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Cnidarians are regarded as one of the earliest-diverging animal phyla. One of the hallmarks of the cnidarian body plan is the evolution of a free-swimming medusa in some medusozoan classes, but the origin of this innovation remains poorly constrained by the fossil record and molecular data. Previously described macrofossils, putatively representing medusa stages of crown-group medusozoans from the Cambrian of Utah and South China, are here reinterpreted as ctenophore-grade organisms. Other putative Ediacaran to Cambrian medusozoan fossils consist mainly of microfossils and tubular forms. Here we describe Burgessomedusa phasmiformis gen. et sp. nov., the oldest unequivocal macroscopic free-swimming medusa in the fossil record. Our study is based on 182 exceptionally preserved body fossils from the middle Cambrian Burgess Shale (Raymond Quarry, British Columbia, Canada). Burgessomedusa possesses a cuboidal umbrella up to 20 cm high and over 90 short, finger-like tentacles. Phylogenetic analysis supports a medusozoan affinity, most likely as a stem group to Cubozoa or Acraspeda (a group including Staurozoa, Cubozoa and Scyphozoa). Burgessomedusa demonstrates an ancient origin for the free-swimming medusa life stage and supports a growing number of studies showing an early evolutionary diversification of Medusozoa, including of the crown group, during the late Precambrian–Cambrian transition. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [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]

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-megacheirans Cambrian taxa also illustrates that the specialization of the head as the dedicated foodprocessing 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. [ABSTRACT FROM AUTHOR]

Background: Arthropoda, Tardigrada and Onychophora evolved from lobopodians, a paraphyletic group of disparate Palaeozoic vermiform animals with soft legs. Although the morphological diversity that this group encompasses likely illustrates the importance of niche diversification in the early radiation of panarthropods, the ecology of lobopodians remains poorly characterized. Results: Here we describe a new luolishaniid taxon from the middle Cambrian Burgess Shale (Walcott Quarry) in British Columbia, Canada, whose specialized morphology epitomizes the suspension-feeding ecology of this clade, and is convergent with some modern marine animals, such as caprellid crustaceans. This species possesses two long pairs and four shorter pairs of elongate spinose lobopods at the front, each bearing two slender claws, and three pairs of stout lobopods bearing single, strong, hook-like anterior-facing claws at the back. The trunk is remarkably bare, widening rearwards, and, at the front, extends beyond the first pair of lobopods into a small "head" bearing a pair of visual organs and a short proboscis with numerous teeth. Based on a critical reappraisal of character coding in lobopodians and using Bayesian and parsimony-based tree searches, two alternative scenarios for early panarthropod evolution are retrieved. In both cases, hallucigeniids and luolishaniids are found to be extinct radiative stem group panarthropods, in contrast to previous analyses supporting a position of hallucigeniids as part of total-group Onychophora. Our Bayesian topology finds luolishaniids and hallucigeniids to form two successive clades at the base of Panarthropoda. Disparity analyses suggest that luolishaniids, hallucigeniids and total-group Onychophora each occupy a distinct region of morphospace. Conclusions: Hallucigeniids and luolishaniids were comparably diverse and successful, representing two major lobopodian clades in the early Palaeozoic, and both evolved body plans adapted to different forms of suspension feeding. A Bayesian approach to cladistics supports the view that a semi-sessile, suspension-feeding lifestyle characterized the origin and rise of Panarthropoda from cycloneuralian body plans. [ABSTRACT FROM AUTHOR]

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. [Display omitted] • The radiodont Stanleycaris is reconstructed from a collection of exquisite fossils • Stanleycaris has three eyes, demonstrating evolutionary continuity of median eyes • Radiodont frontal appendages are innervated by the second brain neuromere • Fossils support early differentiation between arthropod head and trunk segmentation Many early arthropods are known only from rare, fragmentary fossils. Moysiuk and Caron describe hundreds of new, exceptional Cambrian fossils of the radiodont Stanleycaris hirpex , revealing surprising features such as a median eye and preserved neuroanatomy. They set out a new theory for the evolution of key features of the arthropod body plan. [ABSTRACT FROM AUTHOR]

Background: The combination of a meager fossil record of vermiform enteropneusts and their disparity with the tubicolous pterobranchs renders early hemichordate evolution conjectural. The middle Cambrian Oesia disjuncta from the Burgess Shale has been compared to annelids, tunicates and chaetognaths, but on the basis of abundant new material is now identified as a primitive hemichordate. Results: Notable features include a facultative tubicolous habit, a posterior grasping structure and an extensive pharynx. These characters, along with the spirally arranged openings in the associated organic tube (previously assigned to the green alga Margaretia), confirm Oesia as a tiered suspension feeder. Conclusions: Increasing predation pressure was probably one of the main causes of a transition to the infauna. In crown group enteropneusts this was accompanied by a loss of the tube and reduction in gill bars, with a corresponding shift to deposit feeding. The posterior grasping structure may represent an ancestral precursor to the pterobranch stolon, so facilitating their colonial lifestyle. The focus on suspension feeding as a primary mode of life amongst the basal hemichordates adds further evidence to the hypothesis that suspension feeding is the ancestral state for the major clade Deuterostomia. [ABSTRACT FROM AUTHOR]

Summary 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. [ABSTRACT FROM AUTHOR]

The molecularly defined clade Ecdysozoa 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 sparsa 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 tardigrades, stem-group euarthropods and cycloneuralian worms. 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

We herein describe Surusicaris elegans gen. et sp. nov. (in Isoxyidae, amended), a middle (Series 3, Stage 5) Cambrian bivalved arthropod from the new Burgess Shale deposit of Marble Canyon (Kootenay National Park, British Columbia). Surusicaris exhibits 12 simple, partly undivided biramous trunk limbs with long tripartite caeca, which may illustrate a plesiomorphic “fused” condition of exopod and endopod. We construe also that the head is made of five somites (= four segments), including two eyes, one pair of anomalocaridid-like frontalmost appendages, and three pairs of poorly sclerotized uniramous limbs. This fossil may therefore be a candidate for illustrating the origin of the plesiomorphic head condition in euarthropods, and questions the significance of the “two-segmented head” in, e.g., fuxianhuiids. The frontalmost appendage in isoxyids is intriguingly disparate, bearing similarities with both dinocaridids and euarthropods. In order to evaluate the relative importance of bivalved arthropods, such as Surusicaris, in the hypothetical structuro-functional transition between the dinocaridid frontal appendage and the pre-oral—arguably deutocerebral—appendage of euarthropods, we chose a phenetic approach and computed morphospace occupancy for the frontalmost appendages of 36 stem and crown taxa. Results show different levels of evolutionary decoupling between frontalmost appendage disparity and body plans. Variance is greatest in dinocaridids and “stem bivalved” arthropods, but these groups do not occupy the morphospace homogeneously. Rather, the diversity of frontalmost appendages in “stem bivalved” arthropods, distinct in its absence of clear clustering, is found to link the morphologies of “short great appendages,” chelicerae and antennules. This find fits the hypothesis of an increase in disparity of the deutocerebral appendage prior to its diversification in euarthropods, and possibly corresponds to its original time of development. The analysis of this pattern, however, is sensitive to the—still unclear—extent of polyphyly of the “stem bivalved” taxa. [ABSTRACT FROM AUTHOR]

To investigate the phylogenetic affinity of Yuknessia simplex Walcott, 1919, scanning electron microscopy was applied to the Burgess Shale (Cambrian Series 3, Stage 5) type material and to new material from the Trilobite Beds (Yoho National Park) and specimens from the Cambrian of Utah. On the basis of fine-scale details observed using this approach, including banding structure interpreted as fusellae, Yuknessia Walcott, 1919 is transferred from the algae, where it resided for nearly a century, to the extant taxon Pterobranchia (Phylum Hemichordata). Considered as such, Yuknessia specimens from the Trilobite Beds and Spence Formation (Utah) are amongst the oldest known colonial pterobranchs. Two morphs regarded herein as two different species are recognized from the Trilobite Beds based on tubarium morphology. Yuknessia simplex has slender erect tubes whereas Yuknessia stephenensis n. sp., which is also known in Utah, has more robust erect tubes. The two paratypes of Y. simplex designated by Walcott (1919) are formally removed from Yuknessia and are reinterpreted respectively as an indeterminate alga and Dalyia racemata Walcott, 1919, a putative red alga. [ABSTRACT FROM AUTHOR]

Knowledge of the early evolution of fish largely depends on soft-bodied material from the Lower (Series 2) Cambrian period of South China. Owing to the rarity of some of these forms and a general lack of comparative material from other deposits, interpretations of various features remain controversial, as do their wider relationships amongst post-Cambrian early un-skeletonized jawless vertebrates. Here we redescribe Metaspriggina on the basis of new material from the Burgess Shale and exceptionally preserved material collected near Marble Canyon, British Columbia, and three other Cambrian Burgess Shale-type deposits from Laurentia. This primitive fish displays unambiguous vertebrate features: a notochord, a pair of prominent camera-type eyes, paired nasal sacs, possible cranium and arcualia, W-shaped myomeres, and a post-anal tail. A striking feature is the branchial area with an array of bipartite bars. Apart from the anterior-most bar, which appears to be slightly thicker, each is associated with externally located gills, possibly housed in pouches. Phylogenetic analysis places Metaspriggina as a basal vertebrate, apparently close to the Chengjiang taxa Haikouichthys and Myllokunmingia, demonstrating also that this primitive group of fish was cosmopolitan during Lower-Middle Cambrian times (Series 2-3). However, the arrangement of the branchial region in Metaspriggina has wider implications for reconstructing the morphology of the primitive vertebrate. Each bipartite bar is identified as being respectively equivalent to an epibranchial and ceratobranchial. This configuration suggests that a bipartite arrangement is primitive and reinforces the view that the branchial basket of lampreys is probably derived. Other features of Metaspriggina, including the external position of the gills and possible absence of a gill opposite the more robust anterior-most bar, are characteristic of gnathostomes and so may be primitive within vertebrates. [ABSTRACT FROM AUTHOR]

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian ( Series 3, Stage 5) Burgess Shale Formation ( Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton-benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown-group arthropods), were found to have occurred among basal bivalved arthropods. [ABSTRACT FROM AUTHOR]

Background: The Burgess Shale is well known for its preservation of a diverse soft-bodied biota dating from the Cambrian period (Series 3, Stage 5). While previous paleoecological studies have focused on particular species (autecology) or entire paleocommunities (synecology), studies on the ecology of populations (demecology) of Burgess Shale organisms have remained mainly anecdotal. Results: Here, we present evidence for mass molting events in two unrelated arthropods from the Burgess Shale Walcott Quarry, Canadaspis perfecta and a megacheiran referred to as Alalcomenaeus sp. Conclusions: These findings suggest that the triggers for such supposed synchronized molting appeared early on during the Cambrian radiation, and synchronized molting in the Cambrian may have had similar functions in the past as it does today. In addition, the finding of numerous juvenile Alalcomenaeus sp. molts associated with the putative alga Dictyophycus suggests a possible nursery habitat. In this nursery habitat a population of this animal might have found a more protected environment in which to spend critical developmental phases, as do many modern species today. [ABSTRACT FROM AUTHOR]

Hemichordates are a marine group that, apart from one monospecific pelagic larval form, are represented by the vermiform enteropneusts and minute colonial tube-dwelling pterobranchs. Together with echinoderms, they comprise the clade Ambulacraria. Despite their restricted diversity, hemichordates provide important insights into early deuterostome evolution, notably because of their pharyngeal gill slits. Hemichordate phylogeny has long remained problematic, not least because the nature of any transitional form that might serve to link the anatomically disparate enteropneusts and pterobranchs is conjectural. Hence, inter-relationships have also remained controversial. For example, pterobranchs have sometimes been compared to ancestral echinoderms. Molecular data identify enteropneusts as paraphyletic, and harrimaniids as the sister group of pterobranchs. Recent molecular phylogenies suggest that enteropneusts are probably basal within hemichordates, contrary to previous views, but otherwise provide little guidance as to the nature of the primitive hemichordate. In addition, the hemichordate fossil record is almost entirely restricted to peridermal skeletons of pterobranchs, notably graptolites. Owing to their low preservational potentials, fossil enteropneusts are exceedingly rare, and throw no light on either hemichordate phylogeny or the proposed harrimaniid-pterobranch transition. Here we describe an enteropneust, Spartobranchus tenuis (Walcott, 1911), from the Middle Cambrian-period (Series 3, Stage 5) Burgess Shale. It is remarkably similar to the extant harrimaniids, but differs from all known enteropneusts in that it is associated with a fibrous tube that is sometimes branched. We suggest that this is the precursor of the pterobranch periderm, and supports the hypothesis that pterobranchs are miniaturized and derived from an enteropneust-like worm. It also shows that the periderm was acquired before size reduction and acquisition of feeding tentacles, and that coloniality emerged through aggregation of individuals, perhaps similar to the Cambrian rhabdopleurid Fasciculitubus. The presence of both enteropneusts and pterobranchs in Middle Cambrian strata, suggests that hemichordates originated at the onset of the Cambrian explosion. [ABSTRACT FROM AUTHOR]

The Middle Cambrian Pikaia gracilens (Walcott) has an iconic position as a Cambrian chordate, but until now no detailed description has been available. Here on the basis of the 114 available specimens we review its anatomy, confirm its place in the chordates and explore with varying degrees of confidence its relationships to both extant and extinct chordates and other deuterostomes. The body of Pikaia is fusiform, laterally compressed and possesses about 100 myomeres. The head is small, bilobed and bears two narrow tentacles. There is no evidence for eyes. Apart from a thin dorsal fin (without finrays) and a series of at least nine bilaterally arranged appendages with possible pharyngeal pores at the anterior end, there are no other external features. In addition to the musculature the internal anatomy includes an alimentary canal, the anterior of which forms a prominent lenticular unit that is almost invariably preserved in positive relief. The cavity is interpreted as pharyngeal, implying that the mouth itself was almost terminal. The posterior extension of the gut is unclear although the anus appears to have been terminal. The most prominent internal structure is a reflectively preserved unit, possibly hollow, termed here the dorsal organ. Although formerly interpreted as a notochord its position and size make this less likely. Its original function remains uncertain, but it could have formed a storage organ. Ventral to the dorsal organ a narrower strand of tissue is interpreted as representing the nerve chord and notochord. In addition to these structures, there is also evidence for a vascular system, including a ventral blood vessel. The position of Pikaia in the chordates is largely based on the presence of sigmoidal myomeres, and the more tentative identification of a notochord. In many other respects, Pikaia differs from the expected nature of primitive chordates, especially as revealed in amphioxus and the Cambrian record (including Cathaymyrus, Haikouichthys, Metaspriggina, Myllokunmingia, and Zhongxiniscus). Whilst the possibility that Pikaia is simply convergent on the chordates cannot be dismissed, we prefer to build a scenario that regards Pikaia as the most stem-ward of the chordates with links to the phylogenetically controversial yunnanozoans. This hypothesis has implications for the evolution of the myomeres, notochord and gills. Finally, the wealth of material of Pikaia indicates that, although by definition there must be some sort of taphonomic imprint, the consistency of preservational details allows a reliable reconstruction of the anatomy and does not significantly erode phylogenetically relevant characters. [ABSTRACT FROM AUTHOR]

Burgess Shale-type deposits provide invaluable insights into the early evolution of body plans and the ecological structure of Cambrian communities, but a number of species, continue to defy phylogenetic interpretations. Here we extend this list to include a new soft-bodied animal, Siphusauctum gregarium n. gen. and n. sp., from the Tulip Beds (Campsite Cliff Shale Member, Burgess Shale Formation) of Mount Stephen (Yoho National Park, British Columbia). With 1,133 specimens collected, S. gregarium is clearly the most abundant animal from this locality. This stalked animal (reaching at least 20 cm in length), has a large ovoid calyx connected to a narrow bilayered stem and a small flattened or bulb-like holdfast. The calyx is enclosed by a flexible sheath with six small openings at the base, and a central terminal anus near the top encircled by indistinct openings. A prominent organ, represented by six radially symmetrical segments with comb-like elements, surrounds an internal body cavity with a large stomach, conical median gut and straight intestine. Siphusauctum gregarium was probably an active filter-feeder, with water passing through the calyx openings, capturing food particles with its comblike elements. It often occurs in large assemblages on single bedding planes suggesting a gregarious lifestyle, with the animal living in high tier clusters. These were probably buried en masse more or less in-situ by rapid mud flow events. Siphusauctum gregarium resembles Dinomischus, another Cambrian enigmatic stalked animal. Principal points of comparison include a long stem with a calyx containing a visceral mass and bract-like elements, and a similar lifestyle albeit occupying different tiering levels. The presence in both animals of a digestive tract with a potential stomach and anus suggest a grade of organization within bilaterians, but relationships with extant phyla are not straightforward. Thus, the broader affinities of S. gregarium remain largely unconstrained. [ABSTRACT FROM AUTHOR]

The article offers information on the objection of D. Mazurek and M. Zaton on Nectocaris. It says that both Mazurek and Zaton objected to the classification due to the differences of crown-group cephalopods and Nectocaris. They claimed that their differences are tangential to the affinity of Nectocaris.

A new Burgess Shale-type assemblage, from the Stephen Formation of the southern Canadian Rocky Mountains, is described herein. It occurs near Stanley Glacier in Kootenay National Park, 40 km southeast of the type area near Field, British Columbia. While at least a dozen Burgess Shale localities are known from the "thick" Stephen Formation, the Stanley Glacier locality represents the first discovery of Burgess Shale-type fossils from the "thin" Stephen Formation. The Cathedral Escarpment, an important regional paleotopographic feature, has been considered important to the paleoecologic setting and the preservation of the Burgess Shale biota. However, the Stanley Glacier assemblage was preserved in a distal ramp setting in a region where no evidence of an escarpment is present. The low-diversity assemblage contains eight new soft-bodied taxa, including the anomalocaridid Stanleycaris hirpex n. gen., n. sp. (new genus, new species). Nektonic or nektobenthic predators represent the most diverse group, whereas in relative abundance, the assemblage is dominated by typical Cambrian shelly benthic taxa. The low diversity of both the benthic taxa and the ichnofauna, which includes diminutive trace fossils associated with carapaces of soft-bodied arthropods, suggests a paleoenvironment with restrictive conditions. The Stanley Glacier assemblage expands the temporal and geographic range of the Burgess Shale biota in the southern Canadian Rockies, and suggests that Burgess Shale-type assemblages may be common in the "thin" Stephen Formation, which is regionally widespread. [ABSTRACT FROM AUTHOR]

The exquisite preservation of soft-bodied animals in Burgess Shale-type deposits provides important clues into the early evolution of body plans that emerged during the Cambrian explosion. Until now, such deposits have remained silent regarding the early evolution of extant molluscan lineages—in particular the cephalopods. Nautiloids, traditionally considered basal within the cephalopods, are generally depicted as evolving from a creeping Cambrian ancestor whose dorsal shell afforded protection and buoyancy. Although nautiloid-like shells occur from the Late Cambrian onwards, the fossil record provides little constraint on this model, or indeed on the early evolution of cephalopods. Here, we reinterpret the problematic Middle Cambrian animal Nectocaris pteryx as a primitive (that is, stem-group), non-mineralized cephalopod, based on new material from the Burgess Shale. Together with Nectocaris, the problematic Lower Cambrian taxa Petalilium and (probably) Vetustovermis form a distinctive clade, Nectocarididae, characterized by an open axial cavity with paired gills, wide lateral fins, a single pair of long, prehensile tentacles, a pair of non-faceted eyes on short stalks, and a large, flexible anterior funnel. This clade extends the cephalopods’ fossil record by over 30 million years, and indicates that primitive cephalopods lacked a mineralized shell, were hyperbenthic, and were presumably carnivorous. The presence of a funnel suggests that jet propulsion evolved in cephalopods before the acquisition of a shell. The explosive diversification of mineralized cephalopods in the Ordovician may have an understated Cambrian ‘fuse’. [ABSTRACT FROM AUTHOR]

Molecular and morphological evidence unite the hemichordates and echinoderms as the Ambulacraria, but their earliest history remains almost entirely conjectural. This is on account of the morphological disparity of the ambulacrarians and a paucity of obvious stem-groups. We describe here a new taxon Herpetogaster collinsi gen. et sp. nov. from the Burgess Shale (Middle Cambrian) Lagerstätte. This soft-bodied vermiform animal has a pair of elongate dendritic oral tentacles, a flexible stolon with an attachment disc, and a re-curved trunk with at least 13 segments that is directed dextrally. A differentiated but un-looped gut is enclosed in a sac suspended by mesenteries. It consists of a short pharynx, a conspicuous lenticular stomach, followed by a narrow intestine sub-equal in length. This new taxon, together with the Lower Cambrian Phlogites and more intriguingly the hitherto enigmatic discoidal eldoniids (Cambrian-Devonian), form a distinctive clade (herein the cambroernids). Although one hypothesis of their relationships would look to the lophotrochozoans (specifically the entoprocts), we suggest that the evidence is more consistent with their being primitive deuterostomes, with specific comparisons being made to the pterobranch hemichordates and pre-radial echinoderms. On this basis some of the earliest ambulacrarians are interpreted as soft-bodied animals with a muscular stalk, and possessing prominent tentacles. [ABSTRACT FROM AUTHOR]

Abstract: To better understand temporal variations in species diversity and composition, ecological attributes, and environmental influences for the Middle Cambrian Burgess Shale community, we studied 50,900 fossil specimens belonging to 158 genera (mostly monospecific and non-biomineralized) representing 17 major taxonomic groups and 17 ecological categories. Fossils were collected in situ from within 26 massive siliciclastic mudstone beds of the Greater Phyllopod Bed (Walcott Quarry — Fossil Ridge). Previous taphonomic studies have demonstrated that each bed represents a single obrution event capturing a predominantly benthic community represented by census- and time-averaged assemblages, preserved within habitat. The Greater Phyllopod Bed (GPB) corresponds to an estimated depositional interval of 10 to 100 KA and thus potentially preserves community patterns in ecological and short-term evolutionary time. The community is dominated by epibenthic vagile deposit feeders and sessile suspension feeders, represented primarily by arthropods and sponges. Most species are characterized by low abundance and short stratigraphic range and usually do not recur through the section. It is likely that these are stenotopic forms (i.e., tolerant of a narrow range of habitats, or having a narrow geographical distribution). The few recurrent species tend to be numerically abundant and may represent eurytopic organisms (i.e., tolerant of a wide range of habitats, or having a wide geographical distribution). Rarefaction curves demonstrate variation in species richness through the GPB and suggest that more stenotopic species could still be discovered with additional sampling. Comparisons between richness and evenness trends suggest that the community is relatively stable overall, despite gradual species turnover through time, especially in assemblages from younger beds. Less diverse assemblages with low species evenness possibly represent the onset of less favourable environmental conditions. Fossil occurrences in individual beds were analysed using a range of statistical techniques (Correspondence Analysis, Canonical Correspondence Analysis, Minimum Spanning Tree, Indicator Species Analysis, Mantel Test) to extract community patterns. Results suggest the presence of four fossil assemblages based on distinct species associations. The different assemblages presumably reflect variations in environmental and ecological conditions, some acting through time, leading to species turnover. “Disturbances” (e.g., changes in paleo-redox conditions), differences in substrate firmness, and limited taphonomic biases are probably the main factors contributing to community structure. The influence of ecological factors, however, is also predicted from non-random patterns of species recurrences in successive events. Preliminary comparisons with Lower Cambrian Chengjiang-type assemblages of southern China suggest that the overall structure and ecology of Cambrian communities remained relatively stable until at least the Middle Cambrian in subtidal siliciclastic soft substrate environments. Comparisons with modern marine benthic ecosystems further suggest the Burgess Shale community was probably highly dependent on immigration from a regional pool of species after each burial event. This could support the view that species availability, habitat characteristics, and recolonisation processes were more important in structuring the community in the long-term than species interactions or environmental variations at a local scale. [Copyright &y& Elsevier]

The morphology of Tuzoia is reinterpreted in the light of abundant new specimens from the Middle Cambrian Burgess Shale (British Columbia, Canada) and Kaili (Guizhou, China) Lagerstatten. Tuzoia was a very large (up to 180 mm long) bivalved arthropod with a nonmineralized domelike carapace strengthened by prominent pointed features and often flanked by a lateral ridge bearing a spiny frill. The reticulate pattern of Tuzoia is comparable with that of present-day crustaceans (e.g., myodocope ostracods) and is interpreted as a structural compromise between exoskeletal lightness and high resistance to mechanical stress. Tuzoia had a pair of large, stalked, spherical, possibly compound eyes facing forward. Flagella-like antennae protruded through the anterior notch. No other appendages are known except possible filamentous setae underlying the carapace. Tuzoia typically occurs as laterally (lc) or dorsoventrally (dye) compacted carapaces or single valves. Each type (lc or dvc) emphasizes particular aspects of the morphology (e.g., spiny lateral ridge, ventral margin) that were often interpreted as specific differences by previous authors. A revision of Tuzoia validates only 7 of the 23 named species. Tuzoia is placed tentatively within a group of large bivalved arthropods along with Isoxys and the possible ancestors of Thylacocephala (Lower Cambrian-Upper Cretaceous). In the Middle Cambrian, Tuzoia occurs across Laurentia, South and North China, and the Perigondwanan area (Bohemia) within a relatively narrow subtropical belt, indicating a high dispersal capability and possible latitudinal control on its distribution. Functional morphology, taphonomy, and the distributional pattern indicate that Tuzoki was a free-swimming arthropod. [ABSTRACT FROM AUTHOR]

Halkieriids and wiwaxiids are cosmopolitan sclerite-bearing metazoans from the Lower and Middle Cambrian. Although they have similar scleritomes, their phylogenetic position is contested. A new scleritomous fossil from the Burgess Shale has the prominent anterior shell of the halkieriids but also bears wiwaxiid-like sclerites. This new fossil defines the monophyletic halwaxiids and indicates that they have a key place in early lophotrochozoan history. [ABSTRACT FROM AUTHOR]

A new genus and species of a Middle Cambrian stem group brachiopod, Acanthotretella spinosa n. gen. and n. sp., is described from the Burgess Shale Formation. Most of the 42 specimens studied came from the Greater Phyllopod bed (Walcott Quarry) and were collected from five bed assemblages, each representing a single obrution event. Specimens are probably preserved within their original habitat. In contrast to all brachiopods known from the Burgess Shale, the shells of the new stem group brachiopod are often deformed and do not show signs of brittle breakage, which suggests that the valves were originally either entirely organic in composition or, more likely, had just a minor mineral component. Acanthotretella spinosa differs from all the other described Cambrian brachiopods in that it is covered by long, slender and possibly partly mineralized spines that are posteriorly inclined at an oblique angle away from the anterior margin. The spines penetrate the shell and are mainly comparable with the thorn-like organic objects that have been inferred from early siphonotretoid brachiopods. The pedicle was slender and was composed of a central coelomic region and emerged from an apical foramen at the end of an internal pedicle tube. The finding of a pedicle attached to the macrobenthic algae Dictyophycus and other epibenthos implies that A. spinosa did not have an infaunal mode of life. The visceral region and interior characters are poorly preserved. [ABSTRACT FROM AUTHOR]

Odontogriphus omalus was originally described as a problematic non-biomineralized lophophorate organism. Here we re-interpret Odontogriphus based on 189 new specimens including numerous exceptionally well preserved individuals from the Burgess Shale collections of the Royal Ontario Museum. This additional material provides compelling evidence that the feeding apparatus in Odontogriphus is a radula of molluscan architecture comprising two primary bipartite tooth rows attached to a radular membrane and showing replacement by posterior addition. Further characters supporting molluscan affinity include a broad foot bordered by numerous ctenidia located in a mantle groove and a stiffened cuticular dorsum. Odontogriphus has a radula similar to Wiwaxia corrugata but lacks a scleritome. We interpret these animals to be members of an early stem-group mollusc lineage that probably originated in the Neoproterozoic Ediacaran Period, providing support for the retention of a biomat-based grazing community from the late Precambrian Period until at least the Middle Cambrian. [ABSTRACT FROM AUTHOR]

The discovery of a new naraoiid nektaspid in the Upper Silurian (Pridolian) of southeastern Ontario significantly extends the range of this unusual group. Nektaspids are nonmineralized arthropods typical of Early and Middle Cambrian soft-bottom communities, but were thought to have become extinct in the Late Ordovician. The unique holotype specimen of Naraoia bertiensis n. sp. comes from a Konservat-Lagerstätte deposit renowned for its eurypterid fauna (the Williamsville Member of the Bertie Formation). Naraoia bertiensis lacks thoracic segments and is morphologically similar to Naraoia compacta from the Middle Cambrian Burgess Shale, save for the presence of a long ventral cephalic doublure and a subtly pointed posterior shield. To examine the phylogenetic relationships of the new naraoiid, we coded characters of the holotype specimen and of nine previously described nektaspids. The results confirm a sister taxon relationship between Naraoia compacta and Naraoia bertiensis and the monophyly of nektaspid forms lacking thoracic segments (family Naraoiidae). This latter group may have arisen from an ancestral segment-bearing form through heterochronic loss of thoracic segments early in the Cambrian. The disjunct occurrence of a naraoiid nektaspid in the Late Silurian resembles the reappearance of other "Lazarus taxa" that were thought to have been eliminated during mass extinction events. The naraoiid lineage survived the Late Ordovician biotic crisis, but in this case the "Lazarus effect" seems likely to be taphonomic in origin. [ABSTRACT FROM AUTHOR]

Hemichordate relationships remain contentious due to conflicting molecular results [ 1–7 ] and the high degree of morphological disparity between the two hemichordate classes, Enteropneusta and Pterobranchia [ 8–11 ]. Additionally, hemichordates have a poor fossil record outside of the Cambrian, with the exception of the collagenous tubes of the pterobranchs (which include graptolites). By the middle Cambrian, tube-dwelling colonial pterobranchs [ 12 , 13 ] and tube-dwelling enteropneusts coexisted [ 14 , 15 ], supporting the origin of the hemichordate body plan earlier in the Cambrian without clarifying the morphology of their last common ancestor. Here, we describe a new hemichordate, Gyaltsenglossus senis , based on 33 specimens from the 506-million-year-old Burgess Shale (Odaray Mountain, British Columbia). G. senis has a unique combination of soft anatomical characters found in both extant classes of hemichordates, namely a trimeric-vermiform body plan with an elongate proboscis and six feeding arms with tentacles. The trunk possesses a long through-gut and terminates with a bulbous structure potentially used for locomotion and/or as a temporary anchor. There is no evidence of a secreted tube. Our phylogenetic analyses retrieve this new taxon as a stem-group hemichordate, supporting the hypothesis that a vermiform body plan preceded both tube building and colonial ecologies. This new taxon suggests that a bimodal feeding ecology using tentacles to filter feed and a proboscis to deposit feed may be plesiomorphic in hemichordates. Finally, the presence of a muscular, post-anal attachment structure in all known Cambrian hemichordates supports this feature as an additional hemichordate plesiomorphy critical for understanding early hemichordate evolution. • A newly described Cambrian hemichordate preserves soft-tissue details • The new species possesses both a long proboscis and feeding arms • A round posterior structure may be homologous with the juvenile acorn worm tail • Phylogenetic analyses suggest the new species is the first stem-hemichordate known Soft-bodied hemichordate fossils are rare, making it difficult to understand early hemichordate evolution. Nanglu et al. describe a new species of exceptionally well-preserved fossil hemichordate from the 506-million-year-old Burgess Shale. This new species suggests that a vermiform body plan and bi-modal feeding were ancestral to Hemichordata. [ABSTRACT FROM AUTHOR]

A response to a letter to the editor about the article concerning Odontogriphus and relatives from the Middle Cambrian Burgess Shale that is published in previous issue is presented.

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. [ABSTRACT FROM AUTHOR]

Phylogenomic studies have greatly improved our understanding of the animal tree of life but the relationships of many clades remain ambiguous. Here we show that the rare soft-bodied animal Amiskwia from the Cambrian of Canada and China, which has variously been considered a chaetognath, a nemertine, allied to molluscs, or a problematica, is related to gnathiferans. New specimens from the Burgess Shale (British Columbia, Canada) preserve a complex pharyngeal jaw apparatus composed of a pair of elements with teeth most similar to gnathostomulids. Amiskwia demonstrates that primitive spiralians were large and unsegmented, had a coelom, and were probably active nekto-benthic scavengers or predators. Secondary simplification and miniaturisation events likely occurred in response to shifting ecologies and adaptations to specialised planktonic habitats. Jean-Bernard Caron and Brittany Cheung provide new evidence that the rare Cambrian animal Amiskwia is closely related to gnathiferans, thereby resolving its ambiguous phylogeny. They describe new specimens from the Burgess Shale that preserve the complex jaw apparatus, which is most similar to gnathostomulids. [ABSTRACT FROM AUTHOR]

The original article [1] had 4 paragraphs which contained erroneous information. In this correction article the correct and incorrect information is shown. [ABSTRACT FROM AUTHOR]

Summary Chaetognaths (arrow worms) are a separate phylum (Chaetognatha) of small carnivorous animals, dominantly pelagic, and a major component of today’s plankton [ 1, 2 ]. The position of Chaetognatha among metazoan phyla remains equivocal—neither morphological nor molecular data provide definitive evidence [ 3 ]. Originating early in the Cambrian period [ 4 ], if not earlier [ 5 ], chaetognaths quickly became important members of marine metazoan communities [ 6 ]. Chaetognath grasping spines, originally reported as conodonts, occur worldwide in many Cambrian marine sediments [ 6, 7 ]. Fossilized chaetognath bodies, in contrast, are very rare: only two unequivocal specimens have been reported, both from the early Cambrian of China [ 8, 9 ]. Here we describe Capinatator praetermissus , a new genus and species, based on ∼50 specimens from several middle Cambrian Burgess Shale localities in British Columbia, many of which preserve evidence of soft tissues. Capinatator praetermissus reached body lengths of nearly 10 cm exclusive of fins, a much larger size than that of most living forms. Clusters of specimens preserving the body indicate that they were rapidly buried, providing indirect evidence that they swam near the seabed. The feeding apparatus comprises up to ∼25 spines in each half, almost double the maximum number in living chaetognaths. Early chaetognaths apparently occupied ecological niches associated with predatory euarthropods. The large body size and high number of grasping spines in C. praetermissus may indicate that miniaturization and migration to a planktonic lifestyle were secondary. [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]

The article discusses the implications of the palaeontology of the machaeridian, ancient worms in scaly armour of mineralized shell plates around a soft body. A study by J. Vinther and colleagues is based on the machaeridian scleritome, a specimen that has withstood the odds of fossil preservation by retaining traces of the elusive soft body. Researchers interpret the protrusions bearing prominent filaments in bundles along both sides of the animal to be parapodia. It is said that identifying machaeridians as primitive polychaetes eradicates problems arising from their assignment to other groups of animals.

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

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian 'explosion'. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution -- but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils -- including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum -- as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the 'middle' Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution. [ABSTRACT FROM AUTHOR]

Ecological interactions, including symbiotic associations such as mutualism, parasitism and commensalism are crucial factors in generating evolutionary novelties and strategies. Direct examples of species interactions in the fossil record generally involve organisms attached to sessile organisms in an epibiont or macroboring relationship. Here we provide support for an intimate ecological association between a calcareous brachiopod (Nisusia) and the stem group mollusc Wiwaxia from the Burgess Shale. Brachiopod specimens are fixed to Wiwaxia scleritomes, the latter showing no signs of decay and disarticulation, suggesting a live association. We interpret this association as the oldest unambiguous example of a facultative ectosymbiosis between a sessile organism and a mobile benthic animal in the fossil record. The potential evolutionary advantage of this association is discussed, brachiopods benefiting from ease of attachment, increased food supply, avoidance of turbid benthic conditions, biofoul and possible protection from predators, suggesting commensalism (benefiting the symbiont with no impact for the host). While Cambrian brachiopods are relatively common epibionts, in particular on sponges, the association of Nisusia with the motile Wiwaxia is rare for a brachiopod species, fossil or living, and suggests that symbiotic associations were already well established and diversified by the ''middle'' (Series 3, Stage 5) Cambrian. [ABSTRACT FROM AUTHOR]

In Cambrian fossil Lagerstätten like the Burgess Shale, exceptionally preserved arthropods constitute a large part of the taxonomic diversity, providing opportunities to study the early evolution of this phylum in detail. The anomalocaridids, large presumed pelagic predators, are particularly relevant owing to their unique combination of morphological characters and basal position in the arthropod stem lineage. Although isolated elements and fragmented specimens were first discovered over 100 years ago, subsequent findings of more complete bodies ofAnomalocarisandPeytoia, especially in the 1980s, allowed for a better understanding of these enigmatic forms. Their evolutionary significance as stem group arthropods was further clarified by the recent discovery of a third anomalocaridid taxon,Hurdia. Here, examination of hundreds ofHurdiaspecimens from different stratigraphical layers within the Burgess Shale and Stephen Formation, combined with statistical analyses, provides a detailed description of the taphonomy, morphology and diversity of the genus and further elucidates anomalocaridid systematics.Hurdiais distinguished from other anomalocaridids in having mouthparts with extra rows of teeth, a large frontal carapace complex and diminutive swimming flaps with prominent setal structures. The two original species,H. victoriaWalcott, 1912 andH. triangulataWalcott, 1912, are confirmed based on morphometric outline analyses of the frontal carapace components combined with stratigraphical evidence; a third species,Hurdia dentataSimonetta & Delle Cave, 1975, is synonymized withH. victoria. Morphology, preservation and stratigraphical distribution suggest thatH. victoriaandH. triangulatashare the same type of frontal appendage; a second type of appendage, previously assigned toHurdia(Morph A), belongs toPeytoia nathorsti. These and other morphological differences between the anomalocaridids may reflect different feeding strategies. Appendages and mouthparts ofHurdiaindet. sp. are also identified from the Spence Shale Member of Utah, makingHurdiaandAnomalocaristhe most common and globally distributed anomalocaridid taxa. [ABSTRACT FROM PUBLISHER]

Pettersson Stolk, S., Holmer, L. E. and Caron, J -B. 2010. First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale. — Acta Zoologica (Stockholm) 91: 150–162 The organophosphatic shells of linguloid brachiopods are a common component of normal Cambrian–Ordovician shelly assemblages. Preservation of linguloid soft-part anatomy, however, is extremely rare, and restricted to a few species in Lower Cambrian Konservat Lagerstätten. Such remarkable occurrences provide unique insights into the biology and ecology of early linguloids that are not available from the study of shells alone. Based on its shells, Lingulella waptaensis Walcott, was originally described in 1924 from the Middle Cambrian Burgess Shale but despite the widespread occurrence of soft-part preservation associated with fossils from the same levels, no preserved soft parts have been reported. Lingulella waptaensis is restudied herein based on 396 specimens collected by Royal Ontario Museum field parties from the Greater Phyllopod Bed (Walcott Quarry Shale Member, British Columbia). The new specimens, including three with exceptional preservation of the pedicle, were collected in situ in discrete obrution beds. Census counts show that L. waptaensis is rare but recurrent in the Greater Phyllopod Bed, suggesting that this species might have been generalist. The wrinkled pedicle protruded posteriorly between the valves, was composed of a central coelomic space, and was slender and flexible enough to be tightly folded, suggesting a thin chitinous cuticle and underlying muscular layers. The nearly circular shell and the long, slender and highly flexible pedicle suggest that L. waptaensis lived epifaunally, probably attached to the substrate. Vertical cross-sections of the shells show that L. waptaensis possessed a virgose secondary layer, which has previously only been known from Devonian to Recent members of the Family Lingulidae. [ABSTRACT FROM AUTHOR]

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 Hardía, based on several hundred specimens from the Burgess Shale in Canada. Hardío 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. [ABSTRACT FROM AUTHOR]

Exceptionally preserved fossils are the product of complex interplays of biological and geological processes including burial, autolysis and microbial decay, authigenic mineralization, diagenesis, metamorphism, and finally weathering and exhumation. Determining which tissues are preserved and how biases affect their preservation pathways is important for interpreting fossils in phylogenetic, ecological, and evolutionary frameworks. Although laboratory decay experiments reveal important aspects of fossilization, applying the results directly to the interpretation of exceptionally preserved fossils may overlook the impact of other key processes that remove or preserve morphological information. Investigations of fossils preserving non-biomineralized tissues suggest that certain structures that are decay resistant (e.g., the notochord) are rarely preserved (even where carbonaceous components survive), and decay-prone structures (e.g., nervous systems) can fossilize, albeit rarely. As we review here, decay resistance is an imperfect indicator of fossilization potential, and a suite of biological and geological processes account for the features preserved in exceptional fossils. [ABSTRACT FROM AUTHOR]

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian ( Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian ( Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral-like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube-like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian ( Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef-building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well-developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle. [ABSTRACT FROM AUTHOR]