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Fossils preserving soft tissues and lightly biomineralized structures are essential for the reconstruction of past ecosystems and their evolution. Understanding fossilization processes, including decay and mineralisation, is crucial for accurately interpreting ancient morphologies. Here we investigate the decay of marine and freshwater shrimps deposited on the surface of three different clay beds. In experimental set ups containing kaolinite, cryogenic scanning electron microscopy shows a black film comprised of newly formed anhedral and cryptocrystalline aluminosilicates on marine shrimp cuticles, which stabilise the overall morphology. This is the first experimental evidence for the replication of arthropod lightly biomineralized structures in aluminosilicates shortly after death, while carcasses are not buried by sediments. The preservation of morphology through aluminosilicates could result in carcasses persisting on the seafloor for weeks without losing much external anatomical information. In this context, instantaneous burial capturing animals alive may not be a prerequisite for exceptional preservation as usually thought., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2025.)

Exoskeletons are a defining character of all arthropods that provide physical support for their segmented bodies and appendages as well as protection from the environment and predation. This ubiquitous yet evolutionarily variable feature has been instrumental in facilitating the adoption of a variety of lifestyles and the exploitation of ecological niches across all environments. Throughout the radiation that produced the more than one million described modern species, adaptability afforded by segmentation and exoskeletons has led to a diversity that is unrivalled amongst animals. However, because of the limited extensibility of exoskeleton chitin and cuticle components, they must be periodically shed and replaced with new larger ones, notably to accommodate the growing individuals encased within. Therefore, arthropods grow discontinuously by undergoing periodic moulting events, which follow a series of steps from the preparatory pre-moult phase to ecdysis itself and post-moult maturation of new exoskeletons. Each event represents a particularly vulnerable period in an arthropod's life cycle, so processes must be tightly regulated and meticulously executed to ensure successful transitions for normal growth and development. Decades of research in representative arthropods provide a foundation of understanding of the mechanisms involved. Building on this, studies continue to develop and test hypotheses on the presence and function of molecular components, including neuropeptides, hormones, and receptors, as well as the so-called early, late, and fate genes, across arthropod diversity. Here, we review the literature to develop a comprehensive overview of the status of accumulated knowledge of the genetic toolkit governing arthropod moulting. From biosynthesis and regulation of ecdysteroid and sesquiterpenoid hormones, to factors involved in hormonal stimulation responses and exoskeleton remodelling, we identify commonalities and differences, as well as highlighting major knowledge gaps, across arthropod groups. We examine the available evidence supporting current models of how components operate together to prepare for, execute, and recover from ecdysis, comparing reports from Chelicerata, Myriapoda, Crustacea, and Hexapoda. Evidence is generally highly taxonomically imbalanced, with most reports based on insect study systems. Biases are also evident in research on different moulting phases and processes, with the early triggers and late effectors generally being the least well explored. Our synthesis contrasts knowledge based on reported observations with reasonably plausible assumptions given current taxonomic sampling, and exposes weak assumptions or major gaps that need addressing. Encouragingly, advances in genomics are driving a diversification of tractable study systems by facilitating the cataloguing of putative genetic toolkits in previously under-explored taxa. Analysis of genome and transcriptome data supported by experimental investigations have validated the presence of an "ultra-conserved" core of arthropod genes involved in moulting processes. The molecular machinery has likely evolved with elaborations on this conserved pathway backbone, but more taxonomic exploration is needed to characterise lineage-specific changes and novelties. Furthermore, linking these to transformative innovations in moulting processes across Arthropoda remains hampered by knowledge gaps and hypotheses based on untested assumptions. Promisingly however, emerging from the synthesis is a framework that highlights research avenues from the underlying genetics to the dynamic molecular biology through to the complex physiology of moulting., (© 2024 The Author(s). Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

The rapid early diversification of arthropods has made understanding internal relationships within the group fiendish. Particularly unresolved is the origin of Euchelicerata, a clade consisting of the Prosomapoda (comprising the extant Xiphosura and Arachnida and the extinct Chasmataspidida, Eurypterida and synziphosurines) and the extinct Offacolidae. Here we describe new material of the Silurian 'synziphosurine' Bunaia woodwardi Clarke, 1919 that reveals previously unknown features of its ventral anatomy: a pair of elongated chelicerae in the prosoma, followed posteriorly by five pairs of biramous appendages, a first pre-abdomen somite bearing a pair of paddle-like uniramous appendages (exopods) and a ventral pretelsonic process. Phylogenetic analyses retrieve B. woodwardi as a member of Offacolidae closely related to Setapedites abundantis from the Early Ordovician Fezouata Biota. An anatomical comparison of the pretelsonic process of B. woodwardi , also present in Setapedites , with the posterior trunk morphologies of other Offacolidae, Habeliida and Vicissicaudata, suggests a possible homologous appendicular origin. This proposed apomorphic character supports a monophyletic Arachnomorpha, formed of Vicissicaudata, Habeliida and Euchelicerata. The establishment of this new homology could help to clarify the highly enigmatic phylogeny at the base of the euchelicerates as well as the sequence of character acquisition during their early evolution., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

The Fezouata Biota (Morocco) is a Burgess Shale-type (BST) assemblage that provides a wealth of information on Early Ordovician ecosystems. Much work has been done to compare the preservation of the Fezouata Biota to other BSTs. However, studies investigating preservation variations within the Fezouata Biota are rare. Here, we use probabilities to investigate the preservation of various ecological categories of Fezouata eumetazoans. Complex taphonomic processes and phylum-specific constraints have led to the better preservation of predators/scavengers in this biota. However, no differences in preservation are observed between vagile and sessile taxa. Importantly, Tremadocian taxa are better preserved than Floian ones. As such, this study highlights the gradual closure of the BST window of preservation in the Zagora region of Morocco and constitutes a benchmark for future palaeoecological and evolutionary studies on the Fezouata Biota., (© 2024. The Author(s).)

Euchelicerata is a clade of arthropods comprising horseshoe crabs, scorpions, spiders, mites and ticks, as well as the extinct eurypterids (sea scorpions) and chasmataspidids. The understanding of the ground plans and relationships between these crown-group euchelicerates has benefited from the discovery of numerous fossils. However, little is known regarding the origin and early evolution of the euchelicerate body plan because the relationships between their Cambrian sister taxa and synziphosurines, a group of Silurian to Carboniferous stem euchelicerates with chelicerae and an unfused opisthosoma, remain poorly understood owing to the scarce fossil record of appendages. Here we describe a synziphosurine from the Lower Ordovician (ca. 478 Ma) Fezouata Shale of Morocco. This species possesses five biramous appendages with stenopodous exopods bearing setae in the prosoma and a fully expressed first tergite in the opisthosoma illuminating the ancestral anatomy of the group. Phylogenetic analyses recover this fossil as a member of the stem euchelicerate family Offacolidae, which is characterized by biramous prosomal appendages. Moreover, it also shares anatomical features with the Cambrian euarthropod Habelia optata, filling the anatomical gap between euchelicerates and Cambrian stem taxa, while also contributing to our understanding of the evolution of euchelicerate uniramous prosomal appendages and tagmosis., (© 2024. The Author(s).)

Early Palaeozoic sites with soft-tissue preservation are predominantly found in Cambrian rocks and tend to capture past tropical and temperate ecosystems. In this study, we describe the diversity and preservation of the Cabrières Biota, a newly discovered Early Ordovician Lagerstätte from Montagne Noire, southern France. The Cabrières Biota showcases a diverse polar assemblage of both biomineralized and soft-bodied organisms predominantly preserved in iron oxides. Echinoderms are extremely scarce, while sponges and algae are abundantly represented. Non-biomineralized arthropod fragments are also preserved, along with faunal elements reminiscent of Cambrian Burgess Shale-type ecosystems, such as armoured lobopodians. The taxonomic diversity observed in the Cabrières Biota mixes Early Ordovician Lagerstätten taxa with Cambrian forms. By potentially being the closest Lagerstätte to the South Pole, the Cabrières Biota probably served as a biotic refuge amid the high-water temperatures of the Early Ordovician, and shows comparable ecological structuring to modern polar communities., (© 2024. The Author(s).)

Fossilization, or the transition of an organism from the biosphere to the geosphere, is a complex mechanism involving numerous biological and geological variables. Bacteria are one of the most significant biotic players to decompose organic matter in natural environments, early on during fossilization. However, bacterial processes are difficult to characterize as many different abiotic conditions can influence bacterial efficiency in degrading tissues. One potentially important variable is the composition and nature of the sediment on which a carcass is deposited after death. We experimentally examined this by decaying the marine shrimp Palaemon varians underwater on three different clay sediments. Samples were then analyzed using 16S ribosomal RNA sequencing to identify the bacterial communities associated with each clay system. Results show that samples decaying on the surface of kaolinite have a lower bacterial diversity than those decaying on the surface of bentonite and montmorillonite, which could explain the limited decay of carcasses deposited on this clay. However, this is not the only role played by kaolinite, as a greater proportion of gram-negative over gram-positive bacteria is observed in this system. Gram-positive bacteria are generally thought to be more efficient at recycling complex polysaccharides such as those forming the body walls of arthropods. This is the first experimental evidence of sediments shaping an entire bacterial community. Such interaction between sediments and bacteria might have contributed to arthropods' exquisite preservation and prevalence in kaolinite-rich Lagerstätten of the Cambrian Explosion., Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-024-00324-7., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2024.)

Cheiruridae is one of the most diverse families of trilobites known from the Ordovician with 453 species assigned. Within Cheiruridae eight subfamilies (Acanthoparyphinae, Cheirurinae, "Cyrtometopinae", Deiphoninae, Eccoptochilinae, Heliomerinae, Pilekiinae, and Sphaerexochinae) have historically been recognised. Insights about the evolution of the family and the relationships within and between subfamilies have been published. However larger scale phylogenetic hypotheses are needed in order to explore the monophyly, the basal structure, the deep nodes and the relationships of the subfamilies. Cheirurinae, Deiphoninae and "Cyrtometopinae" have historically been defined by various morphological features (e.g., anteroposterior constriction of the thoracic pleura, pleural furrow morphology, pygidial morphology) that differentiate them from the rest of Cheiruridae. However, the phylogenetic status of "Cyrtometopinae" is unclear owing to a lack of obvious synapomorphies. Here, we present phylogenetic analyses of Cheirurinae, Deiphoninae, and "Cyrtometopinae". The results indicate that both Cheirurinae and Deiphoninae are monophyletic. "Cyrtometopines" are resolved as a paraphyletic grade at the base of Deiphoninae and Cyrtometopinae should be considered a junior subjective synonym of Deiphoninae. The new phylogenetic hypothesis reveals that paedomorphosis plays an important role in the evolution of Deiphoninae. Within Cheirurinae two major clades are identified, the ' Ceraurus -like' clade and the ' Ceraurinella -like' clade., Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-024-00338-1., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2024.)

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.

Concretions are an interesting mode of preservation that can occasionally yield fossils with soft tissues. To properly interpret these fossils, an understanding of their fossilization is required. Probabilistic models are useful tools to identify variations between different Konservat-Lagerstätten that are separated spatially and temporally. However, the application of probabilistic modeling has been limited to Early Paleozoic Konservat-Lagerstätten preserved in shales. In this paper, the patterns of preservation of three concretionary Konservat-Lagerstätten-the Carboniferous Mazon Creek (USA) and Montceau-les-Mines (France), and the Silurian Herefordshire Lagerstätte (UK)-are analyzed using a statistical approach. It is demonstrated that the degree of biotic involvement, i.e., the degree to which a carcass dictates its own preservation, is connected to internal organ conditional probabilities-the probabilities of finding an internal organ associated with another structure such as biomineralized, sclerotized, cuticularized, or cellular body walls. In concretions that are externally forced with little biological mediation (e.g., Herefordshire), all internal organ conditional probabilities are uniform. As biological mediation in concretion formation becomes more pronounced, heterogeneities in conditional probabilities are introduced (e.g., Montceau-les-Mines and Mazon Creek). The three concretionary sites were also compared with previously investigated Konservat-Lagerstätten preserving fossils in shales to demonstrate how the developed probability framework aids in understanding the broad-scale functioning of preservation in Konservat-Lagerstätten., Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00284-4., Competing Interests: Competing interestsThe authors declare no conflict of interests., (© Akademie der Naturwissenschaften Schweiz (SCNAT) 2023.)

The Fezouata Biota (Morocco) is a unique Early Ordovician fossil assemblage. The discovery of this biota revolutionized our understanding of Earth's early animal diversifications-the Cambrian Explosion and the Ordovician Radiation-by suggesting an evolutionary continuum between both events. Herein, we describe Taichoute, a new fossil locality from the Fezouata Shale. This locality extends the temporal distribution of fossil preservation from this formation into the upper Floian, while also expanding the range of depositional environments to more distal parts of the shelf. In Taichoute, most animals were transported by density flows, unlike the in-situ preservation of animals recovered in previously investigated Fezouata sites. Taichoute is dominated by three-dimensionally preserved, and heavily sclerotized fragments of large euarthropods-possibly representing nektobenthic/nektic bivalved taxa and/or hurdiid radiodonts. Resolving whether this dominance reflects a legitimate aspect of the original ecosystem or a preservational bias requires an in-depth assessment of the environmental conditions at this site. Nevertheless, Taichoute provides novel preservational and palaeontological insights during a key evolutionary transition in the history of life on Earth., (© 2022. The Author(s).)

Biramous appendages are a common feature among modern marine arthropods that evolved deep in arthropod phylogeny. The branched appendage of Cambrian arthropods has long been considered as the ancient biramous limb, sparking numerous investigations on its origin and evolution. Here, we report a new arthropod, Erratus sperare gen. et sp. nov., from the Lower Cambrian (Stage 3, 520 Ma) Chengjiang biota of Yunnan, China, with unique trunk appendages formed of lateral anomalocaridid-type flaps and ventral subconical endopods. These appendages represent an intermediate stage of biramous limb evolution, i.e. from 'two pairs of flap appendages' in radiodonts to 'flap + endopod' in Erratus , to 'exopod + endopod' in the rest of carapace-bearing arthropods that populate the basal region of the upper-stem lineage arthropods (deuteropods). The new species occupies a phylogenetic position at the first node closer to deuteropods than to radiodonts, and therefore pinpoints the earliest occurrence of the endopod within Deuteropoda. The primitive endopod is weakly sclerotized, and has unspecialized segments without endites or claw. The findings might support previous claims that the outer branch of the biramous limb of fossil marine arthropods, such as trilobites, is not a true exopod, but is instead a modified exite. This article is part of the theme issue 'The impact of Chinese palaeontology on evolutionary research'.

The Early Ordovician is a key interval for our understanding of the evolution of life on Earth as it lays at the transition between the Cambrian Explosion and the Ordovician Radiation and because the fossil record of the late Cambrian is scarce. In this study, assembly processes of Early Ordovician trilobite and echinoderm communities from the Central Anti-Atlas (Morocco), the Montagne Noire (France), and the Cordillera Oriental (Argentina) are explored. The results show that dispersal increased diachronically in trilobite communities during the Early Ordovician. Dispersal did not increase for echinoderms. Dispersal was most probably proximally triggered by the planktic revolution, the fall in seawater temperatures, changes in oceanic circulation, with an overall control by tectonic frameworks and phylogenetic constraints. The diachronous increase in dispersal within trilobite communities in the Early Ordovician highlights the complexity of ecosystem structuring during the early stages of the Ordovician Radiation. As Early Ordovician regional dispersal was followed by well-documented continental dispersal in the Middle/Late Ordovician, it is possible to consider that alongside a global increase in taxonomic richness, the Ordovician Radiation is also characterized by a gradual increase in dispersal., (© 2022. The Author(s).)

Once considered 'weird wonders' of the Cambrian, the emblematic Burgess Shale animals Anomalocaris and Opabinia are now recognized as lower stem-group euarthropods and have provided crucial data for constraining the polarity of key morphological characters in the group. Anomalocaris and its relatives (radiodonts) had worldwide distribution and survived until at least the Devonian. However, despite intense study, Opabinia remains the only formally described opabiniid to date. Here we reinterpret a fossil from the Wheeler Formation of Utah as a new opabiniid, Utaurora comosa nov. gen. et sp. By visualizing the sample of phylogenetic topologies in treespace, our results fortify support for the position of U. comosa beyond the nodal support traditionally applied. Our phylogenetic evidence expands opabiniids to multiple Cambrian stages. Our results underscore the power of treespace visualization for resolving imperfectly preserved fossils and expanding the known diversity and spatio-temporal ranges within the euarthropod lower stem group.

The biological pump is crucial for transporting nutrients fixed by surface-dwelling primary producers to demersal animal communities. Indeed, the establishment of an efficient biological pump was likely a key factor enabling the diversification of animals over 500 Myr ago during the Cambrian explosion. The modern biological pump operates through two main vectors: the passive sinking of aggregates of organic matter, and the active vertical migration of animals. The coevolution of eukaryotes and sinking aggregates is well understood for the Proterozoic and Cambrian; however, little attention has been paid to the establishment of the vertical migration of animals. Here we investigate the morphological variation and hydrodynamic performance of the Cambrian euarthropod Isoxys . We combine elliptical Fourier analysis of carapace shape with computational fluid dynamics simulations to demonstrate that Isoxys species likely occupied a variety of niches in Cambrian oceans, including vertical migrants, providing the first quantitative evidence that some Cambrian animals were adapted for vertical movement in the water column. Vertical migration was one of several early Cambrian metazoan innovations that led to the biological pump taking on a modern-style architecture over 500 Myr ago.

Peracarida (e.g. woodlice and side-swimmers) are, together with their sister-group Eucarida (e.g. krill and decapods), the most speciose group of modern crustaceans, suggested to have appeared as early as the Ordovician. While eucarids' incursion onto land consists of mainly freshwater and littoral grounds, some peracarids have evolved fully terrestrial ground-crawling ecologies, inhabiting even our gardens in temperate regions (e.g. pillbugs and sowbugs). Their fossil record extends back to the Carboniferous and consists mainly of marine occurrences. Here, we provide a complete re-analysis of a fossil arthropod- Oxyuropoda- reported in 1908 from the Late Devonian floodplains of Ireland, and left with unresolved systematic affinities despite a century of attempts at identification. Known from a single specimen preserved in two dimensions, we analysed its anatomy using digital microscopy and multispectral macroimaging to enhance the contrast of morphological structures. The new anatomical characters and completeness of Oxyuropoda , together with a phylogenetic analysis with representatives of all major Eumalacostraca groups, indicate that Oxyuropoda is a crown peracarid, part of a clade including amphipods and isopods. As such, Oxyuropoda is the oldest known species Peracarida, and provides evidence that derived peracarids had an incursion into freshwater and terrestrial environments as early as the Famennian, more than 360 Ma.

Radiodonts have long been known from Cambrian deposits preserving non-biomineralizing organisms. In Utah, the presence of these panarthropods in the Spence and Wheeler (House Range and Drum Mountains) biotas is now well-documented. Conversely, radiodont occurrences in the Marjum Formation have remained scarce. Despite the large amount of work undertaken on its diverse fauna, only one radiodont ( Peytoia ) has been reported from the Marjum Biota. In this contribution we quadruple the known radiodont diversity of the Marjum fauna, with the description of the youngest members of two genera, Caryosyntrips and Pahvantia , and that of a new taxon Buccaspinea cooperi gen. et sp. nov. This new taxon can be identified from its large oral cone bearing robust hooked teeth with one, two, or three cusps, and by the unique endite morphology and organisation of its frontal appendages. Appendages of at least 12 podomeres bear six recurved plate-like endites proximal to up to four spiniform distal endites. Pahvantia hastata specimens from the Marjum Formation are particularly large, but otherwise morphologically indistinguishable from the carapace elements of this species found in the Wheeler Formation. One of the two new Caryosyntrips specimens can be confidently assigned to C. camurus . The other bears the largest spines relative to appendage length recorded for this genus, and possesses endites of variable size and unequal spacing, making its taxonomic assignment uncertain. Caryosyntrips, Pahvantia , and Peytoia are all known from the underlying Wheeler Formation, whereas isolated appendages from the Spence Shale and the Wheeler Formation, previously assigned to Hurdia , are tentatively reidentified as Buccaspinea . Notably, none of these four genera occurs in the overlying Weeks Formation, providing supporting evidence of a faunal restructuring around the Drumian-Guzhangian boundary. The description of three additional nektonic taxa from the Marjum Formation further documents the higher relative proportion of free-swimming species in this biota compared to those of the Wheeler and Weeks Lagerstätten. This could be related to a moderate deepening of the basin and/or changing regional ocean circulation at this time., Competing Interests: The authors declare that they have no competing interests., (© 2021 Pates et al.)

It is hypothesized that iron from biological tissues, liberated during decay, may have played a role in inhibiting loss of anatomical information during fossilization of extinct organisms. Most tissues in the animal kingdom contain iron in different forms. A widely distributed iron-bearing molecule is ferritin, a globular protein that contains iron crystallites in the form of ferrihydrite minerals. Iron concentrations in ferritin are high and ferrihydrites are extremely reactive. When ancient animals are decaying on the sea floor under anoxic environmental conditions, ferrihydrites may initialize the selective replication of some tissues in pyrite FeS 2 . This model explains why some labile tissues are preserved, while other more resistant structures decay and are absent in many fossils. A major implication of this hypothesis is that structures described as brains in Cambrian arthropods are not fossilization artifacts, but are instead a source of information on anatomical evolution at the dawn of complex animal life., (© 2020 WILEY Periodicals, Inc.)

Crustacean eggs are rare in the fossil record. Here we report the exquisite preservation of a fossil polychelidan embedded within an unbroken nodule from the Middle Jurassic La Voulte-sur-Rhône Lagerstätte (France) and found with hundreds of eggs attached to the pleon. This specimen belongs to a new species, Palaeopolycheles nantosueltae sp. nov. and offers unique clues to discuss the evolution of brooding behaviour in polychelidan lobsters. In contrast to their development, which now relies on a long-lived planktic larval stage that probably did not exist in the early evolutionary steps of the group, the brood size of polychelidan lobsters seems to have remained unchanged and comparatively small since the Jurassic. This finding is at odds with reproductive strategies in other lobster groups, in which a long-lived planktic larval stage is associated with a large brood size.

Background: Radiodonta, large Palaeozoic nektonic predators, occupy a pivotal evolutionary position as stem-euarthropods and filled important ecological niches in early animal ecosystems. Analyses of the anatomy and phylogenetic affinity of these large nektonic animals have revealed the origins of the euarthropod compound eye and biramous limb, and interpretations of their diverse feeding styles have placed various radiodont taxa as primary consumers and apex predators. Critical to our understanding of both radiodont evolution and ecology are the paired frontal appendages; however, the vast differences in frontal appendage morphology between and within different radiodont families have made it difficult to identify the relative timings of character acquisitions for this body part., Results: Here we describe a new genus of hurdiid, Ursulinacaris , from the middle Cambrian (Miaolingian, Wuliuan) Mount Cap Formation (Northwest Territories, Canada) and Jangle Limestone (Nevada, USA). Ursulinacaris has the same organisation as other hurdiid frontal appendages, with elongate endites on the first five podomeres in the distal articulated region and auxiliary spines on the distal margin of endites only. Unlike all other hurdiid genera, which possess a single row of elongated and blade-like ventral endites, this taxon uniquely bears paired slender endites., Conclusion: The blade-like endite morphology is shown to be a hurdiid autapomorphy. Two other frontal appendage characters known only in hurdiids, namely auxiliary spines on the distal margin of endites only, and elongate endites on the first five podomeres in the distal articulated region only, predate this innovation., Competing Interests: Competing interestsThe authors declare that they have no competing interests.

Ctenophores or 'comb-jellies' are marine animals whose relationship to other phyla is uncertain, yet important for understanding major steps in animal evolution. Fossil ctenophores from the Cambrian indicate that ctenophores may have evolved from a sessile, cnidarian-like ancestor., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Background: Extended parental care is a complex reproductive strategy in which progenitors actively look after their offspring up to - or beyond - the first juvenile stage in order to maximize their fitness. Although the euarthropod fossil record has produced several examples of brood-care, the appearance of extended parental care within this phylum remains poorly constrained given the scarcity of developmental data for Palaeozoic stem-group representatives that would link juvenile and adult forms in an ontogenetic sequence., Results: Here, we describe the post-embryonic growth of Fuxianhuia protensa from the early Cambrian Chengjiang Lagerstätte in South China. Our data demonstrate anamorphic post-embryonic development for F. protensa, in which new tergites were sequentially added from a posterior growth zone, the number of tergites varies from eight to 30. The growth of F. protensa is typified by the alternation between segment addition, followed by the depletion of the anteriormost abdominal segment into the thoracic region. The transformation of abdominal into thoracic tergite is demarcated by the development of laterally tergopleurae, and biramous walking legs. The new ontogeny data leads to the recognition of the rare Chengjiang euarthropod Pisinnocaris subconigera as a junior synonym of Fuxianhuia. Comparisons between different species of Fuxianhuia and with other genera within Fuxianhuiida suggest that heterochrony played a prominent role in the morphological diversification of fuxianhuiids. Functional analogy with the flexible trunk ontogeny of Cambrian and Silurian olenimorphic trilobites suggests an adaptation to sporadic low oxygen conditions in Chengjiang deposits for F. protensa. Finally, understanding the growth of F. protensa allows for the interpretation of an exceptional life assemblage consisting of a sexually mature adult alongside four ontogenetically coeval juveniles, which constitutes the oldest occurrence of extended parental care by prolonged cohabitation in the panarthropod fossil record., Conclusions: Our findings constitute the most detailed characterization of the post-embryonic development in a soft-bodied upper stem-group euarthropod available to date. The new ontogeny data illuminates the systematics, trunk segmentation and palaeoecology of F. protensa, offers insights on the macroevolutionary processes involved in the diversification of this clade, and contributes towards an improved understanding of complex post-embryonic reproductive ecology in Cambrian euarthropods.

Euarthropoda is one of the best-preserved fossil animal groups and has been the most diverse animal phylum for over 500 million years. Fossil Konservat-Lagerstätten, such as Burgess Shale-type deposits (BSTs), show the evolution of the euarthropod stem lineage during the Cambrian from 518 million years ago (Ma). The stem lineage includes nonbiomineralized groups, such as Radiodonta (e.g., Anomalocaris ) that provide insight into the step-by-step construction of euarthropod morphology, including the exoskeleton, biramous limbs, segmentation, and cephalic structures. Trilobites are crown group euarthropods that appear in the fossil record at 521 Ma, before the stem lineage fossils, implying a ghost lineage that needs to be constrained. These constraints come from the trace fossil record, which show the first evidence for total group Euarthropoda (e.g., Cruziana , Rusophycus ) at around 537 Ma. A deep Precambrian root to the euarthropod evolutionary lineage is disproven by a comparison of Ediacaran and Cambrian lagerstätten. BSTs from the latest Ediacaran Period (e.g., Miaohe biota, 550 Ma) are abundantly fossiliferous with algae but completely lack animals, which are also missing from other Ediacaran windows, such as phosphate deposits (e.g., Doushantuo, 560 Ma). This constrains the appearance of the euarthropod stem lineage to no older than 550 Ma. While each of the major types of fossil evidence (BSTs, trace fossils, and biomineralized preservation) have their limitations and are incomplete in different ways, when taken together they allow a coherent picture to emerge of the origin and subsequent radiation of total group Euarthropoda during the Cambrian., Competing Interests: The authors declare no conflict of interest.

Background: Segmental composition and homologies of the head of stem-group Euarthropoda have been the foci of recent studies on arthropod origins. An emerging hypothesis suggests that upper-stem group euarthropods possessed a three-segmented head/brain, including an ocular segment (protocerebrum) followed by the deutocerebrum with associated antennae/raptorial limbs and the tritocerebrum, while in the lower stem, head structures of Radiodonta are wholly associated with the protocerebrum and its preceding part. However, this hypothesis is incompletely tested because detailed knowledge on the head components of radiodontans is patchy, and informative articulated specimens are lacking for many taxa. Amplectobelua symbrachiata is the most common radiodontan species in the Chengjiang biota (ca. 520 Ma), normally known as isolated frontal appendages. Here we present detailed descriptions of new articulated specimens that elucidate the morphology and function of its head structures, and discuss their implications for hypotheses about euarthropod cephalic organisation., Results: In addition to a central oval head shield, A. symbrachiata also bears a pair of P-elements connected by an elongated rod. The mouth consists of sets of smooth and tuberculate plates, in contrast to the typical radial oral cones of other radiodontans. Previously identified 'palm-like teeth' are located external to the mouth in the posterior head region, and are interpreted as segmental gnathobase-like structures (GLSs) associated with at least three reduced transitional flaps in a one (pair)-to-one (pair) pattern, consistent with an appendicular nature. Comparisons with other panarthropods show that GLSs are morphologically similar to the mandibles and other gnathobasic mouthparts of euarthropods, as well as to the jaws of onychophorans, indicating their functional integration into the feeding activities of A. symbrachiata., Conclusions: The functional head of A. symbrachiata must include the reduced transitional segments (and their associated structures), which have been identified in several other radiodontans. This functional view supports the idea that the integration of segments (and associated appendages) into the head region, probably driven by feeding, occurred along the euarthropod stem-lineage. However, the number of reduced transitional segments varies between different groups and it remains uncertain whether GLSs represent proximal or distal parts of appendages. Our study is the first description of appendicular structures other than the frontal appendages in the functional head region of radiodontans, revealing novel feeding structures in the morphological transition from the lower- to the upper- stem-group of Euarthropoda.

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.)

Exceptionally preserved fossils from the Palaeozoic era provide crucial insights into arthropod evolution, with recent discoveries bringing phylogeny and character homology into sharp focus. Integral to such studies are anomalocaridids, a clade of stem arthropods whose remarkable morphology illuminates early arthropod relationships and Cambrian ecology. Although recent work has focused on the anomalocaridid head, the nature of their trunk has been debated widely. Here we describe new anomalocaridid specimens from the Early Ordovician Fezouata Biota of Morocco, which not only show well-preserved head appendages providing key ecological data, but also elucidate the nature of anomalocaridid trunk flaps, resolving their homology with arthropod trunk limbs. The new material shows that each trunk segment bears a separate dorsal and ventral pair of flaps, with a series of setal blades attached at the base of the dorsal flaps. Comparisons with other stem lineage arthropods indicate that anomalocaridid ventral flaps are homologous with lobopodous walking limbs and the endopod of the euarthropod biramous limb, whereas the dorsal flaps and associated setal blades are homologous with the flaps of gilled lobopodians (for example, Kerygmachela kierkegaardi, Pambdelurion whittingtoni) and exites of the 'Cambrian biramous limb'. This evidence shows that anomalocaridids represent a stage before the fusion of exite and endopod into the 'Cambrian biramous limb', confirming their basal placement in the euarthropod stem, rather than in the arthropod crown or with cycloneuralian worms. Unlike other anomalocaridids, the Fezouata taxon combines head appendages convergently adapted for filter-feeding with an unprecedented body length exceeding 2 m, indicating a new direction in the feeding ecology of the clade. The evolution of giant filter-feeding anomalocaridids may reflect the establishment of highly developed planktic ecosystems during the Great Ordovician Biodiversification Event.

Understanding the way in which animals diversified and radiated during their early evolutionary history remains one of the most captivating of scientific challenges. Integral to this is the 'Cambrian explosion', which records the rapid emergence of most animal phyla, and for which the triggering and accelerating factors, whether environmental or biological, are still unclear. Here we describe exceptionally well-preserved complex digestive organs in early arthropods from the early Cambrian of China and Greenland with functional similarities to certain modern crustaceans and trace these structures through the early evolutionary lineage of fossil arthropods. These digestive structures are assumed to have allowed for more efficient digestion and metabolism, promoting carnivory and macrophagy in early arthropods via predation or scavenging. This key innovation may have been of critical importance in the radiation and ecological success of Arthropoda, which has been the most diverse and abundant invertebrate phylum since the Cambrian.

Ecdysozoans have been key components of ecosystems since the early Cambrian, when trilobites and soft-bodied Burgess Shale-type ecdysozoans dominated marine animal communities. Even today, the most abundant animals on Earth are either nematode worms or plankton-forming crustaceans, whereas the most diverse are the insects. Throughout geological time, several ecdysozoan lineages independently colonized land, shaping both marine and terrestrial ecosystems and providing an adequate environment for successive animal terrestrialization. The timing of these events is largely uncertain and has been investigated only partially using molecular data. Here we present a timescale of ecdysozoan evolution based on multiple molecular data sets, the most complete set of fossil calibrations to date, and a thorough series of validation analyses. Results converge on an Ediacaran origin of all major ecdysozoan lineages (∼587-543 million years ago [mya]), followed by a fast Cambrian radiation of the pancrustaceans (∼539-511 mya), a Cambro-Ordovician colonization of land of different arthropod lineages (∼510-471 mya), and a relatively recent radiation of extant nematodes, onychophorans, and tardigrades (∼442 mya). Arthropods colonized land nearly synchronously with land plants. Further diversification within flying insects, nematodes and onychophorans might be related to the evolution of vascular plants and forests., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The Cambro-Ordovician anomalocaridids are large ecdysozoans commonly regarded as ancestors of the arthropods and apex predators. Predation is indicated partly by the presence of an unusual "peytoia"-type oral cone, which is a tetraradial outer ring of 32 plates, four of which are enlarged and in perpendicular arrangement. This oral cone morphology was considered a highly consistent and defining characteristic of well-known Burgess Shale taxa. It is here shown that Anomalocaris has a different oral cone, with only three large plates and a variable number of smaller and medium plates. Its functional morphology suggests that suction, rather than biting, was used for food ingestion, and that anomalocaridids in general employed a range of different scavenging and predatory feeding strategies. Removing anomalocaridids from the position of highly specialized trilobite predators forces a reconsideration of the ecological structure of the earliest marine animal communities in the Cambrian.

As the largest predators of the Cambrian seas, the anomalocaridids had an important impact in structuring the first complex marine animal communities, but many aspects of anomalocaridid morphology, diversity, ecology, and affinity remain unclear owing to a paucity of specimens. Here we describe the anomalocaridid Hurdia, based on several hundred specimens from the Burgess Shale in Canada. Hurdia possesses a general body architecture similar to those of Anomalocaris and Laggania, including the presence of exceptionally well-preserved gills, but differs from those anomalocaridids by possessing a prominent anterior carapace structure. These features amplify and clarify the diversity of known anomalocaridid morphology and provide insight into the origins of important arthropod features, such as the head shield and respiratory exites.