar to that seen among extant arachnates. This, coupled with the fact that trilobites likely possessed ten Hox genes, presents one alternative to a recent model in which Hox gene distribution in trilobites was equated to eight putative divisions of the trilobite body plan.