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

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

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

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

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

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

Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock being extremely abundant, and the larger Redlichia takooensis Lu , being common. Many specimens within the EBS represent complete moulted exoskeletons, which is unusual for Cambrian fossil deposits. The abundance of complete moults provides an excellent record that has allowed the recognition of various recurrent moult configurations for both species, enabling the inference of movement sequences required to produce such arrangements. Moult configurations of E. bilobata are characterized by slight displacement of the joined rostral plate and librigenae, often accompanied by detachment of the cranidium, suggesting ecdysis was achieved by anterior withdrawal via opening of the cephalic sutures. Moulting in R. takooensis often followed the same method, but configurations show greater displacement of cephalic sclerites, suggesting more vigorous movement by the animal during moulting. Both species also show rare examples of Salter's configuration, with the entire cephalon anteriorly inverted, and several other unusual configurations. These results indicate that moulting in trilobites was a more variable process than originally thought. In contrast, other Cambrian Konservat‐Lagerstätten with an abundance of trilobites (e.g. Wheeler Shale, USA, and Mount Stephen Trilobite Beds, Canada) show larger numbers of ‘axial shields’ and isolated sclerites, often interpreted as disarticulated exuviae. This points to a higher level of disturbance from factors, such as animal activity, depositional processes or water movement, compared to that of the EBS, where quiescent conditions and intermittent seafloor anoxia contributed to an unparalleled trilobite moulting record. [ABSTRACT FROM AUTHOR]

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

Banffia constricta is an enigmatic Burgess Shale animal originally described by Charles Walcott in 1911 as an annelid, and more recently as a stem-group deuterostome. Interpreted, on the basis of anatomy, to have been bottom-feeders, there are few other data from which to draw interpretations of Banffia's life habit. A slab of Burgess Shale with a dense aggregation of B. constricta may indicate a gregarious habit for the animal , as taphonomic and stratigraphical data indicate an in situ origin for the assemblage. Clustering of individuals, high density of the individuals and non-random within-cluster orientation support the hypothesis that detritus-feeding B. constricta congregated to feed on a local, rich food source. Presumed opportunistic feeding aggregations have been documented in at least one other Burgess Shale taxon and have been described for other fossil benthic marine invertebrates. Extant benthic marine invertebrates such as holothurians and echinoids exhibit mass feeding behaviour and may serve as modern analogs for the behaviour represented by the B. constricta assemblage. [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: 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]

A comprehensive restudy of all material assigned in previous reports to the noncalcified macroalgal genus Bosworthia Walcott from the Burgess Shale Formation (Cambrian Stage 5) of western Canada and the slightly older Kaili Formation of South China, together with investigation of previously unpublished material, indicates important morphological differences between the two species heretofore included within the genus. On the basis of these differences, B. gyges is removed from the genus and assigned to Walcottophycus gen. nov. Most specimens previously assigned to Bosworthia from the Cambrian of China, including those assigned to B. simulans Walcott, are transferred to W. gyges , and it is concluded that Bosworthia remains unknown in China. Morphological details for this material when evaluated in the context of molecular clock studies and other fossil macroalgae indicate that B. simulans may belong to the extant red algal subclass Rhodymeniophycidae (Florideophyceae) and indicate that W. gyges may be an early form of bryopsidalean alga within the Chlorophyta. [ABSTRACT FROM AUTHOR]

The Middle Cambrian (series 3, Drumian, Bolaspidella Biozone) Ravens Throat River Lagerstätte in the Rockslide Formation of the Mackenzie Mountains, northwestern Canada, contains a Burgess Shale-type biota of similar age to the Wheeler and Marjum formations of Utah. The Rockslide Formation is a unit of deep-water, mixed carbonate and siliciclastic facies deposited in a slope setting on the present-day northwestern margin of Laurentia. At the fossil-bearing locality, the unit is about 175 m thick and the lower part onlaps a fault scarp cutting lower Cambrian sandstones. It consists of a succession of shale, laminated to thin-bedded lime mudstone, debris-flow breccias, minor calcareous sandstone, greenish-coloured calcareous mudstone and dolomitic siltstone, overlain by shallow-water dolostones of the Broken Skull Formation, which indicates an overall progradational sequence. Two ~1-m-thick units of greenish calcareous mudstone in the upper part exhibit soft-bodied preservation, yielding a biota dominated by bivalved arthropods and macrophytic algae, along with hyoliths and trilobites. It represents a low-diversity in situ community. Most of the fossils occur in the lower unit, and only the more robust components are preserved. Branching burrows are present under the carapaces of some arthropods, and common millimetre-sized disruptions of laminae are interpreted as bioturbation. The fossiliferous planar-laminated calcareous mudstone consists of chlorite, illite, quartz silt, calcite and dolomite and is an anomalous facies in the succession. It was deposited via hemipelagic fallout of a mixture of platform-derived and terrestrial mud. Geochemical analysis and trace-element proxies indicate oxic bottom waters that only occasionally might have become dysoxic. Productivity in the water column was dominated by cyanobacteria. Fragments of microbial mats are common as carbonaceous seams. Complete decay of soft tissues was interrupted due to the specific sediment composition, providing support for the role of clay minerals, possibly chlorite, in the taphonomic process. [ABSTRACT FROM AUTHOR]

The feeding ecology of the 505-million-year-old arthropod Sidneyia inexpectans from the middle Cambrian (Series 3, Stage 5) Burgess Shale fauna (British Columbia, Canada) is revealed by three lines of evidence: the structure of its digestive system, the fossilized contents of its gut and the functional anatomy of its appendages. The digestive tract of Sidneyia is straight, tubular and relatively narrow in the trunk region. It is enlarged into a pear-shaped area in the cephalic region and stretches notably to form a large pocket in the abdomen. The mouth is ventral, posteriorly directed and leads to the midgut via a short tubular structure interpreted as the oesophagus. Anteriorly, three pairs of glands with internal, branching tubular structures open into the digestive tract. These glands have equivalents in various Cambrian arthropod taxa (e.g. naraoiids) and modern arthropods. Their primary function was most likely to digest and assimilate food. The abdominal pocket of Sidneyia concentrates undigested skeletal elements and various residues. It is interpreted here as the functional analogue of the stercoral pocket of some extant terrestrial arachnids (e.g. Araneae, Solifugae), whose primary function is to store food residuals and excretory material until defecation. Analysis of the gut contents indicates that Sidneyia fed largely on small ptychopariid trilobites, brachiopods, possibly agnostids, worms and other undetermined animals. Sidneyia was primarily a durophagous carnivore with predatory and/or scavenging habits, feeding on small invertebrates that lived at the water-sediment interface. There is no evidence for selective feeding. Its food items (e.g. living prey or dead material) were grasped and manipulated ventrally by its anterior appendages, then macerated into ingestible fragments and conveyed to the mouth via the converging action of strong molar-like gnathobases. Digestion probably took place within the anterior midgut via enzymes secreted in the glands. Residues were transported through the digestive tract into the abdominal pocket. The storage of faeces suggests infrequent feeding. The early diagenetic three-dimensional preservation of the digestive glands and abdominal pocket may be due to the capacity of Sidneyia to store Phosphorus and Calcium (e.g. spherites) in its digestive tissues during life as do, for example, modern horseshoe crabs. [ABSTRACT FROM AUTHOR]

The controversy about a Cambrian "explosion" of morphological disparity (followed by decimation), cladogenesis and fossilization is of central importance for the history of life. This paper revisits the controversy (with emphasis in onychophorans, which include emblematic organisms such as Hallucigenia), presents new data about the Chengjiang (Cambrian of China) faunal community and compares it and the Burgess Shale (Cambrian of Canada) with an ecologically similar but modern tropical marine site where onychophorans are absent, and with a modern neotropical terrestrial onychophoran community. Biovolume was estimated from material collected in Costa Rica and morphometric measurements were made on enlarged images of fossils. Cambrian tropical mudflats were characterized by the adaptive radiation of two contrasting groups: the vagile arthropods and the sessile poriferans. Arthropods were later replaced as the dominant benthic taxon by polychaetes. Vagility and the exoskeleton may explain the success of arthropods from the Cambrian to the modern marine and terrestrial communities, both in population and biovolume. Food ecological displacement was apparent in the B. Shale, but not in Chengjiang or the terrestrial community. When only hard parts were preserved, marine and terrestrial fossil deposits of tropical origin are even less representative than deposits produced by temperate taxa, Chengjiang being an exception. Nutrient limitations might explain why deposit feeding is less important in terrestrial onychophoran communities, where carnivory, scavenging and omnivory (associated with high motility and life over the substrate) became more important. Fossil morphometry supports the interpretation of "lobopod animals" as onychophorans, whose abundance in Chengjiang was equal to their abundance in modern communities. The extinction of marine onychophorans may reflect domination of the infaunal habitat by polychaetes. We conclude that (1) a mature ecological community structure was generalized during the Cambrian, and even biodiversity and equitability indices were surprisingly close to modern values; (2) the morphological diversity and geographic distribution of onychophorans indicate a significant pre-Cambrian evolutionary history which does not support the "explosion" hypothesis; (3) disparity among phyla was not as important as the explosion-decimation model predicts, but in the case of onychophorans, disparity within the phylum was greater than it is today and its reduction may have been associated with migration into the sediment when large predators evolved.Con base en nuevos hallazgos paleontológicos hechos en el sur de China se analiza la controversia sobre la "explosión del Cámbrico" que supuestamente incluyó un gran aumento en la diversidad morfológica, con la aparición de muchos filos que posteriormente se extinguieron. Se compara las comunidades fósiles de Cheng-yiang (China) y Burgess Shale (Canadá) con una comunidad marina similar moderna (Punta Morales, Costa Rica) y con una comunidad terrestre moderna (Coronado, Costa Rica) que al igual que los dos depósitos fosilíferos incluye al filo Onychophora. Se concluye que (1) durante el Cámbrico las comunidades costeras tropicales eran similares a las modernas al punto de que incluso los índices de biodiversidad y equitabilidad eran semejantes a los actuales; (2) la diversidad morfológica y distribución geográfica de los onicóforos indican una larga historia evolutiva precámbrica que no apoya el modelo de la "explosión"; (3) las diferencias morfológicas entre filos no eran tan importantes como predice el modelo de "explosión seguida de extinción", pero en la caso de los onicóforos, la diversidad morfológica dentro del filo era mayor en el Cámbrico que en la actualidad, y su reducción puede haber estado asociada con la migración al interior del sedimento al evolucionar mayores depredadores.

The Guole biota contains common shelly fossils and some exceptionally well-preserved soft-bodied fossils. Stratigraphically, it is located in the mudstones and siltstones of the Sandu Formation near Guole Town, Jingxi County, Guangxi, South China. It is the first Furongian (late Cambrian) Burgess Shale-type biota found in the world, thereby filling the gap between middle Cambrian and Lower Ordovician Burgess Shale-type Lagerstätten. Preliminary studies suggest that the Guole biota includes approximately seven metazoan groups as well as algae. These will provide important new evolutionary and ecological information. [ABSTRACT FROM AUTHOR]

The controversy about a Cambrian "explosion" of morphological disparity (followed by decimation), cladogenesis and fossilization is of central importance for the history of life. This paper revisits the controversy (with emphasis in onychophorans, which include emblematic organisms such as Hallucigenia), presents new data about the Chengjiang (Cambrian of China) faunal community and compares it and the Burgess Shale (Cambrian of Canada) with an ecologically similar but modern tropical marine site where onychophorans are absent, and with a modern neotropical terrestrial onychophoran community. Biovolume was estimated from material collected in Costa Rica and morphometric measurements were made on enlarged images of fossils. Cambrian tropical mudflats were characterized by the adaptive radiation of two contrasting groups: the vagile arthropods and the sessile poriferans. Arthropods were later replaced as the dominant benthic taxon by polychaetes. Vagility and the exoskeleton may explain the success of arthropods from the Cambrian to the modern marine and terrestrial communities, both in population and biovolume. Food ecological displacement was apparent in the B. Shale, but not in Chengjiang or the terrestrial community. When only hard parts were preserved, marine and terrestrial fossil deposits of tropical origin are even less representative than deposits produced by temperate taxa, Chengjiang being an exception. Nutrient limitations might explain why deposit feeding is less important in terrestrial onychophoran communities, where carnivory, scavenging and omnivory (associated with high motility and life over the substrate) became more important. Fossil morphometry supports the interpretation of "lobopod animals" as onychophorans, whose abundance in Chengjiang was equal to their abundance in modern communities. The extinction of marine onychophorans may reflect domination of the infaunal habitat by polychaetes. We conclude that (1) a mature ecological community structure was generalized during the Cambrian, and even biodiversity and equitability indices were surprisingly close to modern values; (2) the morphological diversity and geographic distribution of onychophorans indicate a significant pre-Cambrian evolutionary history which does not support the "explosion" hypothesis; (3) disparity among phyla was not as important as the explosion-decimation model predicts, but in the case of onychophorans, disparity within the phylum was greater than it is today and its reduction may have been associated with migration into the sediment when large predators evolved.Con base en nuevos hallazgos paleontológicos hechos en el sur de China se analiza la controversia sobre la "explosión del Cámbrico" que supuestamente incluyó un gran aumento en la diversidad morfológica, con la aparición de muchos filos que posteriormente se extinguieron. Se compara las comunidades fósiles de Cheng-yiang (China) y Burgess Shale (Canadá) con una comunidad marina similar moderna (Punta Morales, Costa Rica) y con una comunidad terrestre moderna (Coronado, Costa Rica) que al igual que los dos depósitos fosilíferos incluye al filo Onychophora. Se concluye que (1) durante el Cámbrico las comunidades costeras tropicales eran similares a las modernas al punto de que incluso los índices de biodiversidad y equitabilidad eran semejantes a los actuales; (2) la diversidad morfológica y distribución geográfica de los onicóforos indican una larga historia evolutiva precámbrica que no apoya el modelo de la "explosión"; (3) las diferencias morfológicas entre filos no eran tan importantes como predice el modelo de "explosión seguida de extinción", pero en la caso de los onicóforos, la diversidad morfológica dentro del filo era mayor en el Cámbrico que en la actualidad, y su reducción puede haber estado asociada con la migración al interior del sedimento al evolucionar mayores depredadores.

Extant panarthropods (euarthropods, onychophorans and tardigrades) are hallmarked by stunning morphological and taxonomic diversity, but their central nervous systems (CNS) are relatively conserved. The timing of divergences of the ground pattern CNS organization of the major panarthropod clades has been poorly constrained because of a scarcity of data from their early fossil record. Although the CNS has been documented in three-dimensional detail in insects from Cenozoic ambers, it is widely assumed that these tissues are too prone to decay to withstand other styles of fossilization or geologically older preservation. However, CambrianBurgess Shale-type compressions have emerged as sources of fossilized brains and nerve cords. CNS in these Cambrian fossils are preserved as carbon films or as iron oxides/hydroxides after pyrite in association with carbon. Experiments with carcasses compacted in fine-grained sediment depict preservation of neural tissue for a more prolonged temporal window than anticipated by decay experiments in other media. CNS and compound eye characters in exceptionally preserved Cambrian fossils predict divergences of the mandibulate and chelicerate ground patterns by Cambrian Stage 3 (ca 518 Ma), a dating that is compatible with molecular estimates for these splits. [ABSTRACT FROM AUTHOR]

Palaeoscolecid worms are a ubiquitous group of Early Palaeozoic ecdysozoans that are curiously lacking in the archetypal Cambrian Lagerstätten, the Burgess Shale. Here I describe Scathascolex minor gen. et sp. nov, the first unequivocal palaeoscolecid from this site. Scathascolex is armoured with simple Hadimopanella-like plates, but lacks smaller platelets, pointing to a close affinity with the Palaeoscolecida sensu stricto. Neither preservational nor environmental factors account for the scarcity of palaeoscolecids in the Burgess Shale, which presumably represents an ecological phenomenon. [ABSTRACT FROM AUTHOR]

The morphology of the arthropod Sanctacaris uncata, from the Middle Cambrian Burgess Shale of Canada, is reinterpreted based on a restudy of previously described material. Although originally considered a chelicerate-like arthropod, these affinities were dismissed based primarily on interpretations of the anterior appendages and hypotheses which considered the megacheirans ('great-appendage' arthropods) as putative ancestors of chelicerates. The similarities between megacheirans and chelicerates appear to be overstated however, and this study instead reaffirms the identity of putative chelicerate feature in S. uncata and similar arthropods such as Sidneyia and Emeraldella, both also from the Middle Cambrian Burgess Shale. Newly interpreted features, including the presence of pediform exites, multi-partite trunk exopods, and a trunk differentiated into an anterior limb-bearing area and a differentiated posterior limbless abdomen, were coded into an extensive phylogenetic data set of fossil and recent arthropods. In all analyses, Sanctacaris resolved as the basal-most member of total-group Euchelicerata (the least inclusive group including horseshoe crabs and arachnids but not pycnogonids), thus making it the oldest chelicerate in the fossil record. The vicissicaudates (including Sidneyia, Emeraldella, aglaspidids, and cheloniellids-all of which have previously been allied to chelicerates) resolved as sister-taxon to crown-group Chelicerata. This topology indicates that many purported chelicerate features, such as lamellar gills, and a differentiated posterior abdomen evolved sequentially in the chelicerate stem-lineage. [ABSTRACT FROM AUTHOR]

A new Chengjiang-type fossil assemblage is reported herein from the lower part of the Hongjingshao Formation at Xiazhuang village of Chenggong, Kunming, Yunnan. The fossil assemblage, named as Xiazhuang fossil assemblage, yields predominantly soft-bodied fossils, including arthropods, brachiopods, priapulids, lobopods and some problematic taxa, with arthropods being the most dominant group. Preservation and composition of the fossil assemblage are very similar to the typical Chengjiang biota, which is preserved in the middle Yu'anshan Formation in the large area of eastern Yunnan. The associated trilobites demonstrate that the soft-bodied fossil assemblage belongs to the late Qiongzhusian in age (Stage 3, Cambrian), suggesting that the Hongjingshao Formation is probably a diachronous lithostratigraphic unit ranging from the upper Qiongzhusian to the lower Canglangpuan stages in eastern Yunnan. The fossil assemblage from the Xiazhuang area fills up the missing link between the typical older Chengjiang biota and the younger Malong and Guanshan biotas, making eastern Yunnan a unique area in the world to reveal the early evolutionary history of animals and palaeocommunity dynamics during the 'Cambrian explosion'. [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]

Abstract: Sidneyia inexpectansWalcott, 1911 from the Cambrian Series 3 Burgess Shale of British Columbia is largely accepted as a representative of the artiopodans, an assemblage of Paleozoic arthropod taxa, including trilobites and their immediate relatives. Its appendage morphology was never fully understood, but the exopod seemed to differ from that of other artiopodans, except for the shared presence of lamellae. The head was considered to comprise only the ocular and antennular segments, these being covered entirely on the ventral side by a large doublure. This short head was often taken as an evidence for variability of head segment counts in Cambrian arthropods, and to falsify the hypothesis of a head with three postantennular segments in the euarthropod ground pattern. Restudy of a substantial amount of material of S. inexpectans shows that previous interpretations of a short head were based on taphonomically deformed specimens, where the head was either partly folded, or entirely flipped under the thorax, resulting in the dorsal shield being mistaken for an extensive doublure. Rather than an extensive doublure, there is a broad hypostome, and the head comprises ocular, antennular, and at least two postantennular appendage bearing segments. The appendage morphology is shown to be consistent with artiopodan affinities. The exopod is of the bilobate flap-like type with lamellae inserting on the proximal portion, earlier proposed as a potential autapomorphy of Artiopoda. Reinforcement of artiopodan affinities for S. inexpectans and reinterpretation of its head reconciles this species with current understanding of arthropod phylogeny and head segmentation. [Copyright &y& Elsevier]

Extant arthropods are diverse and ubiquitous, forming a major constituent of most modern ecosystems. Evidence from early Palaeozoic Konservat Lagerstätten indicates that this has been the case since the Cambrian. Despite this, the details of arthropod origins remain obscure, although most hypotheses regard the first arthropods as benthic predators or scavengers such as the fuxianhuiids or megacheirans ('great-appendage' arthropods). Here, we describe a new arthropod from the Tulip Beds locality of the Burgess Shale Formation (Cambrian, series 3, stage 5) that possesses a weakly sclerotized thorax with filamentous appendages, encased in a bivalved carapace, and a strongly sclerotized, elongate abdomen and telson. A cladistic analysis resolved this taxon as the basal-most member of a paraphyletic grade of nekto-benthic forms with bivalved carapaces. This grade occurs at the base of Arthropoda (panarthropods with arthropodized trunk limbs) and suggests that arthrodization (sclerotization and jointing of the exoskeleton) evolved to facilitate swimming. Predatory and fully benthic habits evolved later in the euarthropod stem-lineage and are plesiomorphically retained in pycnogonids (sea spiders) and euchelicerates (horseshoe crabs and arachnids). [ABSTRACT FROM AUTHOR]

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

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates. [ABSTRACT FROM AUTHOR]

Sarotrocercus oblitus is a small arthropod from the Cambrian Burgess Shale. It was originally described with a short head with only two appendage-bearing segments (the first appendage being limb-shaped), a short trunk of nine segments and lamellate trunk limbs. This rather "unusual" morphology inspired various authors to propose evolutionary scenarios concerning segmentation and appendages. The head of S. oblitus served also for scenarios about the evolution of the arthropod head, because it seemed to document the evolutionary step between the level of Arthropoda sensu stricto (head with one appendage-bearing segment) and that of Euarthropoda (head comprising four appendage-bearing segments). Here we report that the morphology of S. oblitus differs in several significant aspects from its original description, e.g., in the composition of the head, number of trunk segments, and appendage morphology. In consequence, many earlier assumptions based on the original description must be rejected. Although the material consists of only seven individuals, ontogenetic variation of the number of trunk segments was observed, pointing to, at least, two developmental stages. Therefore, S. oblitus is morphologically less different from other Cambrian arthropods than previously thought, but possesses a head with three appendage-bearing segments and lacks a prominent antenn(ul)a. These characters point to a position of S. oblitus inside Arthropoda s. str., deriving from the lineage towards Euarthropoda. The morphology also indicates a special life style, e.g., by the presence of large, stalked eyes, apparently in convergence to one of the Cambrian "Orsten" crustacean stem-lineage derivatives, Henningsmoenicaris scutula. [ABSTRACT FROM AUTHOR]

The complex history of description of the anomalocaridids has partly been caused by the fragmentary nature of these fossils. Frontal appendages and mouth parts are more readily preserved than whole-body assemblages, so the earliest work on these animals examined these structures in isolation. After several decades of research, these disarticulated elements were assembled together to reconstruct the anomalocaridid body plan, and a total of three Burgess Shale genera, Anomalocaris, Laggania and Hurdia, were described in full. Here we present new frontal appendage material of additional anomalocaridid taxa from the ‘Middle’ Cambrian (Series 3) Burgess Shale Formation in Canada, showing that the diversity of anomalocaridids in this locality is even higher than previously thought. Material includes Amplectobelua stephenensis sp. nov. , the first known occurrence of this genus outside of China; Caryosyntrips serratus gen. et sp. nov., which is similar to the Anomalocaris appendage but has a straighter outline and a different arrangement of spines; and an appendage that may be either the Laggania appendage or a third morph of the Hurdia appendage. The new anomalocaridid material is contemporaneous with the previously described taxa Anomalocaris, Laggania, and Hurdia, and the differences in morphology between the frontal appendages may reflect different feeding strategies. The stratigraphically lowest locality, S7 on Mount Stephen, yields material from all anomalocaridid taxa, but the assemblages in the younger quarries on Fossil Ridge are dominated by Anomalocaris and Hurdia only. [ABSTRACT FROM AUTHOR]

Abstract: This paper compares two North American occurrences of fossils that exhibit Burgess Shale-type preservation (BST): the Middle Cambrian Burgess Shale Formation of southeastern British Columbia, and the Lower to Middle Cambrian Kinzers Fm of southeastern Pennsylvania. Redox-sensitive trace elements (Mo, Ni/Co, V/Sc) indicate that palaeo-oxygenation conditions varied with respect to faunal assemblage and location. Ogygopsis-bearing fauna in both the Burgess Shale and Kinzers formations are associated with a dysoxic metal signature. Burgess Shale-type preservation in the Kinzers Fm was deposited in a fluctuating oxycline, whereas the BST-fossil beds of Fossil Ridge and Mt Stephen (Burgess Shale Fm) yield a geochemical signature that consistently indicates oxic conditions. Thus, sustained low-oxygen conditions do not correlate with higher quality of fossil preservation at these sites. Ogygopsis is associated with dysoxic conditions in both localities, and may therefore be a faunal indicator of such environments in Middle Cambrian strata. Carbonate units associated with both the Burgess Shale and Kinzers formations contain large, generally concordant, near-surface cavities that contain herringbone calcite cements that show no significant enrichment in the potential calcite inhibitor ions such as reduced Fe, Sr, S, Pb, and Zn. The abundance of micro-inclusions of dolomite suggests that Mg may have been the calcite inhibitor ion responsible for herringbone cement in these Cambrian occurrences. Another distinctive mineralogical feature is the presence of doubly-terminated, euhedral, zoned, authigenic quartz as a conspicuous part of a carbonate host-rock replacement assemblage. The association of both herringbone calcite and zoned quartz crystals in units associated with the Burgess Shale and Kinzers formations suggest a common feature in the depositional and diagenetic environment of these two regions in which Burgess Shale-type fossil preservation occur; syn-depositional brine flow is the likely cause of these distinct mineralogical features. Brine seeps in the depositional environment may have provided nutrients for stable high-abundance faunal communities in a slope setting, thereby accounting for the abundance of fossil material. Furthermore, high-salinity brine in the pore waters of carcass-bearing Cambrian muds may have further limited shallow burrowing, scavenging, and decomposition rates, accounting for the exceptional preservation in these two localities. [Copyright &y& Elsevier]

Abstract: Until recently, research on the renowned fossil animals of the Burgess Shale has advanced to a greater degree than an understanding of the rocks in which they are found. Studies addressing lithostratigraphy and hydrothermal petrography of the so-called ‘inner carbonate belt’ and adjacent Chancellor Group, however, have begun to re-evaluate long-standing hypotheses on the Middle Cambrian of Western Canada. Tectonic activity along the Kicking Horse Rim (buried remnants of Neoproterozoic rifting) during the Cambrian Period had more influence on local sedimentation than previously thought. Notably, large-scale collapse of the Cathedral carbonate platform margin at ∼509 Ma BP is evidence of reactivated basement faults. These failures produced listric Megatruncation Surfaces, having near vertical escarpments (>150 m in height) where they terminate against the platform. The majority of what have been interpreted as shed olistoliths from the Cathedral platform margin is herein shown instead to be carbonate mud mounds. These grew in situ along the face of the Cathedral Escarpment, and are associated with fossiliferous intervals in overlying basinal mudstones at three distinct stratigraphic horizons (two within the Burgess Shale) during the Delamaran and Marjuman stages. The mounds nucleated where deep-seated normal faults intersected the seafloor, building primarily during periods of relative sea level transgression. Mound growth is associated with syndepositional exhalative activity, inferred from stratigraphic relations, sedimentology, and geochemistry. The new name Monarch Formation is proposed for the initial post-collapse beds and mounds deposited against the Cathedral Escarpment (early Glossopleura Zone); the variably-defined term ‘Takakkaw Tongue’ is thereby confined to pre-collapse slope deposits coeval to and correlative with the Cathedral Formation. [Copyright &y& Elsevier]

Abstract: Elemental mapping offers rapid qualitative analysis of the mineralogy of exceptionally preserved fossils and, importantly, can be undertaken on unprepared samples. The results of such analyses are sensitive to several variables including the operating conditions (particularly accelerating voltage) and the composition of the material analysed: the higher the accelerating voltage employed and the lower the atomic number of the material impacted by the electron beam the larger the excitation volume (in effect a greater thickness of the specimen) that contributes to the resultant elemental map. Fossil non-biomineralised arthropods are commonly preserved as ‘carbonaceous compresssions’: the cuticle is organically preserved and, in hand specimen, essentially two dimensional. The external and internal surfaces of this cuticle may be coated by authigenic minerals. Examples include arthropods from the Middle Cambrian Burgess Shale exceptional fauna. It is theoretically possible that the proportion of the total counts generated by a thin surficial film of low atomic number (e.g., a carbonaceous cuticle) will be insufficient to distinguish it as a peak above the background spectrum; i.e., it will be effectively transparent in elemental maps. Simulated energy dispersive X-ray spectra for a windowless ED detector suggest that at an accelerating voltage of 15 keV a carbon layer less than about 0.2 μm thick will be undetectable, and at 5 keV a layer less than a few hundred angstroms thick. These values will increase if the energy dispersive spectrometer is fitted with a Super Ultra Thin Window. [Copyright &y& Elsevier]

Continuing debate over the evolution and morphology of the arthropod head has led to considerable interest in the relevance of the evidence from the fossil record. However, dispute over homology and even presence of appendages and sclerites in Cambrian arthropods has resulted in widely differing views of their significance. The head structures of several important taxa, Fuxianhuia, Canadaspis, Odaraia, Chengjiangocaris and Branchiocaris are redescribed, revealing the essential similarity between them. In particular, all possessed an anterior sclerite, probably followed by a large posterior, ventral sclerite that is likely to be homologous to the hypostome of trilobites. The presence of a similar feature in Sanctacaris is also possible, but less well-supported. An anterior sclerite, usually bearing eyes, as in Fuxianhuia, appears to be a widespread feature of basal arthropods. Whether or not this sclerite represents an original articulating protocerebral segment on its own is, however, open to debate. [ABSTRACT FROM AUTHOR]

The collection, since 1975, of over 1500 specimens of Leanchoilia Walcott by the Royal Ontario Museum has prompted reassessment of the genus and its species from the Middle Cambrian Burgess Shale. Among new characters in Leanchoilia superlata, the type species, are the presence of two pairs of eyes, a dorsal double carina bracketing the axis of the body segments, segmentation of the gill branch of the appendages, and serration along the body edges from the posterior third of the cephalic shield to the last body segment. Leanchoilia persephone Simonetta, previously synonymized with L. superlata, is also well represented in the Burgess Shale, and is re-established as a valid species, owing to conspicuous differences from the type species. These are the absence of the diagnostic up-curving snout of the cephalic shield, the absence of carina, the shorter ‘great appendages’, the smooth edges of the body, and its overall shape in dorsal aspect. Leanchoilia superlata and L. persephone may be sexual dimorphs of each other. The ROM collections extend considerably the geographical distribution and stratigraphic range of Leanchoilia in western Canada. [ABSTRACT FROM AUTHOR]

Abstract: Cambrian konservat-lagerstätten are the most significant fossil deposits for our understanding of the initiation of Phanerozoic life. Although many modes of preservation may occur, these deposits most frequently contain nonmineralized fossils preserved in the form of kerogenized carbon films, a rare yet important taphonomic pathway that has not previously been explained for any unit by a comprehensive model. The middle Cambrian Wheeler Formation of Utah, one of these lagerstätten, contains abundant kerogenized preservation of nonmineralized tissues, which occurs within a distinctive taphofacies that accumulated under the following conditions: (1) domination of the siliciclastic fraction by clay-sized particles, (2) close proximity to a carbonate platform, which resulted in mixed carbonate-clay sediments, (3) a well-developed oxygen minimum precluding benthic colonization and burrowing, and (4) relative proximity to oxic bottom-waters, facilitating transport of organisms from a habitable environment to one that favored their preservation. We propose that preservation of nonmineralized tissues in the Wheeler Formation may have resulted from a combination of influences that reduced permeability and, thus, lowered oxidant flux, which in turn may have restricted microbial decomposition of some nonmineralized tissues. Those influences include near bottom anoxia, preventing sediment irrigation by restriction of bioturbation; reducing conditions near the sediment–water interface that may have acted to deflocculate aggregations of clay minerals, resulting in low permeability face-to-face contacts; early diagenetic pore occluding carbonate cements; and an absence of coarse grains such as silt, skeletonized microfossils, fecal pellets, or bioclasts. This model may be applicable to kerogenized preservation of macrofossils in other fossil lagerstätten. [Copyright &y& Elsevier]

Abstract: As the depth and intensity of bioturbation increased through the Proterozoic–Phanerozoic transition, the substrates on which marine benthos lived changed from being relatively firm with a sharp sediment–water interface to having a high water content and blurry sediment–water interface. Microbial mats, once dominant on normal marine Proterozoic seafloors, were relegated to stressed settings lacking intense metazoan activity. This change in substrates has been termed the agronomic revolution [Seilacher, A., Pflüger, F., 1994. From biomats to benthic agriculture: a biohistoric revolution. In: Krumbein, W.E. (Ed.), Biostabilization of Sediments. Bibliotheks and Informationsystem del Carl von Ossietzky Universitat, Oldenburg, 97–105], and the impact of the subsequent development of the mixed layer on benthic metazoans has been termed the Cambrian substrate revolution [Bottjer, D.J., Hagadorn, J.W., Dornbos, S.Q., 2000. The Cambrian substrate revolution. GSA Today 10 (9), 1–7]. Because the Early Cambrian was a transitional time in this substrate revolution, it is hypothesized that benthic metazoans adapted to typical Proterozoic-style soft substrates co-existed with benthic metazoans adapted to more typical Phanerozoic-style soft substrates. Paleoecological examination of the benthic metazoans of the Early Cambrian Maotianshan Shale (Chengjiang) biota and Middle Cambrian Burgess Shale biota, combined with examination of core samples from the rocks in which the Maotianshan Shale biota is preserved, was performed in order to test this hypothesis. Results of the core analysis reveal that the rocks in which the Maotianshan Shale biota was preserved, and therefore the substrate on which it lived, contains almost no evidence for bioturbation of any kind, while the immediately underlying rocks of similar lithology are dominated by intermediate levels of horizontal bioturbation and contain some evidence for the beginning of mixed layer development. These results suggest that benthic metazoans of the Maotianshan Shale biota lived on a typical Proterozoic-style soft substrate, and that environmental conditions associated with the exceptional preservation of the Maotianshan Shale biota may have also suppressed bioturbation levels. Results of the paleoecological analysis indicate that the Maotianshan Shale and Burgess Shale biotas contain mobile and sessile benthic metazoans adapted to typical Proterozoic-style soft substrates and others adapted to typical Phanerozoic-style soft substrates. In addition, the Maotianshan Shale biota is dominated by benthic suspension feeding genera adapted to typical Proterozoic-style soft substrates, indicating that this biota lived during the early stages of the Cambrian substrate revolution. The Burgess Shale biota, however, contains more benthic suspension feeding genera adapted to typical Phanerozoic-style soft substrates than the Maotianshan Shale biota, which suggests that the soft substrate on which they lived had more advanced levels of mixed layer development. This research suggests that the adaptive radiation of benthic metazoans during the “Cambrian explosion” was driven in part by the Cambrian substrate revolution, as benthic metazoans were forced to adapt to the development of the mixed layer in subtidal siliciclastic soft substrate environments during the Cambrian. [Copyright &y& Elsevier]

A slab of Burgess Shale (Middle Cambrian), displaying an incomplete exoskeleton of the large arthropod Sidneyia inexpectans and encompassed by nine specimens of the priapulid worm Ottoia prolifica, is interpreted as a death assemblage, with the worms once living off or feeding around a carcass or freshly moulted instar of Sidneyia. Death is thought to have been caused by an obrution event that preserved the organisms in situ. [ABSTRACT FROM AUTHOR]