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1.
    
Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).  相似文献   

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

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Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia was originally described as the earliest crinoid(?) known from the fossil record. Recently, Conway Morris and Ausich & Babcock have questioned whether Echmatocrinus is in fact an echinoderm, comparing it instead to cnidarians with a polyp-like body and pinnate tentacles, and other authors are beginning to use this reinterpretation. We studied the well-preserved holotype of Echmatocrinus brachiatus, two paratypes, and 18 new specimens recovered from different levels in the Burgess Shale sequence at three localities. All are preserved as pyrite films in dark shale with relatively little relief, suggesting a lightly skeletized body. Complete specimens have a long, slightly tapering, large-plated attachment stalk, a conical cup or calyx with numerous small to medium-sized irregular plates, and 7–10 short arms with heavier plating and (in the holotype) soft appendages alternating from opposite sides of several arms. Several morphologic features indicate that Echmatocrinus is an echinoderm and has crinoid affinities: (1) Sutured plates, shown by darker depressed sutures, slightly raised plate centers, and oriented plate ornament, cover all major parts of the body; (2) reticulate surface ornament in the pyrite film on the plates of all specimens matches the ornament in the Burgess Shale edrioasteroid Walcottidiscus, an undoubted echinoderm, but not the pyritized surfaces of other metazoans in the fauna; (3) this distinctive ornament may represent the surface expression of microporous stereom; (4) possible ligament or muscle pads are present between the arm ossicles to fold and unfurl the more heavily plated arms. Within the echinoderms, only crinoids commonly have a calyx attached by a stalk or stem to the substrate and bear erect, moveable, uniserial arms for feeding. Although Echmatocrinus shows some resemblance to octocorals in overall body shape as an attached suspension feeder, almost all the details are different, indicating that Echmatocrinus is most likely unrelated to this group. All complete specimens of Echmatocrinus are attached to hard substrates, either another fossil or skeletal debris. The new specimens indicate that Echmatocrinus was twice as common (about 0.02%) in the Burgess Shale fauna as previously recorded and represents one of the earliest attached, medium-level, skeletized, suspension feeders or microcarnivores in the fossil record.  相似文献   

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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.  相似文献   

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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, Cambrian Burgess 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.  相似文献   

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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.  相似文献   

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Abstract:  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.  相似文献   

10.
    
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.  相似文献   

11.
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.  相似文献   

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《Zoologischer Anzeiger》2014,253(2):164-178
Sidneyia inexpectans Walcott, 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.  相似文献   

14.
    
The stem‐group priapulid Ottoia Walcott, 1911, is the most abundant worm in the mid‐Cambrian Burgess Shale, but has not been unambiguously demonstrated elsewhere. High‐resolution electron and optical microscopy of macroscopic Burgess Shale specimens reveals the detailed anatomy of its robust hooks, spines and pharyngeal teeth, establishing the presence of two species: Ottoia prolifica Walcott, 1911, and Ottoia tricuspida sp. nov. Direct comparison of these sclerotized elements with a suite of shale‐hosted mid‐to‐late Cambrian microfossils extends the range of ottoiid priapulids throughout the middle to upper Cambrian strata of the Western Canada Sedimentary Basin. Ottoiid priapulids represented an important component of Cambrian ecosystems: they occur in a range of lithologies and thrived in shallow water as well as in the deep‐water setting of the Burgess Shale. A wider survey of Burgess Shale macrofossils reveals specific characters that diagnose priapulid sclerites more generally, establishing the affinity of a wide range of Small Carbonaceous Fossils and demonstrating the prominent role of priapulids in Cambrian seas.  相似文献   

15.
    
Exceptional fossil specimens with preserved soft parts from the Maotianshan Shale (ca 520 Myr ago) and the Burgess Shale (505 Myr ago) biotas indicate that the worldwide distributed bivalved arthropod Isoxys was probably a non-benthic visual predator. New lines of evidence come from the functional morphology of its powerful prehensile frontal appendages that, combined with large spherical eyes, are thought to have played a key role in the recognition and capture of swimming or epibenthic prey. The swimming and steering of this arthropod was achieved by the beating of multiple setose exopods and a flap-like telson. The appendage morphology of Isoxys indicates possible phylogenetical relationships with the megacheirans, a widespread group of assumed predator arthropods characterized by a pre-oral ‘great appendage’. Evidence from functional morphology and taphonomy suggests that Isoxys was able to migrate through the water column and was possibly exploiting hyperbenthic niches for food. Although certainly not unique, the case of Isoxys supports the idea that off-bottom animal interactions such as predation, associated with complex feeding strategies and behaviours (e.g. vertical migration and hunting) were established by the Early Cambrian. It also suggests that a prototype of a pelagic food chain had already started to build-up at least in the lower levels of the water column.  相似文献   

16.
    
The early Cambrian Indian Springs biota of western Nevada, USA, exhibits Burgess Shale‐type (BST) preservation of a diverse array of animal phyla, including the earliest definitive echinoderms. It therefore provides an important window on animal life during the Cambrian radiation. The objective of this study was to analyse the trace metal palaeoredox geochemistry and bioturbation levels of this BST deposit in order to characterize the palaeoenvironmental conditions in which these animals lived and their remains preserved. A total of 28 rock samples were collected from outcrops at three previously reported intervals of exceptional preservation at the Indian Springs locality, as well as from one interval not exhibiting such preservation. An additional 20 random samples were collected from talus for comparison. In the laboratory, the samples were analysed for trace metal palaeoredox indices (V/Cr and V/(V + Ni) ratios). Bioturbation levels were assessed through X‐radiography and petrographic thin sections using the ichnofabric index (ii) method. Additional samples from coeval strata of the Poleta Formation in the White‐Inyo Mountains, CA, that lack BST preservation were also analysed with the same methodology. Results indicate that oxic bottom water conditions dominated during deposition of these strata, despite consistently low bioturbation levels. This pattern holds for intervals with BST preservation and those without. Although ephemeral incursions of low‐oxygen waters may have taken place, there is no evidence for persistent oxygen restriction in these palaeoenvironments. The low levels of bioturbation indicate limited mixed layer development and a redox boundary near the sediment–water interface, likely allowing post‐burial BST preservation to occur even in this setting dominated by oxic bottom waters. Palaeoecological reconstructions and taphonomic hypotheses relating to the Indian Springs Lagerstätte must consider the palaeoredox conditions revealed in this study. With the dispensing of anoxic bottom waters as a requirement for BST preservation, other models proposing a role for clay minerals, the presence of hypersaline brines and the actions of Fe‐reducing bacteria as mechanisms for exceptional preservation warrant renewed consideration.  相似文献   

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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.  相似文献   

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As sessile, benthic filter feeders, brachiopods share an intimate relationship with their chosen substrate. Individuals of Micromitra burgessensis in the Burgess Shale Formation are preserved in life position, attached to a range of hard substrates, including skeletal debris, conspecific brachiopods, sponges and enigmatic tubes. Here we investigate the phenotypic variability of Mburgessensis associated with differing substrate attachments. We apply geometric morphometrics to test for variation by plotting landmarks on the exterior of ventral and dorsal valves of Mburgessensis specimens that are preserved attached to different substrates. Using principal component, canonical variate analyses and anova , we determine that there is some variation in shape related to substrate. Canonical variate analyses, for ventral valves and dorsal valves, indicate that specimens attached to the same substrate are recognizable in shape from specimens attached to other substrate types. The strength of differentiation however, is not robust and combined with our discriminate analysis of separate populations suggests that there is the potential for substrates to exercise only weak control over the morphology of Brachiopoda.  相似文献   

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