Thoracic structure and enrolment style in middle Cambrian Eccaparadoxides pradoanus presages caudalization of the derived trilobite trunk

The ability to enrol effectively evolved several times among trilobites. Here, we show that, unlike most redlichiid trilobites that could not enrol, both morphotypes of Eccaparadoxides pradoanus from the middle Cambrian of Spain enrolled so as to enclose most of the ventral surface beneath the exoskeleton and possessed specialized articulating devices that facilitated this behaviour. The holaspid thorax of all E.pradoanus was divided into two principal regions. The boundary between these marked a transition from anterior segments with short pleural spines, fulcra and ridge-and-groove inner pleural regions to posterior segments with longer, acuminate pleural spines that lack fulcra and inner pleural regions. Devices that aid articulation, such as fulcra with short articulating pleural surfaces, the petaloid articulating facet and long articulating half rings, are concentrated in the anterior region. These features, and the large number of specimens preserved in various degrees of enrolment, suggest an enrolment procedure in which the rear part of the trunk, containing both the posterior thorax and the pygidium, rotated as a single unit without significant internal flexure. As these posterior trunk articulations were apparently not required to permit enrolment, concentrating flexure in the anterior may have presaged the caudalized condition seen in many derived trilobite groups that encapsulated, in which a larger proportion of the trunk segments were allocated to the mature pygidium, and therefore unable to articulate.