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Ordovician open marine Lagerstätten are relatively rare and widely dispersed, producing a patchy picture of the diversity and biogeography of nonmineralized marine organisms and challenging our understanding of the fate of Cambrian groups. Here, for the first time, we report soft-bodied fossils, including a well-preserved marrellomorph arthropod, fragmentary carapaces, and macroalgae, from the Late Ordovician (Katian) Upper Member of the Kirkfield Formation near Brechin, Ontario. The unmineralized elements and associated exceptionally preserved shelly biota were entombed rapidly in storm deposits that smothered the shallow, carbonate-dominated shelf. The marrellomorph, Tomlinsonus dimitrii n. gen. n. sp., is remarkable for its ornate, curving cephalic spines and pair of hypertrophied appendages, suggesting a slow-moving, benthic lifestyle. Reevaluation of marrellomorph phylogeny using new data favors an arachnomorph affinity, although internal relationships are robust to differing outgroup selection. Clades Marrellida and Acercostraca are recovered, but the monophyly of Marrellomorpha is uncertain. The new taxon is recovered as sister to the Devonian Mimetaster and, as the second-youngest known marrellid, bridges an important gap in the evolution of this clade. More generally, the Brechin biota represents a rare window into Ordovician open marine shelf environments in Laurentia, representing an important point of comparison with contemporaneous Lagerstätten from other paleocontinents, with great potential for further discoveries.UUID: http://zoobank.org/884589d0-08f7-4398-ab42-1f5d459be9e9

Exceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

By contrast to many previously enigmatic Palaeozoic fossils, the Carboniferous metazoan

Radiodonts (stem Euarthropoda) were ecologically diverse, but species generally displayed limited functional specialization of appendages along the body axis compared with crown group euarthropods. This is puzzling, because such functional specialization is considered to have been an important driver of euarthropod ecological diversification. One way to circumvent this constraint could have been the functional specialization of different parts of the frontal appendages, known to have been ecologically important in radiodonts. This hypothesis has yet to be tested explicitly. Here we redescribe the poorly known mid-Cambrian hurdiid radiodont Stanleycaris hirpex from the Burgess Shale (Stephen Formation) and quantitatively assess functional specialization of the frontal appendages of stem euarthropods. The appendages of Stanleycaris are composed of 14 podomeres, variously differentiated by their possession of pectinate endites, mono- to trifurcate medial gnathites, and outer spines. The oral cone is tetraradially organized and can be uniquely distinguished from those of other hurdiids by the presence of 28 rather than 32 smooth tridentate plates. Our phylogenetic analysis finds Stanleycaris in a grade of hurdiids retaining plesiomorphic raptorial appendicular functionality alongside derived adaptations for sweep feeding and large, bilaterally opposed gnathites. We conclude that the latter performed a masticatory function, convergent with gnathal structures like mandibles in various panarthropods. Taken together, Stanleycaris and similar hurdiids provide an extreme example of the evolution of division of labor within the appendage of a stem euarthropod and suggest that this innovation may have facilitated the functional transition, from raptorial to sweep feeding, at the origin of the hurdiid clade.

In addition to being among the most iconic and bizarre-looking Cambrian animals, radiodonts are a group that offers key insight into the acquisition of the arthropod body plan by virtue of their phylogenetic divergence prior to all living members of the phylum. Nonetheless, radiodont fossils are rare and often fragmentary, and contentions over their interpretation have hindered resolution of important evolutionary conundrums. Here, we describe 268 specimens of Stanleycaris hirpex from the Cambrian Burgess Shale, including many exceptionally preserved whole-body specimens, informing the most complete reconstruction of a radiodont to date. The trunk region of Stanleycaris has up to 17 segments plus two pairs of filiform caudal blades. The recognition of dorsal sclerotic segmentation of the trunk cuticle and putative unganglionated nerve cords provides new insight into the relative timing of acquisition of segmental traits, the epitome of the arthropod body plan. In addition to the pair of stalked lateral eyes, the short head unexpectedly bears a large median eye situated behind a preocular sclerite on an anteriorly projecting head lobe. Upon re-evaluation, similar median eyes can be identified in other Cambrian panarthropods demonstrating a deep evolutionary continuity. The exquisitely preserved brain of Stanleycaris is consistent with the hypothesized deutocerebral innervation of the frontal appendages, reconciling neuroanatomical evidence with external morphology in support of an ancestrally bipartite head and brain for arthropods. We propose that the integration of this bipartite head prior to the acquisition of most segmental characters exclusively in the arthropod trunk may help explain its developmental differentiation.

The middle (Wuliuan Stage) Cambrian Burgess Shale is famous for its exceptional preservation of diverse and abundant soft-bodied animals through the “thick” Stephen Formation. However, with the exception of the Walcott Quarry (Fossil Ridge) and the stratigraphically older Tulip Beds (Mount Stephen), which are both in Yoho National Park (British Columbia), quantitative assessments of the Burgess Shale have remained limited. Here we first provide a detailed quantitative overview of the diversity and structure of the Marble Canyon Burgess Shale locality based on 16,438 specimens. Located 40 km southeast of the Walcott Quarry in Kootenay National Park (British Columbia), Marble Canyon represents the youngest site of the “thick” Stephen Formation. We then combine paleoecological data sets from Marble Canyon, Walcott Quarry, Tulip Beds, and Raymond Quarry, which lies approximately 20 m directly above the Walcott Quarry, to yield a combined species abundance data set of 77,179 specimens encompassing 234 species-level taxa. Marble Canyon shows significant temporal changes in both taxonomic and ecological groups, suggesting periods of stasis followed by rapid turnover patterns at local and short temporal scales. At wider geographic and temporal scales, the different Burgess Shale sites occupy distinct areas in multivariate space. Overall, this suggests that the Burgess Shale paleocommunity is far patchier than previously thought and varies at both local and regional scales through the “thick” Stephen Formation. This underscores that our understanding of Cambrian diversity and ecological networks, particularly in early animal ecosystems, remains limited and highly dependent on new discoveries.

The chelicerates are a ubiquitous and speciose group of animals that has a considerable ecological effect on modern terrestrial ecosystems—notably as predators of insects and also, for instance, as decomposers1. The fossil record shows that chelicerates diversified early in the marine ecosystems of the Palaeozoic era, by at least the Ordovician period2. However, the timing of chelicerate origins and the type of body plan that characterized the earliest members of this group have remained controversial. Although megacheirans3–5 have previously been interpreted as chelicerate-like, and habeliidans6 (including Sanctacaris7,8) have been suggested to belong to their immediate stem lineage, evidence for the specialized feeding appendages (chelicerae) that are diagnostic of the chelicerates has been lacking. Here we use exceptionally well-preserved and abundant fossil material from the middle Cambrian Burgess Shale (Marble Canyon, British Columbia, Canada) to show that Mollisonia plenovenatrix sp. nov. possessed robust but short chelicerae that were placed very anteriorly, between the eyes. This suggests that chelicerae evolved a specialized feeding function early on, possibly as a modification of short antennules. The head also encompasses a pair of large compound eyes, followed by three pairs of long, uniramous walking legs and three pairs of stout, gnathobasic masticatory appendages; this configuration links habeliidans with euchelicerates (‘true’ chelicerates, excluding the sea spiders). The trunk ends in a four-segmented pygidium and bears eleven pairs of identical limbs, each of which is composed of three broad lamellate exopod flaps, and endopods are either reduced or absent. These overlapping exopod flaps resemble euchelicerate book gills, although they lack the diagnostic operculum9. In addition, the eyes of M. plenovenatrix were innervated by three optic neuropils, which strengthens the view that a complex malacostracan-like visual system10,11 might have been plesiomorphic for all crown euarthropods. These fossils thus show that chelicerates arose alongside mandibulates12 as benthic micropredators, at the heart of the Cambrian explosion. Mollisonia plenovenatrix, a small predatory arthropod from the Burgess Shale dated to about 508 million years ago, is morphologically close to horseshoe crabs, which suggests chelicerates arose as micropredators early during the Cambrian explosion.

Cambrian deposits bearing exceptional preservation of soft-bodied fossils offer unique insights into the diversity and paleobiology of the Cambrian biota. In recent years, a growing number of studies have provided evidence that highly labile tissues, such as nervous tissues, may be more common in fossils from Burgess Shale-type deposits than previously appreciated. These discoveries have provided novel insights into the anatomy and phylogeny of Cambrian animals and have also challenged the basic understanding of the limits on the resolution of anatomical information that may be captured by Burgess Shale-type preservation. Evidence used to infer primary blood chemistry was recently reported from the iconic Burgess Shale fossil Marrella splendens, the first such report from a Cambrian Lagerstatte. Diminutive crystals of a copper sulfide mineral found in association with Marrella were interpreted as evidence that the animal's blood contained the Cu-bearing protein hemocyanin. Copper minerals in shales in general, and in the Burgess Shale in particular, however, may have a complex geologic history. Using microscopy and SEM-EDS elemental mapping of material from the Walcott Quarry as well as from the Burgess Shale fossil quarry at Marble Canyon, we assess the timing of emplacement of Cu-sulfides in fossil-bearing mudrocks of the Burgess Shale. These data demonstrate that copper minerals are most conspicuously concentrated within veins that are oriented sub-perpendicular to bedding. These late-stage fluid flow paths allowed Cu-mineralizing fluids to infiltrate bedding plane-parallel cracks, which commonly run through Burgess Shale fossils and extend into the matrix around them. Based upon the relationships of mineral phases associated with fossils, it appears that Cu-mineralization occurred after metamorphic volatilization of a large proportion of the carbonaceous material that originally comprised the fossils. We find that copper minerals in the Burgess Shale are of demonstrably post-Cambrian origin, a secondary overprint that cannot be used to infer original blood chemistry or tissue composition of its fossils.

Synopsis We use a series of hydrodynamic experiments on abstracted models to explore whether primitive vertebrates may have swum under various conditions without a clearly-differentiated tail fin. Cambrian vertebrates had post-anal stubby tails, some had single dorsal and ventral fins, but none had yet evolved a clearly differentiated caudal fin typical of post-Cambrian fishes, and must have relied on their long and flexible laterally-compressed bodies for locomotion, i.e., by bending their bodies side-to-side in order to propagate waves from head to tail. We approach this problem experimentally based on an abstracted model of Metaspriggina walcotti from the 506-million-year old Burgess Shale by using oscillating thin flexible plates while varying the tail fin geometry from rectangular to uniform, and finally to a no tail–fin condition. Despite a missing tail fin, this study supports the observation that the abstracted Metaspriggina model can generate a strong propulsive force in cruise conditions, both away from, and near the sea bed (in ground effect). However, when the abstracted Metaspriggina model moves in ground effect, a weaker performance is observed, indicating that Metaspriggina may not necessarily have been optimized for swimming near the sea bed. When considering acceleration from rest, we find that the Metaspriggina model's performance is not significantly different from other morphological models (abstracted truncate tail and abstracted heterocercal tail). Statistical analysis shows that morphological parameters, swimming modes, and ground effect all play significant roles in thrust performance. While the exact relationships of Cambrian vertebrates are still debated, as agnathans, they share some general characteristics with modern cyclostomes, in particular an elongate body akin to lampreys. Lampreys, as anguilliform swimmers, are considered to be some of the most efficient swimmers using a particular type of suction thrust induced by the traveling body wave as it travels from head to tail. Our current experiments suggest that Metaspriggina’s ability in acceleration from rest, through possibly a similar type of suction thrust, which is defined as the ability to generate low pressure on upstream facing sections of the body, might have evolved early in response to increasing predator pressure during the Cambrian Explosion.

Anomalocaris canadensis, a soft-bodied stem-group arthropod from the Burgess Shale, is considered the largest predator of the Cambrian period. Thanks to a series of lateral flexible lobes along its dorso-ventrally compressed body, it is generally regarded as an efficient swimmer, well-adapted to its predatory lifestyle. Previous theoretical hydrodynamic simulations have suggested a possible optimum in swimming performance when the lateral lobes performed as a single undulatory lateral fin, comparable to the pectoral fins in skates and rays. However, the role of the unusual fan-like tail of Anomalocaris has not been previously explored. Swimming efficiency and maneuverability deduced from direct hydrodynamic analysis are here studied in a towing tank facility using a three-vane physical model designed as an abstraction of the tail fin. Through direct force measurements, it was found that the model exhibited a region of steady-state lift and drag enhancement at angles of attack greater than 25° when compared with a triangular-shaped reference model. This would suggest that the resultant normal force on the tail fin of Anomalocaris made it well-suited for turning maneuvers, giving it the ability to turn quickly and through small radii of curvature. These results are consistent with an active predatory lifestyle, although detailed kinematic studies integrating the full organism, including the lateral lobes, would be required to test the effect of the tail fin on overall swimming performance. This study also highlights a possible example of evolutionary convergence between the tails of Anomalocaris and birds, which, in both cases, are well-adapted to efficient turning maneuvers.

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 thatTitanokorysbridges 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 knownPahvantia. Despite distinct carapaces,Titanokorysshares similar rake-like appendages for sediment-sifting withCambroraster,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.

Tokummia katalepsis from the Burgess Shale had a pair of mandibles and maxilliped claws, showing that large bivalved arthropods from the Cambrian period are forerunners of myriapods and pancrustaceans, thereby providing a basis for the origin of the hyperdiverse mandibulate body plan. Fossils from the famous 508-million-year-old Burgess Shale in Canada have been vital for shaping our understanding of the origin and early evolution of arthropods, the group of invertebrate animals recognized by their segmented bodies with jointed limbs and an exoskeleton. In recent years, research has found support for a single group of arthropods known as mandibulates that comprises insects, crustaceans and myriapods (centipedes and millipedes) but excludes chelicerates (spiders, scorpions and their allies). Few fossils have been found to illuminate the earliest mandibulates. Cedric Aria and Jean-Bernard Caron now show that this gap is partially filled by the arrival of the Burgess Shale fossil Tokummia katalepsis, whose anatomy allows the reconstruction of the anatomical and evolutionary history of this important animal group. Retracing the evolutionary history of arthropods has been one of the greatest challenges in biology1. During the past decade, phylogenetic analyses of morphological and molecular data2,3,4 have coalesced towards the conclusion that Mandibulata, the most diverse and abundant group of animals, is a distinct clade from Chelicerata, in that its members possess post-oral head appendages specialized for food processing, notably the mandible2,5. The origin of the mandibulate body plan, however, which encompasses myriapods, crustaceans and hexapods, has remained poorly documented1,6. Here we show that Tokummia katalepsis gen. et sp. nov., a large bivalved arthropod from the 508 million-year-old Marble Canyon fossil deposit (Burgess Shale, British Columbia), has unequivocal mandibulate synapomorphies, including mandibles and maxillipeds, as well as characters typically found in crustaceans, such as enditic, subdivided basipods and ring-shaped trunk segments. Tokummia and its closest relative, Branchiocaris (in Protocarididae, emended), also have an anteriormost structure housing a probable bilobed organ, which could support the appendicular origin of the labrum7. Protocaridids are retrieved with Canadaspis and Odaraia (in Hymenocarina, emended) as part of an expanded mandibulate clade, refuting the idea that these problematic bivalved taxa, as well as other related forms, are representatives of the basalmost euarthropods8,9. Hymenocarines now illustrate that the subdivision of the basipod and the presence of proximal endites are likely to have been ancestral conditions critical for the evolution of coxal and pre-coxal features in mandibulates10,11. The presence of crustaceomorph traits in the Cambrian larvae of various clades basal to Mandibulata is reinterpreted as evidence for the existence of distinct ontogenetic niches among stem arthropods. Larvae would therefore have constituted an important source of morphological novelty during the Cambrian period, and, through heterochronic processes, may have contributed to the rapid acquisition of crown-group characters and thus to greater evolutionary rates during the early radiation of euarthropods12.

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 nereidisgen. 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 nereidisis 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 nereidisis 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.

Exceptional fossils reveal annelid nervous system evolution., Annelid worms are a disparate, primitively segmented clade of bilaterians that first appear during the early Cambrian Period. Reconstructing their early evolution is complicated by the extreme morphological diversity in early diverging lineages, rapid diversification, and sparse fossil record. Canadia spinosa, a Burgess Shale fossil polychaete, is redescribed as having palps with feeding grooves, a dorsal median antenna and biramous parapodia associated with the head and flanking a ventral mouth. Carbonaceously preserved features are identified as a terminal brain, circumoral connectives, a midventral ganglionated nerve cord and prominent parapodial nerves. Phylogenetic analysis recovers neuroanatomically simple extant taxa as the sister group of other annelids, but the phylogenetic position of Canadia suggests that the annelid ancestor was reasonably complex neuroanatomically and that reduction of the nervous system occurred several times independently in the subsequent 500 million years of annelid evolution.

The in situ preservation of animal behaviour in the fossil record is exceedingly rare, but can lead to unique macroecological and macroevolutionary insights, especially regarding early representatives of major animal clades. We describe a new complex ecological relationship from the middle Cambrian Burgess Shale (Raymond Quarry, Canada). More than 30 organic tubes were recorded with multiple enteropneust and polychaete worms preserved within them. Based on the tubicolous nature of fossil enteropneusts, we suggest that they were the tube builders while the co-preserved polychaetes were commensals. These findings mark, to our knowledge, the first record of commensalism within Annelida and Hemichordata in the entire fossil record. The finding of multiple enteropneusts sharing common tubes suggests that either the tubes represent reproductive structures built by larger adults, and the enteropneusts commonly preserved within are juveniles, or these enteropneusts were living as a pseudo-colony without obligate attachment to each other, and the tube was built collaboratively. While neither hypothesis can be ruled out, gregarious behaviour was clearly an early trait of both hemichordates and annelids. Further, commensal symbioses in the Cambrian may be more common than currently recognized.

Bayesian phylogenetic analysis using a Markov k model on a morphological dataset with 90 taxa and 213 characters (see Methods). Numbers next to nodes are posterior probabilities. Yellow box highlights the group Hymenocarina. Loricicaris may represent a junior synonym of Protocaris (3). Green box highlights the group Hymenocarina.

Bayesian analysis using a Markov k model on a morphological dataset with 90 taxa and 209 characters (without characters 206, 207, 208, 209 and 205 as originally defined (28) (see Methods, See Supplementary Material 3). Numbers next to nodes are posterior probabilities. Loricicaris may represent a junior synonym of Protocaris (3).

Size of the measured specimens (n=50), lengths and heights (mm). Most specimens follow a similar proportion of length-width. Although some variation is present regarding widths, the number of measured specimens is too small to detect any morphotype patterns. The specimen that appears as an outlier (at 15L, 10H) is ROMIP 65386.

Clusters of individuals of Fibulacaris nereidis, preserved in multiple views ROMIP 65457. A) Part. B) Counterpart. C) Telson of specimen 4, figured in Figure 4 C. The total number is approximately 11 individuals, as indicated. Abbreviations: ca) carapace, c.r) caudal rami, ro) rostrum, th) thorax.

Three-dimensional internal structures and comparisons with other arthropods from the Burgess Shale. A) Odaraia, lateral view ROMIP 61121. Odaraia was also referred as an invert swimmer and suspension feeder and shares multiple traits with Fibulacaris nereidis, including the absence of developed antennae and a highly multisegmented body. B, C) F. nereidis, ROMIP 65363, focus on the indeterminate rostrum tissue. D) Elemental maps of F. nereidis, ROMIP 65363: aluminium, carbon, calcium, phosphorous. E) Elemental maps of Marrella splendens ROMIP 60748: carbon, phosphorous. The tissue seen in the cephalic spines of M. splendens could represent cavities or unrecognised organs (79), and may have been similar to the internal tissue on the rostrum of F. nereidis. Abbreviations: ca) carapace, ey) eye, gu) gut, in?) indeterminate, le) legs, ro) rostrum. Scale bars: 10 mm (A), 2 mm (B), 1 mm (C, D, E).

Includes: Supp. Material 1-Specimens figured, measured or referenced. Supp. Material 2: added taxa. Supp. Material 3: list character matrix. Supp. Material 4: applied constraints

Brood care, including the carrying of eggs or juveniles, is a form of parental care, which, like other parental traits [1], enhances offspring fitness with variable costs and benefits to the parents [2]. Attempts to understand why and how parental care evolved independently in numerous animal groups often emphasize the role of environmental pressures such as predation, ephemeral resources, and, more generally, the harshness of environment. The fossil record can, in principle, provide minimum age constraints on the evolution of life-history traits, including brood care and key information on the reproductive strategies of extinct organisms. New, exceptionally preserved specimens of the weakly sclerotized arthropod Waptia fieldensis from the middle Cambrian (ca. 508 million years ago) Burgess Shale, Canada, provide the oldest example of in situ eggs with preserved embryos in the fossil record. The relatively small clutch size, up to 24 eggs, and the relatively large diameter of individual eggs, some over 2 mm, contrast with the high number of small eggs-found without preserved embryos-in the bivalved bradoriid arthropod Kunmingella douvillei from the Chengjiang biota (ca. 515 million years ago). The presence of these two different parental strategies suggests a rapid evolution of a variety of modern-type life-history traits, including extended investment in offspring survivorship, soon after the Cambrian emergence of animals. Together with previously described brooded eggs in ostracods from the Upper Ordovician (ca. 450 million years ago), these new findings suggest that the presence of a bivalved carapace played a key role in the early evolution of parental care in arthropods.

The Tulip Beds locality on Mount Stephen (Yoho National Park, British Columbia) yields one of the most abundant and diverse (~10,000 specimens in 110 taxa) Burgess Shale fossil assemblages in the Canadian Rockies. Detailed semi quantitative and quantitative analyses of this assemblage suggest strong similarities with the Walcott Quarry on Fossil Ridge. Both assemblages are dominated by epibenthic, sessile, and suspension feeding taxa, mostly represented by arthropods and sponges and have comparable diversity patterns, despite sharing only about half the genera. However, the Tulip Beds has a higher relative abundance of suspension feeders and taxa of unknown affinity compared to the Walcott Quarry. These biotic variations are probably largely attributable to ecological and evolutionary differences between the two temporally distinct communities that adapted to similar, but not identical, environmental settings. For instance, the Tulip Beds is farther away from the Cathedral Escarpment than the Walcott Quarry. The Tulip Beds and Walcott Quarry assemblages are more similar to each other than either one is to the assemblages of the Chengjiang biota, although the relative diversity of major taxonomic groups and ecological patterns are similar in all assemblages. The conserved diversity patterns and ecological structures among sites suggest that the ecological composition of Cambrian Burgess Shale-type communities was relatively stable across wide geographic and temporal scales.

Yuknessia Walcott, 1919 recently was transferred from the green algae to the Phylum Hemichordata on the basis of new details observed for the type species, Y. simplex, from the Burgess Shale Formation (Cambrian Stage 5) of British Columbia. This has prompted reexamination of material attributed to Yuknessia from various Cambrian localities in South China. Findings preclude both a Yuknessia and a hemichordate affinity for all of the Chinese study material, and most of this material is formally transferred to Fuxianospira Chen and Zhou, 1997, a taxon common in the Chengjiang biota. Comparable material from the Cambrian Marjum, Wheeler, and Burgess Shale formations of North America is also assigned to Fuxianospira, and this reassignment expands both the paleogeographic and stratigraphic range of this taxon. All aspects of the study specimens, including details obtained from scanning electron microscopy, are consistent with an algal affinity, as proposed in the original descriptions of the Chinese material.

The molecularly defined clade Ecdysozoa1 comprises the panarthropods (Euarthropoda, Onychophora and Tardigrada) and the cycloneuralian worms (Nematoda, Nematomorpha, Priapulida, Loricifera and Kinorhyncha). These disparate phyla are united by their means of moulting, but otherwise share few morphological characters—none of which has a meaningful fossilization potential. As such, the early evolutionary history of the group as a whole is largely uncharted. Here we redescribe the 508-million-year-old stem-group onychophoran Hallucigenia sparsa2, 3, 4, 5, 6 from the mid-Cambrian Burgess Shale. We document an elongate head with a pair of simple eyes, a terminal buccal chamber containing a radial array of sclerotized elements, and a differentiated foregut that is lined with acicular teeth. The radial elements and pharyngeal teeth resemble the sclerotized circumoral elements and pharyngeal teeth expressed in tardigrades7, 8, 9, stem-group euarthropods10, 11, 12 and cycloneuralian worms13. Phylogenetic results indicate that equivalent structures characterized the ancestral panarthropod and, seemingly, the ancestral ecdysozoan, demonstrating the deep homology of panarthropod and cycloneuralian mouthparts, and providing an anatomical synapomorphy for the ecdysozoan supergroup.

Decay experiments are becoming a more widespread tool in evaluating the fidelity of the fossil record. Character interpretations of fossil specimens stand to benefit from an understanding of how decay can result in changes in morphology and, potentially, total character loss. We performed a decay experiment for the Class Enteropneusta to test the validity of anatomical interpretations of the Burgess Shale enteropneust Spartobranchus tenuis and to determine how the preservation of morphological features compares with the sequence of character decay in extant analogues. We used three species of enteropneust (Saccoglossus pusillus, Harrimania planktophilus, and Balanoglossus occidentalis) representing the two major families of Enteropneusta. Comparisons between decay sequences suggest that morphological characters decay in a consistent and predictable manner within Enteropneusta, and do not support the hypothesis of stemward slippage. The gill bars and nuchal skeleton were the most decay resistant, whereas the gill pores and pre-oral ciliary organ were unequivocally the most decay prone. Decay patterns support the identification of the nuchal skeleton, gill bars, esophageal organ, trunk, and proboscis in Spartobranchus tenuis and corroborate a harrimaniid affinity. Bias due to the taphonomic loss of taxonomically informative characters is unlikely. The morphologically simple harrimaniid body plan can be seen, therefore, to be plesiomorphic within the enteropneusts. Discrepancies between the sequence of decay in a laboratory setting and fossil preservation also exist. These discrepancies are highlighted not to discredit the use of modern decay studies but rather to underline their non-actualistic nature. Paleoenvironmental variables besides decay, such as the timeframe between death and early diagenesis as well as postmortem transport, are discussed relative to decay data. These experiments reinforce the strength of a comprehensive understanding of decay sequences as a benchmark against which to describe fossil taxa and understand the conditions leading to fossilization.

Characterized by atypical frontalmost appendages, leanchoiliids are early arthropods whose phylogenetic placement has been much debated. Morphological interpretations have differed, some of which concern critical characters such as the number of eyes and head appendages, but methodological approaches also have diverged. Here, we describe a new leanchoiliid, Yawunik kootenayi gen. et sp. nov., based on 42 specimens from the newly discovered Marble Canyon locality of the Burgess Shale (Kootenay National Park, British Columbia; middle Cambrian). This new morphotype demonstrates the presence of a four-segmented head in leanchoiliids, along with two small antero-median eyes in addition to lateral eyes. Yawunik is characterized by a 12-segmented trunk and a carinate, lanceolate telson adorned with minute spines. The ‘great appendages’ of the animal bear teeth on their two distal rami, which would have enhanced their ability to grasp prey. Attitudes of specimens, resulting from burial at multiple aspects of bedding, suggest the ‘great appendages’ were flexible and capable of antero-posterior rotation. We also discuss the nature of intersegmental tissues and filaments present within the ‘great appendages’. Our phylogenetic analyses extend the monophyly of leanchoiliids to include Haikoucaris and Yohoia in a new clade, the Cheiromorpha nom. nov. (within Heptopodomera nom. nov.). Other nodes are poorly resolved unless implied weights are used, and in this case, the topology is critically sensitive to the coding prerogative of inapplicable states (NAs). Both the traditional ‘Arachnomorpha’ hypothesis (NAs as additional states) and the more recently favoured ‘Artiopoda + Crustacea’ (NAs as uncertainties) were obtained using the same data set and outgroup. This result stresses, first, the historical importance of polarization over data content in scenarios of early arthropod evolution, and second, a pressing need to investigate the impacts of coding inapplicables, especially given the inflating effect of implied weights.

S1- W. fieldensis, geographic and stratigraphic distribution. S2- Nebalia bipes (Phyllocarida), decay experiments. S3- W. fieldensis, fossil assemblage. S4- W. fieldensis, synonymy list. S5- W. fieldensis, type specimens. S6- W. fieldensis, measurements (1). S7- Nebalia bipes (Phyllocarida), morphology. S8- W. fieldensis, frontal part. S9- Vargula hilgendorfii (Ostracoda), labrum. S10- Cirolana harfordi (Isopoda), mandibles. S11- W. fieldensis, mandibles and maxillules. S12- W. fieldensis, post-maxillular cephalothoracic appendages. S13- W. fieldensis, post-maxillular cephalothoracic appendages (frontal view). S14- W. fieldensis, lamellate appendages. S15- Crangon crangon (Decapoda), gills. S16- W. fieldensis, measurements (2). S17- W. fieldensis, digestive system (1). S18- W. fieldensis, digestive system (2). S19- Cladogram (adult taxa). S20- List of characters. S22- Cladogram (larval and adult taxa).

Electronic Supplementary Material Figure 2. Cross sections of a Tuzoia carapace. ROM 64825, Stanley Glacier, Kootenay National Park. (a) Low angle light emphasizing the presence of burrows. (b) Specimens photographed using cross-polarized light after cuts – the line in red represents area mapped. (c) Polished cross-section of the part and counterpart. The carapace occurs at the interface along an horizontal yellow zone which likely represents a weathering effect. (d) Maps of Ca, Ca+Si and Ca+Fe showing a weathered layer poor in Ca (framed area in c). (e) Different elemental maps showing no evidence of variations in mineralogy between the traces fossils and the matrix. The bright carbon zone at the interface between the part and the counterpart is glue holding the two pieces together. The carapace itself is not visible, and the Carbon film has probably weathered out or is too thin to be detected in cross-section. Note that part of the cut probably included both valves, but it is impossible to tell based on cross sections. Scale bars are 1 cm in (a, b) 1 mm in (c) and 0.5 mm in (d, e).

Electronic Supplementary Material Figure 1. Trace fossils associated with Tuzoia carapaces. (a) General view of two specimens of Tuzoia sp.; annulated burrow in fragmented specimen (white box), ROM 64814, Tuzoia Bed, Fossil Ridge. (b) Close-up of annulated burrow (arrow) shown in a. Scale bar is 0.5 cm. (c) General view of a high-density trace-fossil assemblage associated with Tuzoia sp. Note two clear sets of trace fossils: thin simple trails and burrows mostly confined to the carapace outline and U-shaped burrows clearly visible as positive ridges. These two distinctive set of structures are interpreted as recording a palimpsest assemblage. Although polarity of the bed was not recorded (this is talus material), based on the preservation of U-shaped burrows it is suggested that this surface records a bed sole. Accordingly, U-shaped burrows are overlying the Tuzoia carapace and possibly superimposed due to the compactional effect. U-shaped burrows, although not as visible, are also present in the host rock. ROM 57533, Walcott Talus, Fossil Ridge. Specimen most likely coming from levels of the Raymond Quarry or higher in the stratigraphic section. Scale bar is 1 cm.

Waptia fieldensisWalcott, 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 thatW. fieldensisshares 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.Waptiaconfirms 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 thatWaptiawas 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.