Long-term clade-wide shifts in trilobite segment number and allocation during the Palaeozoic.

Arthropods are characterized by having an exoskeleton, paired jointed appendages and segmented body. The number and shape of those segments vary dramatically and unravelling the evolution of segmentation is fundamental to our understanding of arthropod diversification. Because trilobites added segments to the body post-hatching which were expressed and preserved in biomineralized exoskeletal sclerites, their fossil record provides an excellent system for understanding the early evolution of segmentation in arthropods. Over the last 200 years, palaeontologists have hypothesized trends in segment number and allocation in the trilobite body, but they have never been rigorously tested. We tabulated the number of segments in the post-cephalic body for over 1500 species, selected to maximize taxonomic, geographical and temporal representation. Analysis reveals long-term shifts in segment number and allocation over the 250-million-year evolutionary history of the clade. For most of the Palaeozoic, the median number of segments in the body did not change. Instead, the total range decreased over time and there was long-term increase in the proportion of segments allocated to the fused terminal sclerite relative to the articulated thoracic region. There was also increased conservation of thoracic segment number within families. Neither taxonomic turnover nor trends in functionally relevant defensive behaviour sufficiently explain these patterns.