Quantifying intra-and interspecific variability in trilobite moulting behaviour across the Palaeozoic

Abstract

Moulting of a protective exoskeleton is a defining characteristic of Euarthropoda, and its evolution can be explored through analysing moults preserved in the fossil record. Our most complete record comes from the Trilobita, which were uniquely flexible in moulting compared to other arthropod groups. This study presents the first broad-scale quantitative analysis of trilobite moulting. Trends in moulting variability with taxonomy and through the Palaeozoic are explored by looking at the occurrences of six moulting characteristics: opening of the facial and ventral sutures; and disarticulation of the cephalon, cranidium, thorax, and pygidium. Significant differences in moulting across taxonomic and temporal groups were identified using chi-squared analyses, and biases with sampling and diversity identified through correlation analyses. Occurrences of facial and ventral suture opening, and cephalic disarticulation, significantly varied between orders and Epochs. These likely result from the prevalence of ventral suture moulting in Redlichiida and cephalic disarticulation in Phacopida, and their relevant diversity patterns. The data show high levels of intraspecific variability in moulting; ~40% of species showed multiple moulting characteristics. Redlichiida and the early Cambrian are the most intraspecifically variable, with greater variability likely an adaptation to the initial radiation and establishment of trilobites into new niches. The lon-gest-lived group, Proetida, showed the lowest levels of intraspecific variability, which may suggest greater specialism later in the Palaeozoic. Ultimately, datasets such as this advocate the need to study behaviours in the fossil record on a broad-scale, because they help us build a comprehensive picture of extinct groups as living animals.