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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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Trilobites are an iconic Paleozoic group of biomineralizing marine euarthropods that appear abruptly in the fossil record (c. 521 million years ago) during the Cambrian 'explosion'. This sudden appearance has proven controversial ever since Darwin puzzled over the lack of pre-trilobitic fossils in the Origin of Species, and it has generally been assumed that trilobites must have an unobserved cryptic evolutionary history reaching back into the Precambrian. Here we review the assumptions behind this model, and suggest that a cryptic history creates significant difficulties, including the invocation of rampant convergent evolution of biomineralized structures and the abandonment of the synapomorphies uniting the clade. We show that a vicariance explanation for early Cambrian trilobite palaeobiogeographic patterns is inconsistent with factors controlling extant marine invertebrate distributions, including the increasingly-recognized importance of long-distance dispersal. We suggest that survivorship bias may explain the initial rapid diversification of trilobites, and conclude that the group's appearance at c. 521 Ma closely reflects their evolutionary origins. A reassessment of early trilobite phylogenetic relationships and palaeobiogeographic patterns suggests that a cryptic evolutionary history is unlikely for this group. The abrupt appearance of trilobites is likely to closely reflect their evolutionary origins, and may be explained by survivorship biases inherent in the fossil record. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Trilobites are one of the most diverse and abundant fossil groups from the early Palaeozoic, and as such are useful for answering important questions about early animal evolution, including developmental processes. Ontogenetic information for a large number of trilobite species has been published, but cases where multiple articulated specimens are known across the full range of developmental stages are rare. The early Cambrian (Series 2, Stage 4) Emu Bay Shale biota from Kangaroo Island (South Australia) is numerically dominated by trilobites, particularly articulated specimens of the ellipsocephaloid Estaingia bilobata, which are present in densities of >600 individuals per square metre on certain bedding planes. Here we describe the essentially complete post‐embryonic ontogenetic series of E. bilobata from the Emu Bay Shale, and investigate patterns of growth relating to articulation and segmentation in this early Cambrian arthropod. Estaingia bilobata exhibits the hypoprotomeric mode of growth, with the epimorphic phase (the cessation of trunk segment generation) reached prior to the onset of the holaspid period. The meraspid pygidium had an extended equilibrium period in which anterior segment release into the thorax was matched by subterminal segment generation. Previously undocumented morphological features of E. bilobata, including the hypostome and bispinose pleural tips in holaspides, are also described. The growth characteristics and morphological features of E. bilobata documented herein strengthen close phylogenetic relationships between the Estaingiidae, Ellipsocephalidae and Xystriduridae. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The trilobite Redlichia Cossmann, 1902 is an abundant element of the lower Cambrian (Series 2, Stage 4) Emu Bay Shale (EBS) Konservat-Lagerstätte on Kangaroo Island, South Australia. Well-preserved, fully articulated specimens from this deposit are known to reach lengths of up to 25 cm, representing one of the largest known Cambrian trilobites. Until now, all Redlichia specimens from the EBS have been referred to Redlichia takooensis Lu, 1950, a species originally described from South China. Previous work recognized considerable differences in exoskeletal morphology among specimens of varying sizes, which was attributed to ontogeny. However, close examination of a large collection of recently acquired specimens shows that this variation actually represents two distinct morphs, interpreted here as separate species: R. takooensis, and a large, new species, Redlichia rex sp. nov. An analysis of morphological variation in holaspides ('adults') of the more common R. takooensis reveals considerable ontogenetic change occurred even during this later phase of growth. Some specimens of both Redlichia species from the EBS also exhibit exceptionally preserved soft-part anatomy, particularly the antennae and biramous appendages. Here, appendages (antenniform and biramous) and digestive structures are described, and biramous appendage reconstructions of R. rex sp. nov. are presented, which show a striking resemblance to some early Cambrian trilobites from South China. In particular, R. rex has a tripartite exopodite, as well as a dorsoventrally deep protopodite with gnathobasic spines used to shred or crush food items. Based on recent phylogenetic analyses, it is possible that an exopodite with tripartite subdivisions represents the plesiomorphic condition for Artiopoda (trilobites and kin). The digestive system of R. takooensis exhibits a series of paired digestive glands in the cephalon and anterior thorax, similar to those described for a number of other Cambrian and Ordovician trilobites. [ABSTRACT FROM AUTHOR]

Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock being extremely abundant, and the larger Redlichia takooensis Lu , 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 characterized 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. [ABSTRACT FROM AUTHOR]

The University of Mississippi, Oxford, Mississippi is extremely fortunate in having a computer installation comprised of the IBM 630 computer and auxiliary equipment. A course in electronic data processing was established in the School of Commerce and Business Administration in February, 1959. The course is entitled Business Data Processing, and carries three semester hours credit. It is being offered at the junior level to any student having a minimum of nine hours of accounting. Since the majority of students in the School are required to take nine hours of accounting, no particular major field of study is precluded from participation in the course. Based on the assumption that not only accounting but other fields would be represented in the student enrollment, it appeared that the introductory portion of the course should be similar to the systems course required of all accounting majors. It is believed that the student will have a better background for the computer portion of the course if he has first been grounded in manual and punched card systems. Also, it will insure that all students will be exposed to an equal amount of material regarding manual and punched card systems.