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Malformed trilobite specimens present important insight into understanding how this extinct arthropod group recovered from developmental or moulting malfunctions, pathologies, and injuries. Previously documented examples of malformed trilobite specimens are often considered in isolation, with few studies reporting on multiple malformations in the same species. Here we report malformed specimens of the ellipsocephaloid trilobite Estaingia bilobata from the Emu Bay Shale Konservat-Lagerstätte (Cambrian Series 2, Stage 4) on Kangaroo Island, South Australia. Ten malformed specimens exhibiting injuries, pathologies, and a range of teratologies are documented. Furthermore, five examples of mangled exoskeletons are presented, indicative of predation on E. bilobata. Considering the position of malformed and normal specimens of E. bilobata in bivariate space, we demonstrate that the majority of malformed specimens cluster among the larger individuals. Such specimens may exemplify larger forms successfully escaping predation attempts, but could equally represent individuals exhibiting old injuries that were made during earlier (smaller) growth stages that have healed through subsequent moulting events. The available evidence from the Emu Bay Shale suggests that this small, extremely abundant trilobite likely played an important role in the structure of the local ecosystem, occupying a low trophic level and being preyed upon by multiple durophagous arthropods. Furthermore, the scarcity of malformed E. bilobata specimens demonstrates how rarely injuries, developmental malfunctions, and pathological infestations occurred within the species.

Trilobites were notably flexible in the moulting behaviours they employed, producing a variety of moult configurations preserved in the fossil record. Investigations seeking to explain this moulting variability and its potential impacts are few, despite abundant material being available for study. We present the first quantitative study on moulting in a single trilobite species using a dataset of almost 500 moult specimens of Estaingia bilobata from the Cambrian (Series 2, Stage 4) Emu Bay Shale, South Australia. Specimens were categorized by moulting mode (Salter's or Sutural Gape) and their associated configurations, and their body proportions measured from both a museum collection (including a bycatch sample) and a randomly-collected field sample. This enabled analysis of the proportion of E. bilobata specimens displaying the Sutural Gape and Salter's modes of moulting and their different configurations, and tests for association between moulting behaviour and body proportions. The results show a wide range of E. bilobata moulting configurations in all samples, suggesting that configurations represent definable instances in a largely continuous spectrum of variation. Analyses comparing body proportions of specimens showing the two modes of moulting were non-significant, suggesting there is no true association between moulting behaviour and body proportion, except for a single significant result for body length. All results were relatively consistent between the museum and field samples. However, removing accessioned specimens from the museum sample brought results even further in line with the field sample, supporting the need for consideration of museum collection bias in palaeontological analyses.

Trilobites are an abundant group of Palaeozoic marine euarthropods that appear abruptly in the fossil record c. 521 million years ago. Quantifying the development of morphological variation (or 'disparity') through time in fossil groups like trilobites is critical in understanding evolutionary radiations such as the Cambrian 'explosion'. Here, I use geometric morphometrics to quantify 'cumulative disparity' in functionally-important structures within the trilobite cephalon across their initial radiation during Cambrian Series 2. Overall cephalic disparity increased rapidly and attained a maximum within several million years. This pattern is dominated by the cephalic outline (in particular the genal spines), reflecting rapid, convergent expansion to the extremes of morphospace in a few early families. In contrast, removing the outline and focusing on structures such as the glabella and eye ridges (associated with feeding and vision, respectively) showed a more gradual increase in disparity, closer in line with taxonomic diversity and supporting the hypothesis of a relatively accurate trilobite fossil record. These contrasting patterns suggest that disparity in different structures was constrained in different ways, with extrinsic (ecological) factors probably having the major impact on overall disparity. It also implies that patterns of disparity in isolated substructures cannot necessarily be taken individually as representative of overall morphologies.

Supplementary Materials (text and 2 figures) toRe-assessing growth and mortality estimates for the Ordovician trilobite Triarthus eatonito be published in the journal Paleontology (June/July 2022)

The fossiliferous lower Cambrian (Series 2) successions along the north coast of Kangaroo Island, South Australia—known collectively as the Kangaroo Island Group—can be divided into two main areas: a western succession located between Snelling Beach and Smith Bay, which comprises the Mt McDonnell Formation (base), Stokes Bay Sandstone and Smith Bay Shale; and an eastern succession that extends from Emu Bay to Point Marsden, represented by exposures of the White Point Conglomerate (base), Marsden Sandstone, Emu Bay Shale and Boxing Bay Formation (top), with an overlap in the Cape d’Estaing/Emu Bay area. Some previous interpretations of the Kangaroo Island Group stratigraphy suggest that the western succession stratigraphically underlies the eastern succession. Most of the previously reported and described Cambrian fossils come from the eastern succession, especially the internationally significant Emu Bay Shale Konservat-Lagerstätte. Here we report the trilobites Redlichia takooensis and Balcoracania dailyi from the Smith Bay Shale near Freestone Creek, indicating that the eastern and western successions are at least partly contemporaneous. The present investigation indicates that the Smith Bay Shale of the western succession can be correlated with the stratigraphic interval represented by the Marsden Sandstone and the overlying Emu Bay Shale of the eastern succession. The Kangaroo Island Group was deposited as part of a fan delta system with the eastern succession representing the proximal part, and the western succession representing the distal part of the fan delta. The lack of substantial conglomerate units within the Kangaroo Island Group to the west of Cape d’Estaing suggests that the tectonic uplift that led to the deposition of the White Point Conglomerate was concentrated in the area immediately to the north of Emu Bay. Refereed/Peer-reviewed

A length–frequency sample (n = 295) from a fossil population of the Ordovician trilobite Triarthrus eatoni Hall, 1838, assembled and analyzed by J. L. Cisne in 1973 is here reexamined using methods of length–frequency analysis commonly used in fishery science and marine biology. Theoretical considerations and the empirical data at hand suggest that the growth of T. eatoni was not “linear,” but asymptotic, as is the growth of most Recent marine invertebrates. The parameters of the von Bertalanffy growth function (L∞ = 41 mm, K = 0.29 yr−1) suggest that T. eatoni, which apparently lived in a challenging environment, grew somewhat more slowly than the extant marine isopod Ceratoserolis trilobitoides (Eights, 1833), used here as Recent analogue to T. eatoni. This trilobite probably lived up to 10 years, rather than the suggested 4 years, and its mortality rate was 15%–20% per year rather than 30%–40% per year. These represent the first estimates of trilobite absolute growth characteristics using methods known to accurately model growth in extant water-breathing ectotherms. These provide a baseline for trilobite growth that can be used to make inferences about growth in other species. The approach used here may also be applied to other trilobites for which suitable length–frequency data exist.

The exceptional fossil record of trilobites provides our best window on developmental processes in early euarthropods, but data on growth dynamics are limited. Here, we analyse post-embryonic axial growth in the Cambrian trilobite Estaingia bilobata from the Emu Bay Shale, South Australia. Using threshold models, we show that abrupt changes in growth trajectories of different body sections occurred in two phases, closely associated with the anamorphic/epimorphic and meraspid/holaspid transitions. These changes are similar to the progression to sexual maturity seen in certain extant euarthropods and suggest that the onset of maturity coincided with the commencement of the holaspid period. We also conduct hypothesis testing to reveal the likely controls of observed axial growth gradients and suggest that size may better explain growth patterns than moult stage. The two phases of allometric change in E. bilobata , as well as probable differing growth regulation in the earliest post-embryonic stages, suggest that observed body segmentation patterns in this trilobite were the result of a complex series of changing growth controls that characterized different ontogenetic intervals. This indicates that trilobite development is more complex than previously thought, even in early members of the clade.

The trilobite Redlichia Cossmann, 1902 is an abundant element of the lower Cambrian (Series 2, Stage 4) Emu Bay Shale (EBS) Konservat-Lagerstatte on Kangaroo Island, South Australia. Well-preserved...

Studies that reveal detailed information about trilobite growth, particularly early developmental stages, are crucial for improving our understanding of the phylogenetic relationships within this iconic group of fossil arthropods. Here we document an essentially complete ontogeny of the trilobite Redlichia cf. versabunda from the Cambrian Series 2 (late Stage 4) Ramsay Limestone of Yorke Peninsula in South Australia, including some of the best-preserved protaspides (the earliest biomineralized trilobite larval stage) known for any Cambrian trilobite. These protaspid stages exhibit similar morphological characteristics to many other taxa within the Suborder Redlichiina, especially to closely related species such as Metaredlichia cylindrica from the early Cambrian period of China. Morphological patterns observed across early developmental stages of different groups within the Order Redlichiida are discussed. Although redlichiine protaspides exhibit similar overall morphologies, certain ontogenetic characters within this suborder have potential phylogenetic signal, with different superfamilies characterized by unique trait combinations in these early growth stages.

In order to maximize the utility of future studies of trilobite ontogeny, we propose a set of standard practices that relate to the collection, nomenclature, description, depiction, and interpretation of ontogenetic series inferred from articulated specimens belonging to individual species. In some cases, these suggestions may also apply to ontogenetic studies of other fossilized taxa.

The Ediacara Member of the Flinders Ranges (South Australia) preserves body and trace fossils of the Ediacara biota. Fossils span five lithofacies representative of a range of shallow-marine environments and are preserved as in situ and transported material. Previous work has demonstrated a relationship between paleo environment and taxa at the Nilpena fossil site. We expand the analysis to include facies-taxa data from a further nine localities across the Flinders Ranges to assess if the taxa–paleo environment relationship is site specific or valid at a regional scale. The new analysis demonstrates that the distribution of taxa within the lithofacies, as a proxy for paleo environment, is non-random. This preliminary analysis presents a beta diversity-like spatial turnover across the range of shallow marine Ediacaran environments, and demonstrates taxonomic assemblages are specific to given paleo environmental zones. These assemblages are consistent over a broad spatial extent and also a presumed temporal distribution. This specificity indicates that a marked sensitivity to environmental parameters was present in these communities, as demonstrated by the non-random distribution of taxa and spatial turnover of biotic assemblage throughout the gradational environments of the Ediacara Member. This study highlights the variability and heterogeneity that is a characteristic of shallow marine settings, and offers a novel approach to the future investigation of the relationship between Ediacaran environments and taxa assemblages. Refereed/Peer-reviewed

Michael S.Y. Lee, Alessandro Palci, Marc E.H. Jones, Michael W. Caldwell, James D. Holmes, Robert R. Reisz

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases— Redlichia rex and Olenoides serratus —and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus . Results show that L. polyphemus , S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus , were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the ‘arms race' between predators and biomineralized prey.

Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock 1964 being extremely abundant, and the larger Redlichia takooensis Lu 1950 being common. Many specimens within the EBS represent complete moulted exoskeletons, which is unusual for Cambrian fossil deposits. The abundance of complete moults provides an excellent record that has allowed the recognition of various recurrent moult configurations for both species, enabling the inference of movement sequences required to produce such arrangements. Moult configurations of E. bilobata are characterised by slight displacement of the joined rostral plate and librigenae, often accompanied by detachment of the cranidium, suggesting ecdysis was achieved by anterior withdrawal via opening of the cephalic sutures. Moulting in R. takooensis often followed the same method, but configurations show greater displacement of cephalic sclerites, suggesting more vigorous movement by the animal during moulting. Both species also show rare examples of Salter’s configuration, with the entire cephalon anteriorly inverted, and several other unusual configurations. These results indicate that moulting in trilobites was a more variable process than originally thought. In contrast, other Cambrian Konservat-Lagerstätten with an abundance of trilobites (e.g., Wheeler Shale, USA, and Mount Stephen Trilobite Beds, Canada)show larger numbers of ‘axial shields’ and isolated sclerites, often interpreted as disarticulated exuviae. This points to a higher level of disturbance from factors such as animal activity, depositional processes, or water movement, compared to that of the EBS, where quiescent conditions and intermittent seafloor anoxia contributed to an unparalleled trilobite moulting record.

© 2017 Elsevier. This manuscript version is made available under the CC-BY-NC-ND 4.0 license: http://creativecommons.org/licenses/by-nc-nd/4.0/ This author accepted manuscript is made available following 24 month embargo from date of publication (Dec 2017) in accordance with the publisher’s archiving policy, Exceptional fossil deposits exhibiting soft-part preservation, or Konservat-Lagerstätten, are prevalent in Cambrian rocks and provide detailed information on fossil assemblages not available from conventional deposits. It has long been recognised that many of these assemblages exhibit certain taxonomic similarities, with many elements seemingly having cosmopolitan distributions. These types of assemblages, particularly those of Cambrian age, have become known as Burgess Shale-type (BST) biotas, named for the famous deposit in the Canadian Rocky Mountains where fossils preserved in this way were first discovered. This study provides the first broad-scale analysis of the assemblage relationships between all major BST biotas. We compiled a database of the presences and absences of over 600 genera within 12 Lagerstätten from Laurentia, Siberia, South China and East Gondwana, ranging in age from Cambrian Series 2 through Series 3 (late-early to middle Cambrian; c. 518–502 Ma), and analysed this using a variety of quantitative methods in order to investigate the relationships between these sites. Non-metric multidimensional scaling (NMDS) ordination, cluster analysis and Parsimony Analysis of Endemicity (PAE) were used to group localities and examine relationships. We also used Bayesian inference and illustrate the benefits of this approach to biogeographic studies. Results suggest that both space and time have important effects on the taxonomic constitution of BST biotas, and that the similarity of these assemblages appears to increase from Series 2 through Series 3, largely driven by increases in cosmopolitanism of biomineralised taxa such as trilobites and brachiopods. There is also evidence of higher-level taxonomic turnover across this period. Endemic taxa help amplify these patterns, despite their frequent exclusion from biogeographic analyses.