Development, Trait Evolution, and the Evolution of Development in Trilobites.

Trilobites offer one of the best fossil records of any arthropod group. This is due to a number of factors, most notably the combination of (1) having inhabited areas where organisms are more likely to be buried and ultimately fossilized; and (2) having had a highly biomineralized exoskeleton more likely to survive the stresses of fossilization. This biomineralized exoskeleton was also morphologically complex, bearing traits that had ecological significance, and was present throughout postembryonic development, from larval to adult stages. Because the morphology of the exoskeleton changed gradually across molts during development, it is possible to reconstruct ontogenetic series for many species. Over the last decade, studies have documented both variation in modularity among closely related species and conserved developmental patterns among modules. In the latter case, trait evolution could still occur through modification of rates of morphological change along otherwise conserved ontogenetic trajectories. At the clade level, the pattern of expression and release of new exoskeletal segments during post-embryonic development was generally conserved across most species, but the relative timing of different segmentation events could vary, and developmental traits appear to have been relatively labile across the clade's evolutionary history. Most recently, comparative analyses indicate that the association between segmentation events and the timing of shifts in the rate of ontogenetic shape change varies across species. Despite these advances, we still know relatively little about how development constrained or contributed to trait evolution in trilobites, and almost nothing about the origin of novel traits in trilobites. A major (but removable) obstacle is the current lack of well-supported trilobite phylogenies that span higher taxonomic levels.