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The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott-Rust Quarry in New York State, USA. Walcott-Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages.

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.

Trilobites represent a model for 'evo-devo' studies in fossil euarthropods, owing to a rare developmental trait: the biomineralization of the dorsal exoskeleton soon after hatching. Many fossilized trilobite ontogenies thus feature early stages - the protaspides - characterized by non-articulated, calcified dorsal exoskeletons. The recent discovery of a protaspid-like fossil occurring with aglaspidid remains in Middle Ordovician strata of Siberia has been interpreted as evidence for the presence of protaspides in these distant relatives of trilobites. In this contribution, we demonstrate that this Siberian protaspis actually belongs to the asaphid trilobite Isotelus (or a related taxon), a genus likely present in the same bed. We conclude that protaspid larvae still represent a developmental trait unique to Trilobita., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

A 365 million year-old trilobite moult-carcass assemblage was described by Błażejowski et al. (2015) as the oldest direct evidence of moulting in the arthropod fossil record. Unfortunately, their suppositions are insufficiently supported by the data provided. Instead, the morphology, configuration and preservational context of the highly fossiliferous locality (Kowala Quarry, Poland) suggest that the specimen consists of two overlapping, queued carcasses. The wider fossil record of moulting actually extends back 520 million years, providing an unparalleled opportunity to study behaviour, ecology and development in early animals. Taking cues from modern analogues, it is possible to quantify precise details about moulting behaviour to determine broad-scale evolutionary trends, ontogenetic sequences and morphological selection pressures. In this review, we argue that this rich source of data has been underused in evolutionary studies, though has great potential for investigating the life history and evolution of arthropods in deep time., (© 2016 The Authors. Bioessays published by WILEY Periodicals, Inc.)

Ecdysis, the process of moulting an exoskeleton, is one of the key characters uniting arthropods, nematodes and a number of smaller phyla into Ecdysozoa. The arthropod fossil record, particularly trilobites, eurypterids and decapod crustaceans, yields information on moulting, although the current focus is predominantly descriptive and lacks a broader evolutionary perspective. We here review literature on the fossil record of ecdysis, synthesising research on the behaviour, evolutionary trends, and phylogenetic significance of moulting throughout the Phanerozoic. Approaches vary widely between taxonomic groups, but an overall theme uniting these works suggests that identifying moults in the palaeontological record must take into account the morphology, taphonomy and depositional environment of fossils. We also quantitatively analyse trends in trilobite ecdysis based on a newly generated database of published incidences of moulting behaviour. This preliminary work reveals significant taxonomic and temporal signal in the trilobite moulting fossil record, with free cheek moulting being prevalent across all Orders and throughout the Phanerozoic, and peaks of cephalic moulting in Phacopida during the Ordovician and rostral plate moulting in Redlichiida during the Cambrian. This study and a review of the literature suggest that it is feasible to extract large-scale evolutionary information from the fossil record of moulting., (Copyright © 2015 Elsevier Ltd. All rights reserved.)