The Emu Bay Shale: A unique early Cambrian Lagerstätte from a tectonically active basin.

The Emu Bay Shale (EBS) of South Australia is anomalous among Cambrian Lagerstätten because it captures anatomical information that is rare in Burgess Shale-type fossils, and because of its inferred nearshore setting, the nature of which has remained controversial. Intensive study, combining outcrop and borehole data with a compilation of >25,000 fossil specimens, reveals that the EBS biota inhabited a fan delta complex within a tectonically active basin. Preservation of soft-bodied organisms in this setting is unexpected and further underscores differences between the EBS and other Cambrian Lagerstätten. Environmental conditions, including oxygen fluctuations, slope instability, high suspended sediment concentrations, and episodic high-energy events, inhibited colonization of the lower prodelta by all but a few specialist species but favored downslope transportation and preservation of other largely endemic, shallow-water benthos. The EBS provides extraordinary insight into early Cambrian animal diversity from Gondwana. These results demonstrate how environmental factors determined community composition and provide a framework for understanding this unique Konservat-Lagerstätte.

Rapid volcanic ash entombment reveals the 3D anatomy of Cambrian trilobites.

Knowledge of Cambrian animal anatomy is limited by preservational processes that result in compaction, size bias, and incompleteness. We documented pristine three-dimensional (3D) anatomy of trilobites fossilized through rapid ash burial from a pyroclastic flow entering a shallow marine environment. Cambrian ellipsocephaloid trilobites from Morocco are articulated and undistorted, revealing exquisite details of the appendages and digestive system. Previously unknown anatomy includes a soft-tissue labrum attached to the hypostome, a slit-like mouth, and distinctive cephalic feeding appendages. Our findings resolve controversy over whether the trilobite hypostome is the labrum or incorporates it and establish crown-group euarthropod homologies in trilobites. This occurrence of moldic fossils with 3D soft parts highlights volcanic ash deposits in marine settings as an underexplored source for exceptionally preserved organisms.

Malformed trilobites from the Cambrian, Ordovician, and Silurian of Australia.

Biomineralised remains of trilobites provide important insight into the evolutionary history of a diverse, extinct group of arthropods. Their exoskeletons are also ideal for recording malformations, including evidence of post-injury repair. Re-examination of historic collections and the study of new specimens is important for enhancing knowledge on trilobite malformations across this diverse clade. To expand the records of these abnormalities and present explanations for their formation, we document eight malformed trilobite specimens, as well as one carcass, housed within the Commonwealth Palaeontological Collection at Geoscience Australia in Canberra. We present examples of Asthenopsis, Burminresia, Centropleura, Coronocephalus, Dolicholeptus, Galahetes, Papyriaspis, and Xystridura from Cambrian, Ordovician, and Silurian deposits of Australia. Most of the malformed specimens show W-, U-, or L-shaped indentations that reflect injuries from either failed predation or complications during moulting, and a mangled carcass is ascribed to either successful predation or post-mortem scavenging. We also uncover examples of teratologies, such as bifurcated pygidial ribs and pygidial asymmetry, in addition to evidence of abnormal recovery (i.e., fusion of thoracic segments) from a traumatic incident.

Hidden Markov movement models reveal diverse seasonal movement patterns in two North American ungulates.

Animal movement is the mechanism connecting landscapes to fitness, and understanding variation in seasonal animal movements has benefited from the analysis and categorization of animal displacement. However, seasonal movement patterns can defy classification when movements are highly variable. Hidden Markov movement models (HMMs) are a class of latent-state models well-suited to modeling movement data. Here, we used HMMs to assess seasonal patterns of variation in the movement of pronghorn (Antilocapra americana), a species known for variable seasonal movements that challenge analytical approaches, while using a population of mule deer (Odocoileus hemionus), for whom seasonal movements are well-documented, as a comparison. We used population-level HMMs in a Bayesian framework to estimate a seasonal trend in the daily probability of transitioning between a short-distance local movement state and a long-distance movement state. The estimated seasonal patterns of movements in mule deer closely aligned with prior work based on indices of animal displacement: a short period of long-distance movements in the fall season and again in the spring, consistent with migrations to and from seasonal ranges. We found seasonal movement patterns for pronghorn were more variable, as a period of long-distance movements in the fall was followed by a winter period in which pronghorn were much more likely to further initiate and remain in a long-distance movement pattern compared with the movement patterns of mule deer. Overall, pronghorn were simply more likely to be in a long-distance movement pattern throughout the year. Hidden Markov movement models provide inference on seasonal movements similar to other methods, while providing a robust framework to understand movement patterns on shorter timescales and for more challenging movement patterns. Hidden Markov movement models can allow a rigorous assessment of the drivers of changes in movement patterns such as extreme weather events and land development, important for management and conservation.

Raptorial appendages of the Cambrian apex predator Anomalocaris canadensis are built for soft prey and speed.

The stem-group euarthropod Anomalocaris canadensis is one of the largest Cambrian animals and is often considered the quintessential apex predator of its time. This radiodont is commonly interpreted as a demersal hunter, responsible for inflicting injuries seen in benthic trilobites. However, controversy surrounds the ability of A. canadensis to use its spinose frontal appendages to masticate or even manipulate biomineralized prey. Here, we apply a new integrative computational approach, combining three-dimensional digital modelling, kinematics, finite-element analysis (FEA) and computational fluid dynamics (CFD) to rigorously analyse an A. canadensis feeding appendage and test its morphofunctional limits. These models corroborate a raptorial function, but expose inconsistencies with a capacity for durophagy. In particular, FEA results show that certain parts of the appendage would have experienced high degrees of plastic deformation, especially at the endites, the points of impact with prey. The CFD results demonstrate that outstretched appendages produced low drag and hence represented the optimal orientation for speed, permitting acceleration bursts to capture prey. These data, when combined with evidence regarding the functional morphology of its oral cone, eyes, body flaps and tail fan, suggest that A. canadensis was an agile nektonic predator that fed on soft-bodied animals swimming in a well-lit water column above the benthos. The lifestyle of A. canadensis and that of other radiodonts, including plausible durophages, suggests that niche partitioning across this clade influenced the dynamics of Cambrian food webs, impacting on a diverse array of organisms at different sizes, tiers and trophic levels.

Biomechanical analyses of pterygotid sea scorpion chelicerae uncover predatory specialisation within eurypterids.

Eurypterids (sea scorpions) are extinct aquatic chelicerates. Within this group, members of Pterygotidae represent some of the largest known marine arthropods. Representatives of this family all have hypertrophied, anteriorly-directed chelicerae and are commonly considered Silurian and Devonian apex predators. Despite a long history of research interest in these appendages, pterygotids have been subject to limited biomechanical investigation. Here, we present finite element analysis (FEA) models of four different pterygotid chelicerae-those of Acutiramus bohemicus, Erettopterus bilobus, Jaekelopterus rhenaniae, and Pterygotus anglicus-informed through muscle data and finite element models (FEMs) of chelae from 16 extant scorpion taxa. We find that Er. bilobus and Pt. anglicus have comparable stress patterns to modern scorpions, suggesting a generalised diet that probably included other eurypterids and, in the Devonian species, armoured fishes, as indicated by co-occurring fauna. Acutiramus bohemicus is markedly different, with the stress being concentrated in the proximal free ramus and the serrated denticles. This indicates a morphology better suited for targeting softer prey. Jaekelopterus rhenaniae exhibits much lower stress across the entire model. This, combined with an extremely large body size, suggests that the species likely fed on larger and harder prey, including heavily armoured fishes. The range of cheliceral morphologies and stress patterns within Pterygotidae demonstrate that members of this family had variable diets, with only the most derived species likely to feed on armoured prey, such as placoderms. Indeed, increased sizes of these forms throughout the mid-Palaeozoic may represent an 'arms race' between eurypterids and armoured fishes, with Devonian pterygotids adapting to the rapid diversification of placoderms.

Evaluating the summer landscapes of predation risk and forage quality for elk (Cervus canadensis).

The recovery of carnivore populations in North American has consequences for trophic interactions and population dynamics of prey. In addition to direct effects on prey populations through killing, predators can influence prey behavior by imposing the risk of predation. The mechanisms through which patterns of space use by predators are linked to behavioral response by prey and nonconsumptive effects on prey population dynamics are poorly understood. Our goal was to characterize population- and individual-level patterns of resource selection by elk (Cervus canadensis) in response to risk of wolves (Canis lupus) and mountain lions (Puma concolor) and evaluate potential nonconsumptive effects of these behavioral patterns. We tested the hypothesis that individual elk risk-avoidance behavior during summer would result in exposure to lower-quality forage and reduced body fat and pregnancy rates. First, we evaluated individuals' second-order and third-order resource selection with a used-available sampling design. At the population level, we found evidence for a positive relationship between second- and third-order selection and forage, and an interaction between forage quality and mountain lion risk such that the relative probability of use at low mountain lion risk increased with forage quality but decreased at high risk at both orders of selection. We found no evidence of a population-level trade-off between forage quality and wolf risk. However, we found substantial among-individual heterogeneity in resource selection patterns such that population-level patterns were potentially misleading. We found no evidence that the diversity of individual resource selection patterns varied predictably with available resources, or that patterns of individual risk-related resource selection translated into biologically meaningful changes in body fat or pregnancy rates. Our work highlights the importance of evaluating individual responses to predation risk and predator hunting technique when assessing responses to predators and suggests nonconsumptive effects are not operating at a population scale in this system.

Complex axial growth patterns in an early Cambrian trilobite from South Australia.

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.

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy.

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.

Disparate compound eyes of Cambrian radiodonts reveal their developmental growth mode and diverse visual ecology.

Radiodonts are nektonic stem-group euarthropods that played various trophic roles in Paleozoic marine ecosystems, but information on their vision is limited. Optical details exist only in one species from the Cambrian Emu Bay Shale of Australia, here assigned to Anomalocaris aff. canadensis We identify another type of radiodont compound eye from this deposit, belonging to 'Anomalocaris' briggsi This ≤4-cm sessile eye has >13,000 lenses and a dorsally oriented acute zone. In both taxa, lenses were added marginally and increased in size and number throughout development, as in many crown-group euarthropods. Both species' eyes conform to their inferred lifestyles: The macrophagous predator A. aff. canadensis has acute stalked eyes (>24,000 lenses each) adapted for hunting in well-lit waters, whereas the suspension-feeding 'A.' briggsi could detect plankton in dim down-welling light. Radiodont eyes further demonstrate the group's anatomical and ecological diversity and reinforce the crucial role of vision in early animal ecosystems.

Active surveillance of low-risk papillary thyroid cancer: A meta-analysis.

BACKGROUND: This study evaluates the safety and efficacy of active surveillance for low-risk papillary thyroid carcinoma. METHODS: MEDLINE, EMBASE, and PubMed were searched from inception for relevant studies of active surveillance for low-risk papillary thyroid carcinoma, defined as T1a or T1b, N0, M0 disease. Main outcomes of interest were growth of primary tumor, metastatic spread, thyroid cancer-related mortality, and disease recurrence after delayed thyroid surgery. RESULTS: Nine publications with 4,156 patients were included. Primary analysis of the 9 studies revealed pooled proportion of tumor growth during active surveillance to be 4.4% (95% confidence interval 3.2-5.8%). The pooled rate of metastatic spread to cervical nodes was 1.0% (95% confidence interval 0.7-1.4%), and pooled mortality due to thyroid cancer was 0.03% (95% confidence interval 0.0005-0.2%). Eight studies assessed incidence of delayed thyroid surgery with pooled proportion of 9.9% (95% confidence interval 6.4-14.0%). The main indication for surgery was patient preference, not disease progression, at 51.9% (95% confidence interval 44.9-58.9%). The pooled proportion of recurrence after delayed thyroid surgery was 1.1% (95% confidence interval 0.1-3.8%). CONCLUSION: Active surveillance appears to be a safe alternative to surgery for the management of low-risk papillary thyroid carcinoma, without increased risk of recurrence or death. This strategy allows for avoidance of exposure to surgical risk and need for subsequent thyroid replacement therapy.

Incremental healthcare utilisation and costs among new senior high-cost users in Ontario, Canada: a retrospective matched cohort study.

OBJECTIVES: To describe healthcare use and spending before and on becoming a new (incident) senior high-cost user (HCU) compared with senior non-HCUs; to estimate the incremental costs, overall and by service category, attributable to HCU status; and to quantify its monetary impact on the provincial healthcare budget in Ontario, Canada. DESIGN: We conducted a retrospective, population-based comparative cohort study using administrative healthcare records. Incremental healthcare utilisation and costs were determined using the method of recycled predictions allowing adjustment for preincident and incident year values, and covariates. Estimated budget impact was computed as the product of the mean annual total incremental cost and the number of senior HCUs. PARTICIPANTS: Incident senior HCUs were defined as Ontarians aged ≥66 years who were in the top 5% of healthcare cost users during fiscal year 2013 (FY2013) but not during FY2012. The incident HCU cohort was matched with senior non-HCUs in a ratio of 1 HCU:3 non-HCU. RESULTS: Senior HCUs (n=175 847) reached the annual HCU threshold of CAD$10 192 through different combinations of incurred costs. Although HCUs had higher healthcare utilisation and costs at baseline, HCU status was associated with a substantial spike in both, with prolonged hospitalisations playing a major role. Twelve per cent of HCUs reached the HCU expenditure threshold without hospitalisation. Compared with non-HCUs (n=5 27 541), HCUs incurred an additional CAD$25 527 per patient in total healthcare costs; collectively CAD$4.5 billion or 9% of the 2013 Ontario healthcare budget. Inpatient care had the highest incremental costs: CAD$13 427, 53% of the total incremental spending. CONCLUSIONS: Costs attributable to incident senior HCU status accounted for almost 1/10 of the provincial healthcare budget. Prolonged hospitalisations made a major contribution to the total incremental costs. A subgroup of patients that became HCU without hospitalisation requires further investigation.

Efficacy of 177Lu Peptide Receptor Radionuclide Therapy for the Treatment of Neuroendocrine Tumors: A Meta-analysis.

OBJECTIVE: The purpose of this study was to assess the efficacy of Lu-labeled peptide receptor radionuclide therapy (PRRT) induction treatments for patients with unresectable metastatic neuroendocrine tumors. METHODS: MEDLINE, EMBASE, and Ovid were systematically searched with keywords "lutetium," "Lu-177," "PRRT," "neuroendocrine," and "prognosis." Studies evaluating treatment with Lu-labeled PRRT were assessed for disease response and/or disease control rate by Response Evaluation Criteria in Solid Tumors (RECIST) 1.0 or 1.1, modified RECIST, Southwest Oncology Group (SWOG), or modified SWOG criteria. Pooled proportions of disease response and control rates were calculated for both fixed- and random-effects models. RESULTS: Eighteen studies with 1920 patients were included (11 with 1268 patients using RECIST and 6 with 804 patients using SWOG). By RECIST criteria, the pooled disease response rate by random-effects model was 29.1% (95% confidence interval [CI], 20.2%-38.9%), and disease control rate was 74.1% (95% CI, 67.8%-80.0%). By SWOG criteria, the pooled disease response rate by random-effects model was 30.6% (95% CI, 20.7%-41.5%), and disease control rate was 81.1% (95% CI, 76.4%-85.4%). CONCLUSIONS: Induction therapy, typically 4 treatments, with Lu PRRT is an effective method of treating unresectable metastatic neuroendocrine tumors with significant disease response and control rates.

Trilobite evolutionary rates constrain the duration of the Cambrian explosion.

Trilobites are often considered exemplary for understanding the Cambrian explosion of animal life, due to their unsurpassed diversity and abundance. These biomineralized arthropods appear abruptly in the fossil record with an established diversity, phylogenetic disparity, and provincialism at the beginning of Cambrian Series 2 (∼521 Ma), suggesting a protracted but cryptic earlier history that possibly extends into the Precambrian. However, recent analyses indicate elevated rates of phenotypic and genomic evolution for arthropods during the early Cambrian, thereby shortening the phylogenetic fuse. Furthermore, comparatively little research has been devoted to understanding the duration of the Cambrian explosion, after which normal Phanerozoic evolutionary rates were established. We test these hypotheses by applying Bayesian tip-dating methods to a comprehensive dataset of Cambrian trilobites. We show that trilobites have a Cambrian origin, as supported by the trace fossil record and molecular clocks. Surprisingly, they exhibit constant evolutionary rates across the entire Cambrian, for all aspects of the preserved phenotype: discrete, meristic, and continuous morphological traits. Our data therefore provide robust, quantitative evidence that by the time the typical Cambrian fossil record begins (∼521 Ma), the Cambrian explosion had already largely concluded. This suggests that a modern-style marine biosphere had rapidly emerged during the latest Ediacaran and earliest Cambrian (∼20 million years), followed by broad-scale evolutionary stasis throughout the remainder of the Cambrian.

Incidence of new-onset diabetes mellitus and association with mortality in childhood solid organ transplant recipients: a population-based study.

BACKGROUND: Precise estimates of the long-term risk of new-onset diabetes and its impact on mortality among transplanted children are not known. METHODS: We conducted a cohort study comparing children undergoing solid organ (kidney, heart, liver, lung and multiple organ) transplant (n = 1020) between 1991 and 2014 with healthy non-transplanted children (n = 7 134 067) using Ontario health administrative data. Outcomes included incidence of diabetes among transplanted and non-transplanted children, the relative hazard of diabetes among solid organ transplant recipients, overall and at specific intervals posttransplant, and mortality among diabetic transplant recipients. RESULTS: During 56 019 824 person-years of follow-up, the incidence rate of diabetes was 17.8 [95% confidence interval (CI) 15-21] and 2.5 (95% CI 2.5-2.5) per 1000 person-years among transplanted and non-transplanted children, respectively. The transplant cohort had a 9-fold [hazard ratio (HR) 8.9; 95% CI 7.5-10.5] higher hazard of diabetes compared with those not transplanted. Risk was highest within the first year after transplant (HR 20.7; 95% CI 15.9-27.1), and remained elevated even at 5 and 10 years of follow-up. Lung and multiple organ recipients had a 5-fold (HR 5.4; 95% CI 3.0-9.8) higher hazard of developing diabetes compared with kidney transplant recipients. Transplant recipients with diabetes had a three times higher hazard of death compared with those who did not develop diabetes (HR 3.3; 95% CI 2.3-4.8). CONCLUSIONS: The elevated risk of diabetes in transplant recipients persists even after a decade, highlighting the importance of ongoing surveillance. Diabetes after transplantation increases the risk of mortality among childhood transplant recipients.

Computational biomechanical analyses demonstrate similar shell-crushing abilities in modern and ancient arthropods.

The biology of the American horseshoe crab, Limulus polyphemus, is well documented-including its dietary habits, particularly the ability to crush shell with gnathobasic walking appendages-but virtually nothing is known about the feeding biomechanics of this iconic arthropod. Limulus polyphemus is also considered the archetypal functional analogue of various extinct groups with serial gnathobasic appendages, including eurypterids, trilobites and other early arthropods, especially Sidneyia inexpectans from the mid-Cambrian (508 Myr) Burgess Shale of Canada. Exceptionally preserved specimens of S. inexpectans show evidence suggestive of durophagous (shell-crushing) tendencies-including thick gnathobasic spine cuticle and shelly gut contents-but the masticatory capabilities of this fossil species have yet to be compared with modern durophagous arthropods. Here, we use advanced computational techniques, specifically a unique application of 3D finite-element analysis (FEA), to model the feeding mechanics of L. polyphemus and S. inexpectans: the first such analyses of a modern horseshoe crab and a fossil arthropod. Results show that mechanical performance of the feeding appendages in both arthropods is remarkably similar, suggesting that S. inexpectans had similar shell-crushing capabilities to L. polyphemus This biomechanical solution to processing shelly food therefore has a history extending over 500 Myr, arising soon after the first shell-bearing animals. Arrival of durophagous predators during the early phase of animal evolution undoubtedly fuelled the Cambrian 'arms race' that involved a rapid increase in diversity, disparity and abundance of biomineralized prey species.

Trilobites and agnostids from the Goyder Formation (Cambrian Series 3, Guzhangian; Mindyallan), Amadeus Basin, central Australia.

A new assemblage containing twenty-two species of trilobites and agnostids is described from the Goyder Formation (Cambrian Series 3) in the Ross River Syncline and Gardiner Ranges of the Amadeus Basin, Northern Territory, central Australia. New trilobite taxa described include the genus, Trephina gen. nov., and four new species Adelogonus prichardi sp. nov., Hebeia stewarti sp. nov., Liostracina joyceae sp. nov., and Trephina ranfordi sp. nov. Two agnostid taxa previously known only from Antarctica, Ammagnostus antarcticus Bentley, Jago Cooper, 2009 and Hadragnostus helixensis Jago Cooper, 2005, are also documented. Of the two agnostid species, the latter is the most age diagnostic, previously reported from the Cambrian Series 3 (Guzhangian; late Mindyallan; Glyptagnostus stolidotus Zone) Spurs Formation in Northern Victoria Land. This age for the Goyder Formation assemblage is supported by the co-occurrence of the trilobites Biaverta reineri Öpik, 1967, Blackwelderia repanda Öpik, 1967, Henadoparia integra Öpik, 1967, Monkaspis cf. travesi (Öpik, 1967), Nomadinis pristinus Öpik, 1967, Paraacidaspis? priscilla (Öpik, 1967), and Polycyrtaspis cf. flexuosa Öpik, 1967, also known from the late Mindyallan (G. stolidotus Zone) successions of the neighbouring Georgina Basin (Northern Territory and Queensland). The generic assemblage of the Goyder Formation is also similar to those from the Guzhangian (Mindyallan) of other parts of Australia (New South Wales, South Australia, and Western Australia), in addition to East Antarctica and North and South China.

A 3D anatomical atlas of appendage musculature in the chelicerate arthropod Limulus polyphemus.

Limulus polyphemus, an archetypal chelicerate taxon, has interested both biological and paleontological researchers due to its unique suite of anatomical features and as a useful modern analogue for fossil arthropod groups. To assist the study and documentation of this iconic taxon, we present a 3D atlas on the appendage musculature, with specific focus on the muscles of the cephalothoracic appendages. As L. polyphemus appendage musculature has been the focus of extensive study, depicting the muscles in 3D will facilitate a more complete understanding thereof for future researchers. A large museum specimen was CT scanned to illustrate the major exoskeletal features of L. polyphemus. Micro-CT scans of iodine-stained appendages from fresh, non-museum specimens were digitally dissected to interactively depict appendage sections and muscles. This study has revealed the presence of two new muscles: one within the pushing leg, located dorsally relative to all other patella muscles, and the other within the male pedipalp, located in the modified tibiotarsus. This atlas increases accessibility to important internal and external morphological features of L. polyphemus and reduces the need for destructive fresh tissue dissection of specimens. Scanning, digitally dissecting, and documenting taxa in 3D is a pivotal step towards creating permanent digital records of life on Earth.

Reappraising the early evidence of durophagy and drilling predation in the fossil record: implications for escalation and the Cambrian Explosion.

The Cambrian Explosion is arguably the most extreme example of a biological radiation preserved in the fossil record, and studies of Cambrian Lagerstätten have facilitated the exploration of many facets of this key evolutionary event. As predation was a major ecological driver behind the Explosion - particularly the radiation of biomineralising metazoans - the evidence for shell crushing (durophagy), drilling and puncturing predation in the Cambrian (and possibly the Ediacaran) is considered. Examples of durophagous predation on biomineralised taxa other than trilobites are apparently rare, reflecting predator preference, taphonomic and sampling biases, or simply lack of documentation. The oldest known example of durophagy is shell damage on the problematic taxon Mobergella holsti from the early Cambrian (possibly Terreneuvian) of Sweden. Using functional morphology to identify (or perhaps misidentify) durophagous predators is discussed, with emphasis on the toolkit used by Cambrian arthropods, specifically the radiodontan oral cone and the frontal and gnathobasic appendages of various taxa. Records of drill holes and possible puncture holes in Cambrian shells are mostly on brachiopods, but the lack of prey diversity may represent either a true biological signal or a result of various biases. The oldest drilled Cambrian shells occur in a variety of Terreneuvian-aged taxa, but specimens of the ubiquitous Ediacaran shelly fossil Cloudina also show putative drilling traces. Knowledge on Cambrian shell drillers is sorely lacking and there is little evidence or consensus concerning the taxonomic groups that made the holes, which often leads to the suggestion of an unknown 'soft bodied driller'. Useful methodologies for deciphering the identities and capabilities of shell drillers are outlined. Evidence for puncture holes in Cambrian shelly taxa is rare. Such holes are more jagged than drill holes and possibly made by a Cambrian 'puncher'. The Cambrian arthropod Yohoia may have used its frontal appendages in a jack-knifing manner, similar to Recent stomatopod crustaceans, to strike and puncture shells rapidly. Finally, Cambrian durophagous and shell-drilling predation is considered in the context of escalation - an evolutionary process that, amongst other scenarios, involves predators (and other 'enemies') as the predominant agents of natural selection. The rapid increase in diversity and abundance of biomineralised shells during the early Cambrian is often attributed to escalation: enemies placed selective pressure on prey, forcing phenotypic responses in prey and, by extension, in predator groups over time. Unfortunately, few case studies illustrate long-term patterns in shelly fossil morphologies that may reflect the influence of predation throughout the Cambrian. More studies on phenotypic change in hard-shelled lineages are needed to convincingly illustrate escalation and the responses of prey during the Cambrian.

The gnathobasic spine microstructure of recent and Silurian chelicerates and the Cambrian artiopodan Sidneyia: Functional and evolutionary implications.

Gnathobasic spines are located on the protopodal segments of the appendages of various euarthropod taxa, notably chelicerates. Although they are used to crush shells and masticate soft food items, the microstructure of these spines are relatively poorly known in both extant and extinct forms. Here we compare the gnathobasic spine microstructures of the Silurian eurypterid Eurypterus tetragonophthalmus from Estonia and the Cambrian artiopodan Sidneyiainexpectans from Canada with those of the Recent xiphosuran chelicerate Limulus polyphemus to infer potential variations in functional morphology through time. The thickened fibrous exocuticle in L. polyphemus spine tips enables effective prey mastication and shell crushing, while also reducing pressure on nerve endings that fill the spine cavities. The spine cuticle of E. tetragonophthalmus has a laminate structure and lacks the fibrous layers seen in L. polyphemus spines, suggesting that E. tetragonophthalmus may not have been capable of crushing thick shells, but a durophagous habit cannot be precluded. Conversely, the cuticle of S. inexpectans spines has a similar fibrous microstructure to L. polyphemus, suggesting that S. inexpectans was a competent shell crusher. This conclusion is consistent with specimens showing preserved gut contents containing various shelly fragments. The shape and arrangement of the gnathobasic spines is similar for both L. polyphemus and S. inexpectans, with stouter spines in the posterior cephalothoracic or trunk appendages, respectively. This differentiation indicates that crushing occurs posteriorly, while the gnathobases on anterior appendages continue mastication and push food towards and into the mouth. The results of recent phylogenetic analyses that considered both modern and fossil euarthropod clades show that xiphosurans and eurypterids are united as crown-group euchelicerates, with S. inexpectans placed within more basal artiopodan clades. These relationships suggest that gnathobases with thickened fibrous exocuticle, if not homoplasious, may be plesiomorphic for chelicerates and deeper relatives within Arachnomorpha. This study shows that the gnathobasic spine microstructure best adapted for durophagy has remained remarkably constant since the Cambrian.