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The evolution of articulated sclerites via soft membranes, termed arthrodization, is arguably one of the most critical innovations in animals. Defining the megaphylum Arthropoda, the arthrodization of appendages, or arthropodization, likely predated that of the body, the combination of both being diagnostic of true arthropods (Euarthropoda) – all of these innovations occurring during the Cambrian explosion. Here, thanks to dozens of exceptionally preserved fossils from the Cambrian Wuliuan Stage Burgess Shale (Tulip Beds locality on Mount Stephen, British Columbia, Canada), we show that a distinct but comparable system of imbricated sclerotic elements evolved in the paraphyletic sister group of arthropods, the lobopodians. Entothyreos synnaustrus gen. et sp. nov. has characteristic body plan features of the Collinsovermidae (order Luolishaniida), including anterior limbs for suspension-feeding and stout anchoring posterior limbs. Uniquely, however, E. synnaustrus also displays segmental sclerotic sheets along the trunk, covered in a thin layer of integument, as well as overlapping sclerotized annuli on posterior-most limbs. While the latter elements likely served a protective function, the dorsolateral trunk sheets, which also carry spines, may have facilitated body erection and suspension-feeding. Other luolishaniids possess separate ring-like structures connecting the base of metameric spines which are covered by the apical layer of the lobopodian integument. E. synnaustrus and related taxa illustrate, therefore, an arguably parallel evolution of arthropod-like morphoanatomical features early during the rise of panarthropods. This finding broadens our perspectives on the uniqueness of major synapomorphies and the importance of including canalization in macroevolutionary narratives. [ABSTRACT FROM AUTHOR]

The diversity of cephalic morphologies in mandibulates (myriapods and pancrustaceans) was key to their evolutionary success. A group of Cambrian bivalved arthropods called hymenocarines exhibit diagnostic mandibulate traits that illustrate this diversity, but many forms are still poorly known. These include the odaraiids, typified by Odaraia alata from the Burgess Shale (Wuliuan), characterized by its unique tubular carapace and rudder-like tail fan, and one of the largest Cambrian euarthropods at nearly 20 cm in length. Unfortunately, odaraiid cephalic anatomy has been largely unknown, limiting evolutionary scenarios and putting their mandibulate affinities into question. Here, we reinvestigate Odaraia based on new specimens from the Burgess Shale and describe exquisitely preserved mandibles with teeth and adjacent structures: a hypostome, maxillae and potential paragnaths. These structures can be homologized with those of Cambrian fuxianhuiids and extant mandibulates, and suggest that the ancestral mandibulate head could have had a limbless segment but retained its plasticity, allowing for limb re-expression within Pancrustacea. Furthermore, we show the presence of limbs with spinose endites which created a suspension-feeding structure. This discovery provides morphological evidence for suspension feeding among large Cambrian euarthropods and evinces the increasing exploitation of planktonic resources in Cambrian pelagic food webs. [ABSTRACT FROM AUTHOR]

Injured trilobites present insight into how a completely extinct group of arthropods responded to traumatic experiences, such as failed predation and moulting complications. These specimens are therefore important for more thoroughly understanding the Paleozoic predator-prey systems that involved trilobites. To expand the record of injured trilobites, we present new examples of injured Ogygopsis klotzi and Olenoides serratus from the Campsite Cliff Shale Member of the Burgess Shale Formation (Cambrian, Miaolingian, Wuliuan), Paradoxides (Paradoxides) paradoxissimus gracilis from the Jince Formation (Cambrian, Miaolingian, Drumian), Ogygiocarella angustissima from the Llanfawr Mudstones Formation (Middle-Late Ordovician, Darriwilian-Sandbian), and Ogygiocarella debuchii from the Meadowtown Formation, (Middle-Late Ordovician, Darriwilian-Sandbian). We consider the possible origins of these malformations and conclude that most injuries reflect failed predation. Within this framework, possible predators are presented, and we uncover a marked shift in the diversity of animals that targeted trilobites in the Ordovician. We also collate other records of injured Ogygo. klotzi and Ol. serratus, and Ogygi. debuchii, highlighting that these species are targets for further understanding patterns and records of trilobite injuries. [ABSTRACT FROM AUTHOR]

Injured trilobites present insight into how a completely extinct group of arthropods responded to traumatic experiences, such as failed predation and moulting complications. These specimens are therefore important for more thoroughly understanding the Paleozoic predator-prey systems that involved trilobites. To expand the record of injured trilobites, we present new examples of injured Ogygopsis klotzi and Olenoides serratus from the Campsite Cliff Shale Member of the Burgess Shale Formation (Cambrian, Miaolingian, Wuliuan), Paradoxides (Paradoxides) paradoxissimus gracilis from the Jince Formation (Cambrian, Miaolingian, Drumian), Ogygiocarella angustissima from the Llanfawr Mudstones Formation (Middle–Late Ordovician, Darriwilian–Sandbian), and Ogygiocarella debuchii from the Meadowtown Formation, (Middle–Late Ordovician, Darriwilian–Sandbian). We consider the possible origins of these malformations and conclude that most injuries reflect failed predation. Within this framework, possible predators are presented, and we uncover a marked shift in the diversity of animals that targeted trilobites in the Ordovician. We also collate other records of injured Ogygo. klotzi and Ol. serratus, and Ogygi. debuchii, highlighting that these species are targets for further understanding patterns and records of trilobite injuries.

Early annelid evolution is mostly known from 13 described species from Cambrian Burgess Shale-type Lagerstätten. We introduce a new exceptionally well-preserved polychaete, Ursactis comosa gen. et sp. nov., from the Burgess Shale (Wuliuan Stage). This small species (3–15 mm) is the most abundant Cambrian polychaete known to date. Most specimens come from Tokumm Creek, a new Burgess Shale locality in northern Kootenay National Park, British Columbia, Canada. Ursactis has a pair of large palps, thin peristomial neurochaetae and biramous parapodia bearing similarly sized capillary neurochaetae and notochaetae, except for segments six to nine, which also have longer notochaetae. The number of segments in this polychaete range between 8 and 10 with larger individuals having 10 segments. This number of segments in Ursactis is remarkably small compared with other polychaetes, including modern forms. Specimens with 10 segments show significant size variations, and the length of each segment increases with the body length, indicating that body growth was primarily achieved by increasing the size of existing segments rather than adding new ones. This contrasts with most modern polychaetes, which typically have a larger number of segments through additions of segments throughout life. The inferred growth pattern in Ursactis suggests that annelids had evolved control over segment addition by the mid-Cambrian.

Vetulicolians are an enigmatic phylum of extinct Cambrian marine invertebrates. They are particularly diverse in the Chengjiang Biota of China, but representatives have been recovered from other Fossil-Lagerstätten (Cambrian Stage 3-Drumian). These organisms are characterized by a bipartite body, which is split into an anterior section and a posterior segmented section connected by a narrow constriction. Here we report new material of the genus Pomatrum from the Cambrian Balang Biota (Series 2, Stage 4) of Hunan, southern China. This is the first discovery of this vetulicolian outside of the Chengjiang Biota (Series 2, Stage 3) and the first report of vetulicolians from the Balang Biota. This finding not only suggests that this group had a wider spatial and temporal distribution than previously known, but also adds information to the overall biodiversity of the Balang Biota-one of the most important Stage 4 fossil deposits known from Gondwana., Competing Interests: The authors declare that they have no competing interests., (© 2025 Ma et al.)

The origin of mandibulates, the hyperdiverse arthropod group that includes pancrustaceans and myriapods, dates back to the Cambrian. Bivalved arthropod groups such as hymenocarines have been argued to be early mandibulates, but many species are still poorly known, and their affinities remain uncertain. One of the most common and globally distributed Cambrian bivalved arthropods is Tuzoia. Originally described in 1912 from the Burgess Shale based on isolated carapaces, its full anatomy has remained largely unknown. Here, we describe new specimens of Tuzoia from the Canadian Burgess Shale (Wuliuan, Cambrian) showcasing exceptionally preserved soft tissues, allowing for the first comprehensive reconstruction of its anatomy, ecology and evolutionary affinities. The head bears antennae and differentiated cephalic appendages. The body is divided into a cephalothorax, a homonomous trunk bearing ca 10 pairs of legs with heptopodomerous endopods and enlarged basipods, and a tail fan with two pairs of caudal rami. These traits suggest that Tuzoia swam along the seafloor and used its spinose legs for predation or scavenging. Tuzoia is retrieved by a Bayesian phylogenetic analysis as an early mandibulate hymenocarine lineage, exemplifying the rapid diversification of this group in open marine environments during the Cambrian Explosion.

Radiodonts, stem-group euarthropods that evolved during the Cambrian explosion, were among the largest and most diversified lower palaeozoic predators. These animals were widespread geographically, occupying a variety of ecological niches, from benthic foragers to nektonic suspension feeders and apex predators. Here, we describe the largest Cambrian hurdiid radiodont known so far, Titanokorys gainesi, gen. et sp. nov., from the Burgess Shale (Marble Canyon, Kootenay National Park, British Columbia). Estimated to reach half a metre in length, this new species bears a very large ovoid-shaped central carapace with distinct short posterolateral processes and an anterior spine. Geometric morphometric analyses highlight the high diversity of carapace shapes in hurdiids and show that Titanokorys bridges a morphological gap between forms with long and short carapaces. Carapace shape, however, is prone to homoplasy and shows no consistent relationship with trophic ecology, as demonstrated by new data, including a reappraisal of the poorly known Pahvantia. Despite distinct carapaces, Titanokorys shares similar rake-like appendages for sediment-sifting with Cambroraster, a smaller but much more abundant sympatric hurdiid from the Burgess Shale. The co-occurrence of these two species on the same bedding planes highlights potential competition for benthic resources and the high diversity of large predators sustained by Cambrian communities.

Cambrian bivalved arthropods are a polyphyletic group of carapace‐bearing arthropods that includes stem euarthropods, stem mandibulates and crustaceans. Here, we describe Pakucaris apatis gen. et sp. nov., a new stem mandibulate bivalved arthropod from the middle Cambrian (Wuliuan Stage) Burgess Shale (Marble Canyon, Kootenay National Park, British Columbia, Canada). Two morphotypes of this small arthropod (11.6–26.6 mm long) are recognized, which differ mainly in their size and number of segments, possibly reflecting sexual dimorphism or different anamorphic stages. The carapace presents a dorsal crest extending anteriorly into a small recurved rostrum and two anterolateral processes. Around 20% of the posteriormost body segments and limbs are covered by a large spine‐bearing shield. The head bears a pair of eyes, a possible pair of unsegmented appendicular projections and two pairs of segmented appendages. The thorax is multisegmented, homonomous, with weakly sclerotized segments bearing biramous limbs, composed of a stenopodous endopod with c. 20 podomeres and a paddle‐shaped exopod. Pakucaris is interpreted as a nektobenthic suspension feeder. Bayesian phylogenetic analysis implies a position within Hymenocarina as stem mandibulates. The posterior shield is regarded as a pygidium, and represents a case of morphofunctional convergent evolution between mandibulates, artiopodans and mollisoniids. Furthermore, Pakucaris adds to a growing number of pygidium‐bearing arthropods, potentially hinting at a common developmental pattern across early arthropod evolution. This study not only increases our understanding of the early evolution of mandibulates, but also illustrates a unique case of early evolutionary convergence during the Cambrian Explosion. [ABSTRACT FROM AUTHOR]

Pterobranchs originated in the basal Cambrian (Fortunian) and are mostly known by their tubes preserved in the fossil record. The earliest forms are represented by bushy erect growing colonies that are not widely studied due to their scarcity, preservation quality and species misidentification. For this reason, early phylogenetic relationships within the group are not clearly established. Middle Cambrian Burgess Shale graptolites were poorly known, based on the presence of Chaunograptus scandens, some debatable species of the genus Yuknessia, and other undetermined fragmented material. This study represents a complete description of C. scandens, a consensus for Yuknessia simplex and Y. stephenensis, and new reports of Protohalecium hallianum and Mastigograptus sp. from the Burgess Shale localities, which have also been found in other Burgess Shale type localities in Utah and Australia. Phylogenetic analyses of 34 discrete morphological traits from these Burgess Shale genera and some known benthic and planktic taxa (n = 22), place these Cambrian species as basal forms closer to the pseudocolonial pterobranch Cephalodiscus and the living graptolite Rhabdopleura. [ABSTRACT FROM AUTHOR]

Lobopodians, a paraphyletic group of rare but morphologically diverse Palaeozoic vermiform animals bearing metameric appendages, are key to the origin of extant panarthropods. First discovered in 1983 on Mount Stephen (Yoho National Park, British Columbia), the Cambrian (Wuliuan) Burgess Shale lobopodian nicknamed 'Collins' monster' is formally described as Collinsovermis monstruosus gen. et sp. nov. A formal systematic treatment of the comparable and poorly known lobopodian Acinocricus stichus from Utah is also provided. The body of Collinsovermis is plump and compact but shows the diagnostic suspension‐feeding characters of luolishaniid lobopodians. It possesses 14 contiguous pairs of lobopods, lacking space between them. The 6 anterior pairs are elongate, adorned with about 20 pairs of long and slightly curved ventral spinules arranged in a chevron‐like pattern. These appendages terminate in a pair of thin claws and their dorsal surfaces are covered in minute spines or setae. The 8 posterior lobopod pairs, which attach to a truncated body termination, are stout and smooth, each terminated by a single strong recurved claw. Each somite bears a pair of dorsal spines; somites 4 and posteriad bear an additional median spine. The spines on somites 1–3 are much shorter than the spines on the remaining somites. The head is short, bears a terminal mouth and a pair of antenniform outgrowths, and is covered by an oblong sclerite. Collinsovermis, plus Collinsium and Acinocricus, are found to comprise a sub‐group of stout luolishaniid lobopodians with remarkably long spinules on the front lobopods, interpreted here as a clade (Teratopodidae fam. nov.) This clade is distinct from both the comparatively slenderer Luolishania and a sub‐group composed of Facivermis and Ovatiovermis lacking body sclerites. Luolishaniids were mostly sessile forerunners of arthropods that had coupled efficient suspension‐feeding devices and, as in Collinsovermis, strong defensive or deterrent features. [ABSTRACT FROM AUTHOR]

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 FeS2. 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. [ABSTRACT FROM AUTHOR]

An experimental decay methodology is developed for a cnidarian model organism to serve as a comparison to the many previous such studies on bilaterians. This allows an examination of inherent bias against the fossilisation of cnidarian tissue and their diagnostic characters, under what conditions these occur, and in what way. The decay sequence of Actinia equina was examined under a series of controlled conditions. These experiments show that cnidarian decay begins with an initial rupturing of the epidermis, followed by rapid loss of recognisable internal morphological characters. This suggests that bacteria work quicker on the epidermis than autolysis does on the internal anatomy. The data also show that diploblastic tissue is not universally decayed more slowly under anoxic or reducing conditions than under oxic conditions. Indeed, some cnidarian characters decay more rapidly under anoxic conditions than they do under oxic conditions. This suggests the decay pathways acting may be different to those affecting soft bilaterian tissue such as soft epidermis and internal organs. What is most important in the decay of soft polyp anatomy is the microbial community, which can be dominated by oxic or anoxic bacteria. Different Lagerstätte, even of the same type, will inevitably have subtle difference in their bacterial communities, which among other factors, could be a control on soft polyp preservation leading to either an absence of compelling soft anthozoans (Burgess Shale) or an astonishing abundance (Qingjiang biota). [ABSTRACT FROM AUTHOR]

Abstract Background Chelicerata represents a vast clade of mostly predatory arthropods united by a distinctive body plan throughout the Phanerozoic. Their origins, however, with respect to both their ancestral morphological features and their related ecologies, are still poorly understood. In particular, it remains unclear whether their major diagnostic characters were acquired early on, and their anatomical organization rapidly constrained, or if they emerged from a stem lineage encompassing an array of structural variations, based on a more labile “panchelicerate” body plan. Results In this study, we reinvestigated the problematic middle Cambrian arthropod Habelia optata Walcott from the Burgess Shale, and found that it was a close relative of Sanctacaris uncata Briggs and Collins (in Habeliida, ord. nov.), both retrieved in our Bayesian phylogeny as stem chelicerates. Habelia possesses an exoskeleton covered in numerous spines and a bipartite telson as long as the rest of the body. Segments are arranged into three tagmata. The prosoma includes a reduced appendage possibly precursor to the chelicera, raptorial endopods connected to five pairs of outstandingly large and overlapping gnathobasic basipods, antennule-like exopods seemingly dissociated from the main limb axis, and, posteriorly, a pair of appendages morphologically similar to thoracic ones. While the head configuration of habeliidans anchors a seven-segmented prosoma as the chelicerate ground pattern, the peculiar size and arrangement of gnathobases and the presence of sensory/tactile appendages also point to an early convergence with the masticatory head of mandibulates. Conclusions Although habeliidans illustrate the early appearance of some diagnostic chelicerate features in the evolution of euarthropods, the unique convergence of their cephalons with mandibulate anatomies suggests that these traits retained an unusual variability in these taxa. The common involvement of strong gnathal appendages across non-megacheiran Cambrian taxa also illustrates that the specialization of the head as the dedicated food-processing tagma was critical to the emergence of both lineages of extant euarthropods—Chelicerata and Mandibulata—and implies that this diversification was facilitated by the expansion of durophagous niches.

The origin of the arthropod carapace, an enlargement of cephalic tergites, can be traced back to the Cambrian period. However, its disparity and evolution are still not fully understood. Here, we describe a new ‘bivalved’ arthropod, Fibulacaris nereidis gen. et sp. nov., based on 102 specimens from the middle Cambrian (Wuliuan Stage) Burgess Shale, Marble Canyon area in British Columbia's Kootenay National Park, Canada. The laterally compressed carapace covers most of the body. It is fused dorsally and merges anteriorly into a conspicuous postero-ventrally recurved rostrum as long as the carapace and positioned between a pair of backwards-facing pedunculate eyes. The body is homonomous, with approximately 40 weakly sclerotized segments bearing biramous legs with elongate endopods, and ends in a pair of small flap-like caudal rami. Fibulacaris nereidis is interpreted as a suspension feeder possibly swimming inverted, in a potential case of convergence with some branchiopods. A Bayesian phylogenetic analysis places it within a group closely related to the extinct Hymenocarina. Fibulacaris nereidis is unique in its carapace morphology and overall widens the ecological disparity of Cambrian arthropods and suggests that the evolution of a ‘bivalved' carapace and an upside-down lifestyle may have occurred early in stem-group crustaceans.

Radiodonts, a clade of Cambro-Devonian stem group euarthropods, have classically been regarded as nektonic apex predators. However, many aspects of radiodont morphology and ecology have remained unclear because of the typically fragmentary nature of fossil material. Here, we describe a new hurdiid radiodont based on abundant and exceptionally well-preserved fossils from the Burgess Shale (Marble Canyon area, British Columbia, Canada). Cambroraster falcatus gen. et sp. nov. is characterized by an extra-large horseshoe-shaped head carapace, bearing conspicuous posterolateral spinous processes, and partially covering a short trunk with eight pairs of lateral flaps. Each of the pair of frontal appendages possess five mesially curving rake-like endites equipped with a series of anteriorly directed hooked spines, altogether surrounding the oral cone. This feeding apparatus suggests a micro to macrophagous sediment-sifting feeding ecology. Cambroraster illuminates the evolution of Hurdiidae and evinces the exploitation of the diversifying infauna by these large and specialized nektobenthic carnivores in the aftermath of the Cambrian explosion. [ABSTRACT FROM AUTHOR]

Naraoiids are non‐biomineralized euarthropods characterized by the complete fusion of post‐cephalic tergo‐pleurae into a single shield, as well as an extensively ramified digestive tract. Ranging from the early Cambrian to the late Silurian (Pridoli), these arthropods of simple appearance have traditionally been associated with the early diversification of trilobites and their close relatives, but the interrelationships and affinities of naraoiids within Artiopoda remain poorly characterized. Three new species from the Burgess Shale (middle Cambrian, Stage 5) of British Columbia, Canada, are described here: Misszhouia canadensis sp. nov., from Marble Canyon (Kootenay National Park), the first species belonging to the genus Misszhouia outside of China; Naraoia magna sp. nov., from Marble Canyon and also from the Raymond Quarry (Yoho National Park), the largest species of Naraoia described thus far, reaching up to 9 cm in length; and Naraoia arcana sp. nov., from two sublocalities on Mount Stephen (Yoho National Park), defined by its unusual combination of spines. This new material shows that gut morphology is no longer a reliable character to distinguish Misszhouia from Naraoia. We demonstrate that Naraoia and Misszhouia can instead be discriminated morphometrically, based on simple metrics of the dorsal exoskeleton. Our quantitative results also help with inter‐specific discrimination and illustrate possible cases of sexual dimorphism. Phylogenetically, the inclusion of morphometric data adds resolution to our cladogram, although parsimony and likelihood treatments provide somewhat different evolutionary scenarios. In all cases, liwiines are nested within Naraoiidae, resolved as the most derived clade of trilobitomorph arthropods. [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]

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

Waptia fieldensis Walcott, 1912 is one of the iconic animals from the middle Cambrian Burgess Shale biota that had lacked a formal description since its discovery at the beginning of the twentieth century. This study, based on over 1800 specimens, finds that W. fieldensis shares general characteristics with pancrustaceans, as previous authors had suggested based mostly on its overall aspect. The cephalothorax is covered by a flexible, bivalved carapace and houses a pair of long multisegmented antennules, palp-bearing mandibles, maxillules, and four pairs of appendages with five-segmented endopods—the anterior three pairs with long and robust enditic basipods, the fourth pair with proximal annulations and lamellae. The post-cephalothorax has six pairs of lamellate and fully annulated appendages which appear to be extensively modified basipods rather than exopods. The front part of the body bears a pair of stalked eyes with the first ommatidia preserved in a Burgess Shale arthropod, and a median ‘labral’ complex flanked by lobate projections with possible affinities to hemi-ellipsoid bodies. Waptia confirms the mandibulate affinity of hymenocarines, retrieved here as part of an expanded Pancrustacea, thereby providing a novel perspective on the evolutionary history of this hyperdiverse group. We construe that Waptia was an active swimming predator of soft prey items, using its anterior appendages for food capture and manipulation, and also potentially for clinging to epibenthic substrates.