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By virtue of remarkable tolerance on hypoxia and adaptive specialization in morphology, diverse hexactinellid sponges prospered in an early Cambrian environment characterized by dysoxic bottom waters documented by black shales. New fossils from the black shale of the Niutitang Formation (basal Stage 3 of Cambrian) in Hunan Province of China, reveal for the first time an articulated body of the sponge Hyalosinica archaica Mehl & Reitner in Steiner et al., 1993, which possesses an ovoid main body and an impressive long stalk. The spicular skeleton includes large diactines that are generally organized as fan-shaped clusters, a few small stauractines and hexactines, and twisted bundles of long monaxons that form the stalk/root tuft. This hexactinellid sponge represents the oldest extinct taxon that took advantage of a long stalk to elevate the main body above the sediment surface and thus to adapt to the oxygen-deficient sea-bottom environment. The long root tuft links Hyalosinica to a series of fossil and recent sponge taxa and proves a deep origin of the stalk-bearing morphology, indicating a likely parallel evolution within the Hexactinellida in response to special environmental pressures. Furthermore, the overall skeletal organization indicates that Hyalosinica, as well as related early ‘rossellimorphs’, are basal stem group representatives of Hexactinellida and probably branched before the extinct Reticulosa and before the two extant hexactinellid subclasses. [ABSTRACT FROM AUTHOR]

Though morphological variation among trilobites has been studied for some time, it is difficult to identify differences between interspecific and intraspecific variations. In this study, the intergeneric, interspecific, and intraspecific variations of the Cambrian oryctocephalid trilobites Duyunaspis and Balangia from Jianhe, South China are analysed using geometric morphometrics. In addition to the number of thoracic segments and the relative size of the pygidium compared to the cephalon mentioned in previous studies, the results in this study suggest that the predominant variation of Duyunaspis and Balangia is the presence or absence of the glabellar furrow. The interspecific variations of Duyunaspis (D. duyunensis, D. paiwuensis and D. jianheensis) mainly lie in the types of facial suture, occipital ring and glabella. Although the differences of D. duyunensis or D. jianheensis from adjacent sections have been observed, these variations may be caused by ontogeny, and do not reach the magnitude of interspecific variations in Duyunaspis. This research provides a better quantitative understanding of the variability in oryctocephalid trilobites. [ABSTRACT FROM AUTHOR]

Since 1990, it has been proposed that olenelloid and paradoxidid trilobites overlapped in age, even though the former represents the traditional lower Cambrian of Laurentia and the latter represents the traditional middle Cambrian of the ‘Acadobaltic Realm’ (i.e., Baltica, West Gondwana) and Siberia. Subsequent studies on biostratigraphical correlation and radiometric ages have supported this conclusion. The recent discovery of the oryctocephalid trilobites Oryctocephalops frischenfeldi Lermontova, 1940, Ovatoryctocara cf. O. yaxiensis Yuan et al. 2009, and Protoryctocephalus? aff. P.? arcticus Geyer & Peel 2011 from the mid-Dyeran Stage portion of the Harkless Formation, Nevada, not only verifies a direct overlap of olenelloid and paradoxidine trilobites, but also indicates that the upper six olenelloid biochrons of the Cordilleran margin of Laurentia are age equivalent to the range of the oldest paradoxidines. This new fauna from the Harkless Formation also allows for a precise biostratigraphical correlation of this level to age-equivalent strata in Greenland, the Siberian Platform, and South China. This correlation indicates that the upper Dyeran and a portion of the mid-Dyeran Stage are biostratigraphically equivalent to the Bonnia–Pagetides elegans Biozone of Greenland, the Ovatoryctocara Biozone (which contains Paradoxides (s.l.) in its lower part) of the Siberian Platform, and the Protoryctocephalus arcticus to Bathynotus kueichouensis–Ovatoryctocara sinensis biozones of the South China palaeocontinent. [ABSTRACT FROM AUTHOR]

The ontogenetic sequence of an early Cambrian redlichiid trilobite Bathynotus kueichouensis is presented on the basis of numerous articulated specimens from Guizhou, China. The body regionalization of B. kueichouensis witnessed a wide variation in the morphology of trunk segments between different thoracic regions, referred to as the ‘thoracic region 1-4’. A greater size variation of these long pleural spines is recognized from the earlier meraspid degrees in a gradual appearance or disappearance until the holaspid period. The thoroughly inverse variation of these pleural spines in size and form is seen for the first time during the process of trilobite segment formation, which offers an opportunity to trace the evolutionary path of trilobite body structures, associated with their functional morphology and behavior. The ontogenetic strategy of B. kueichouensis is broadly similar to those of other redlichioid representatives where the full ontogenetic sequence exhibited a balanced rate in segment expression and liberation. Such a developmental mode may be the reason why most redlichiids possess a micropygous body patterning during ontogeny and provides insights into the controls of early arthropod developmental segmentation during evolutionary process. [ABSTRACT FROM AUTHOR]

Archaeocyaths were one of the major animal groups contributing to the Cambrian Explosion. Despite an overall poriferan appearance of their hypercalcified skeletons, they exhibit a number of peculiar features such as the presence of septa and an absence of spicules. This sets archaeocyaths apart from general trends in early poriferan evolution. Fossils recently discovered from the early Cambrian Guanshan Lagerstätte of South China exhibit features typical for the suborder Archaeocyathina whilst possessing a silicified skeleton associated with carbonaceous material. This rigid fibrous skeletal framework, on the other hand, fits the diagnosis of the supposed ancestral aspiculate demosponges, the family Vauxiidae. By a detailed description of this specimen and a direct morphological comparison between the Archaeocyathina and the Vauxiidae, we suggest that the supposed Cambrian representatives of horny demosponges, that is, vauxiids, probably, were a progeny lineage of archaeocyathines. They survived the extinction of hypercalcified archaeocyaths due to their ecological adaption to siliciclastic environments. The existence of hypercalcified dictyoceratid ('keratosan') sponge Vaceletia, well known as a 'living archaeocyath', supports such a possibility. [ABSTRACT FROM AUTHOR]

Microscope, cathodoluminescence, chemical and oxygen isotope studies have been conducted to determine the original mineralogy and diagenetic alteration of phosphatic grasping spines of Cambrian chaetognaths. The obtained data, along with a comparison to the composition of conodont apatite, show the presence of a few generations of diagenetic phosphate phases. A thin, outer layer of the spines is composed of very early diagenetic phosphate, which incrusted the original cuticle layer contributing to the preservation of an original shape and structural details of the fossils. This process can be linked to the 'Orsten'‐type phosphatization mainly known from Lower Palaeozoic strata. Successive phosphate generations are observed within the middle and inner layers as well as internal cavities of the spines. Plastic deformations and diagenetic features of the spines show that they were originally composed of slightly flexible chitinous organic matter similar to that of modern chaetognaths. This conclusion is substantiated by rare findings of purely organic remnants of Cambrian chaetognath grasping spines. [ABSTRACT FROM AUTHOR]

It is important to understand moulting behaviours in trilobites through different growth stages. Most studies have focused on patterns of moulting in trilobites based on a few exuviae. The present study is based on 177 disarticulated specimens of Arthricocephalites xinzhaiheensis from the Balang Formation (Cambrian Series 2, Stage 4, Guizhou Province, South China). The abundance of disarticulated assemblages allows the distinction between carcasses and exuviae to be made with confidence and enables a detailed exploration of moulting behaviours. About 96.6% of exuviae of both meraspides and holaspides of Ar. xinzhaiheensis show a disarticulated cranidium characterizing Henningsmoen's configuration. The trilobites commonly used the cephalic sutures and cephalothoracic joint for moulting, involving lateral rotation or inversion of the cranidium to create a wide exuvial gape. Only one specimen displays the Somersault configuration characterized by an overturned lower cephalic unit and the data also show that trilobites employing the cephalic sutures to create an anterior exuvial gape are rare in holaspides. Though some specimens are similar to Salter's configuration with an inverted cephalon with respect to the thoracopygon, the specimens show closed cephalic sutures and thus dissociated or inverted cephala are likely to represent disturbed carcasses. Several other species of oryctocephalid trilobites display Henningsmoen's configuration indicating that exuviation by disarticulated cranidium to enlarge the exuvial gape was prevalent in the family from Cambrian Series 2 Stage 4 to the succeeding Wuliuan Stage of the Miaolingian Series. [ABSTRACT FROM AUTHOR]

Completely enrolled, phosphatized, 3‐D specimens of the eodiscinid trilobite Tsunyidiscus yanjiazhaiensis from Cambrian Stage 3 of South China exhibit much morphological detail and show variation in delicate coaptative structures associated with their trunk segmentation. The relationship between enrolment mechanism and trunk segmentation during ontogeny confirms a unique developmental pattern among the early Cambrian eodiscoid trilobites, revealing how these animals controlled the rate of segment increase and release during post‐embryonic development in their life cycles. [ABSTRACT FROM AUTHOR]

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

Exceptionally preserved, silicified and articulated complete shells of the rhynchonelliform kutorginate brachiopod Nisusia sulcata are redescribed from the middle Cambrian (Series 3) Marjum Limestone, Utah. Cylindroid sausage‐like protrusions, emerging posteriorly between the valves, were originally interpreted as faecal in origin, but restudy under the SEM shows that these features represent silicified pedicles as they are attached in situ to other Nisusia. The Nisusia host most likely was alive at the time of attachment. Restudy of the pedicles of Nisusia provides new phylogenetic information on the anatomy of the earliest rhynchonelliforms. The silicified pedicles differ considerably from the pedicles of living crown group rhynchonelliforms in being strongly annulated, distally tapering, and were likely to have been rather more flexible. The Nisusia pedicles are more similar to the exceptionally preserved pedicles from other Cambrian rhynchonelliform brachiopods, including Kutorgina, Longtancunella and Alisina, but these emerge from the ventral apical foramen rather than from between the valves as in Nisusia. Although generally similar, these two types of pedicles are unlikely to represent homologous structures as Nisusia is provided with both an apical foramen (possibly larval attachment) and a posterior adult pedicle. The similarities may be explained by similar type of accretionary growth from two different types of epithelia. The Nisusia‐like pedicle appeared early within the kutorginates and rhynchonellates. The discovery of hollow spines in Nisusia sulcata further supports the generic assignation of the species. [ABSTRACT FROM AUTHOR]

Brachiopods are among the first animal phyla to emerge from the Cambrian Explosion, rapidly diversifying to all major palaeocontinental blocks within 20 million years. The group underwent another steep rise in diversity during the Ordovician, and their relative abundance and diversity made them one of the most successful invertebrate groups during the entire Palaeozoic. During this time, brachiopods lived in a range of environments and represented a significant component of marine ecosystems, yet information regarding their modes of life and ecology is somewhat limited. Recent studies, primarily from the Chengjiang and Burgess Shale Lagerstätten, have revealed that by the Middle Cambrian (Series 3, Stage 5) brachiopods from across the phylum had already developed a range of ecological strategies and life modes. Cambrian brachiopods occupied distinct trophic niches on soft and hard substrates and exhibited at least five types of lifestyles: pedicle attachment, pedicle anchoring, cemented, free‐lying and semi‐infaunal. Comparisons with Ordovician benthic assemblages show that despite the explosion of brachiopod taxa witnessed in the Ordovician, with the exception of the appearance of burrowing brachiopods, life strategies of brachiopods remained largely the same. Indicating that the majority of life modes observed in brachiopods had rapidly evolved and was already in place prior to the Great Ordovician Biodiversity Event. [ABSTRACT FROM AUTHOR]

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]

A detailed exploration of growth and trunk segmentation of the oryctocephalid trilobite, Duyunaspis duyunensis Chang & Chien in Zhou et al. 1977, from the lower Cambrian (Stage 4, Series 2) Balang Formation in western Hunan Province, South China, is presented. Because of the excellent preservation, the complete post-protaspid ontogenetic series from merapsid degree 0 to the holaspid phase is described. The ontogenetic series reveals new information on morphological changes such as the migration of the posterior branch of the facial sutures (from proparian to opisthoparian) and contraction of the posteromedial notch in the pygidium. The abundance of articulated specimens available from a narrow stratigraphical interval makes this material singularly useful for studying the morphogenesis and post-embryonic growth of D. duyunensis in comparison with other oryctocephalids. Strong evidence that multiple numbers of pygidial segment are recognized in each meraspid degree as well as in the holaspid period showed unusual intraspecific variability in the rate of trunk segmentation, providing insights into how Cambrian subisopygous trilobites controlled their body patterning, including size, shape and trunk segment number in both thorax and caudal plate during growth. [ABSTRACT FROM AUTHOR]

One of the principal biostratigraphical tools used in the lower Palaeozoic, and especially the Cambrian System, is trilobites. Historically, both polymerids and agnostoids have commonly been included as 'trilobite' groups, although currently the question of whether they comprise a monophyletic group or are polyphyletic is unsettled. Beginning in the late 1800s, the base of the Cambrian System was marked by the lowest occurrence of olenelline trilobites. Discovery of a rich pre-trilobite metazoan record in the mid-1900s led to significant modification of that concept, but polymerids and especially agnostoids still rank among the leading biostratigraphical and chronostratigraphical guides through much of the Cambrian. Chronostratigraphical sub-division of the trilobite-bearing part of the Cambrian System derives largely from biostratigraphical, ecological and evolutionary concepts about agnostoids and polymerids introduced in the 1940s to 1970s by A.H. Westergård, A.R. Palmer and R.A. Robison. Westergård introduced a zonation for Scandinavia that was largely based on agnostoids. Palmer explained the distribution of Cambrian trilobite faunas in terms of restricted and unrestricted access to open oceans. Together, these ideas coalesced in Robison's recognition of separate zonation schemes for restricted-shelf polymerids, open-shelf polymerids and open-shelf (cosmopolitan) agnostoids. Palmer also introduced the concept of biomeres, which placed sharp limits on biostratigraphical intervals recognizable from trilobites. Global correlation in the upper half of the Cambrian today depends to a large extent on the ranges of agnostoids and some polymerids characteristic of open-shelf to slope areas, ones that facilitate identification of precise intercontinental tie points. Agnostoid and polymerid biostratigraphy is now being integrated with information about coastal onlap and eustatic sea-level history, geochemical cycling and other data to provide a more complete understanding of the early Palaeozoic biosphere and its complex physico-chemical context. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Synchrotron radiation X-ray tomographic microscopy ( SRXTM) was used to virtually dissect and peel the shields off of the microscopic, bivalved phosphatocopine crustaceans in the Cambrian 'Orsten' type of preservation of Sweden. Doing so opened up for an array of concealed internal structures to be observed in a fully enclosed specimen of Hesslandona ventrospinata and a semi-enclosed specimen of Hesslandona angustata. For comparison, also a head-larva stage specimen of H. angustata, with shields in 'butterfly position', was analysed. The X-ray tomographic data sets revealed excellently preserved structures, such as labrum, sternum, antennae, mandibular and post-mandibular limbs with their minute setae, all of which were more or less disguised by the enclosing shields. This, moreover, allowed assignment to growth stages of the specimens, which is impossible based solely on external morphology and size. Micro-spherules observed inside the shields of the semi-enclosed H. angustata specimen may represent remains of food particles, and the feeding biology of phosphatocopines is discussed in detail. Our analyses suggest that phosphatocopines were particle feeders. The SRXTM technique offers the ability to three-dimensionally reconstruct the morphology in high resolution, construct virtual serial sections and study concealed structures. The resulting data allow for new structures to be revealed for previously known taxa and for new taxa to be identified, with the added benefit of not destroying the specimens in the process. Hence, we do not longer have to rely on serendipitous finds of broken and/or open phosphatocopine specimens to elucidate their diagnostic ventral morphology. [ABSTRACT FROM AUTHOR]

The problematic mollusc Wiwaxia is perhaps the most widely distributed non-mineralized Cambrian metazoan, but has only been reported from palaeotropical latitudes. Here, we describe mid-Cambrian (Drumian, c. 504 Ma) sclerites and possible tooth arrays from the northern Llanos Basin, Colombia, recovered from drilled ditch cuttings by palynological processing - demonstrating that pristine material and low-manipulation processing are not essential to the recovery of Small Carbonaceous Fossils. This, the first report of Wiwaxia from South America, substantially expands Wiwaxia's geographic range into the high palaeolatitudes. [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]

Fossil skeletal apatites vary in their composition and can yield mixed biochemical, environmental and diagenetic information. Thus, it is important to evaluate the diagenesis spatially inside the skeleton. We study the cross sections of shells of the Furongian lingulate brachiopod Ungula ingrica from Estonia using the Attenuated Total Reflectance -- Fourier Transform Infrared (ATR-FTIR) microspectroscopic and energy dispersive spectroscopic (EDS) mapping and show for the first time that different structural laminae of the shell have different chemical compositions. Compact laminae are rich in PO43-, Na, Mg and poor in F and Ca. Porous (baculate) laminae are rich in carbonate anions, Ca and F, but contain less Na and Mg. The ATR-FTIR spectra show further differences in the v2 carbonate region, where the IR band at 872 cm-1 in compact laminae is replaced by a strong band at 864 cm-1 in baculate laminae. The changes in shell apatite suggest different origins of the apatite phases. Compact laminae are likely chemically less altered and could potentially carry more reliable palaeoenvironmental or geochemical information than the apatite in baculate laminae, which is mostly authigenic in its origin. [ABSTRACT FROM AUTHOR]

The Cambrian Series 3 Zhangxia Formation in Shandong Province, North China, includes small-scale lithistid sponge-microbial reefs. The lithistid sponges grew on oolitic and bioclastic sediments, which were stabilized by microbial activities. The relative abundances of microbial components (e.g. calcimicrobe Epiphyton and stromatolites) vary among the reefs. However, the microbial components commonly encrusted or bound the lithistid sponges, formed remarkable encrustations on the surfaces of the sponges. Epiphyton especially grew upward and downward. The lithistid sponges thus provided substrates for the attachment and development of microbes, and the microbes played essential roles as consolidators, by encrusting reef-building sponges. Additionally, the lithistid sponges were prone to degradation via microbial activities and diagenetic processes, and were thus preserved as micritic bodies, showing faint spicular networks or abundant spicules. Such low preservation potential within the reef environment obscured the presence of the sponges and their widespread contribution as reef-building organisms during the Cambrian. During the prolonged interval after the demise of archaeocyaths, purely microbial reefs, such as stromatolites and thrombolites have been considered to be the principal reef builders, in association with rare lithistid sponge-microbial associations. However, recent findings, including those from Shandong Province and Korea, suggest that the lithistid sponge-bearing reefs were more extensive during the Epoch 3 to the Furongian than previously thought. These lithistid sponge-microbial reefs were precursors of the sponge-microbial reefs that dominated worldwide in the Early Ordovician. [ABSTRACT FROM AUTHOR]

Feeding behavioural patterns have the potential to shed light on evolutionary trends, especially at the critical point of the Cambrian Explosion. In this context, we present specimens of the tubiform priapulid Selkirkia sinica, preserved in situ, collected from the lower Cambrian (Stage 3) Xiaoshiba Lagerstätte, which show the diverse morphology of its intestinal contents within the dark-coloured gut and the formation of faecal pellets. The two digestive modes displayed by S. sinica suggest that it is a putative dietary generalist rather than a typical scavenger as previously thought. Moreover, the asymmetry exhibited by the paired caudal appendages of S. sinica provides direct evidence of the development of dorsoventral ( DV) differentiation in Selkirkia, which aids our understanding of the lifestyle of this tubiculous worm. [ABSTRACT FROM AUTHOR]

The use of carbon isotope excursion in Cambrian stratigraphical correlation is a standard practice at both the intercontinental and intracontinental scales. The Steptoean positive carbon isotope excursion ( SPICE) is one of the prime examples in this regard in correlating the base of the Paibian Stage and Furongian Series. A lack of definite SPICE evidence in the North China craton has been a challenge in precision correlation between North China and other palaeo-continents. This study provides new carbonates carbon isotope data from the type locality of the Changshan Formation in Hebei Province, North China. Our new δ13Ccarb data provide new objective evidence for the presence of the SPICE in North China. The sampling section is relatively condensed, and the interval of the SPICE curve is less than one and half-metres after analysing 64 samples (the sampling interval within the SPICE is less than 10 cm). The onset of the SPICE curve in Tangshan, Hebei, occurs in the barren interval between the Neodrepanura and Chuangia trilobite zones. Based on this study and previous work, this could imply the middle part of the Prochuangia-Paracoosia trilobite Zone in North China and can be correlated with the base of the Paibian Stage and Furongian Series. [ABSTRACT FROM AUTHOR]

Enrolment was a major defensive strategy for trilobites that significantly contributed to their evolutionary success. The ability to enrol also helped to constrain the morphological evolution of trilobites, which in part was driven by the need to improve this capability to encapsulate the soft parts of the body within the mineralized dorsal exoskeleton. Here, we describe a unique example of gut content fossilization in an enrolled trilobite from the Cambrian of China, and we propose a taphonomic scenario that considers the possible implication of enrolment in this exceptional preservation. A micro-facies analysis indicates that the specimen was entombed during an obrution event and remained intact due to limited infaunal activity. The encapsulation of the body did not prevent the decay of soft tissues, but it permitted the delicate gut content to be protected during diagenesis. In isolating the decaying soft tissues from the external environment, enrolment might also have favoured the establishment of microenvironmental conditions conducive to the precipitation of pyrite framboids. Within the gut, the formation of such crystals may have led to the consolidation of the ingested material. These results suggest that fossilized gut contents might be quite common in enrolled trilobites. Textural and compositional analyses reveal that the gut content is similar to the sediment surrounding the fossil except for the presence of pyrite framboids that indicate higher initial organic matter content. The complete enrolment of the body argues against an accidental ingestion of this material or a diagenetic origin for it. Accordingly, detritus feeding habits are inferred for this ptychopariid trilobite. [ABSTRACT FROM AUTHOR]

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages ( SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well-preserved isolated plates of Onychodictyon sp. from the Lower Cambrian ( Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny. [ABSTRACT FROM AUTHOR]

Terfelt, F., Ahlberg, P. & Eriksson, M.E. 2011: Complete record of Furongian polymerid trilobites and agnostoids of Scandinavia - a biostratigraphical scheme. Lethaia, Vol. 44, pp. 8-14. So far, 112 polymerid trilobite species/subspecies and 13 agnostoid species/subspecies have been recorded from the Furongian (upper Cambrian) of Scandinavia. For the first time, their zonal occurrences are summarized in a biostratigraphical scheme serving as a practical synopsis for students of this interval in time. Ninety-six of the recorded polymerid trilobite species/subspecies belong to the family Olenidae whereas the remaining 16 are distributed across eight other families. Levels of increased speciation and low diversity (including stratigraphical range gaps) are conspicuous and these may be correlated with recorded physical and chemical anomalies. □ Agnostoids, biostratigraphy, Cambrian, Furongian, polymerids, Scandinavia, trilobites. [ABSTRACT FROM AUTHOR]

Among 381 specimens of Cambrian stalked echinoderms from eastern Guizhou, China examined, several slabs ( n = 19) contain either dumbbell-shaped or v-shaped echinoderm clusters. Four slabs of Globoeocrinus globules Zhao, Parsley & Peng, 2008 from the middle-upper part (Cambrian Series 3 portion) of the Kaili Formation are prepared to reveal the attachment sites. Articulated gogiid echinoderms are reported to be attached to both sides of inarticulate (organophosphatic) brachiopods; thus, allowing me to interpret that the larvae of these gogiids were capable of attaching to live benthic brachiopods. This study documents the one of the earliest examples of echinoderms employing secondary tiering, which elevates an organism higher into the benthic boundary layer. Many of the gogiid echinoderm pairs attached to a live brachiopod are similar in size, indicating they were from a single larval spatfall. [ABSTRACT FROM AUTHOR]

Simultaneously with the development of animal body plans, probably before the Precambrian, there was an explosive diversification of visual systems. Competition of performance in these visual systems was a critical factor in the evolution of life systems. Here we analyse the visual system in the lobopod Miraluolishania haikouensis ( Liu et al., 2004 ) from the Lower Cambrian Chengjiang Lagerstätte, Kunming, China. It consists of a very small eye with a miniscule lens. A physical problem lies in the fact that due to the usual refractive conditions of such a lens, it hardly represents an improvement of the visual quality over the basal pit- or pinhole camera eyes. To develop such a lavish visual system, however, would not have been of any value, if it achieved no more than an equal level or represented even a retrograde step in evolutionary progress. We show how this system may have allowed pattern recognition even under poor light conditions. Optimization of such a tiny eye is costly but is not ‘a wasted effort’ in evolution. In M. haikouensis ( Liu et al., 2004 ), an excellently adapted miniscule visual system has become possible. [ABSTRACT FROM AUTHOR]

The early Middle Cambrian Monastery Creek Phosphorite (Beetle Creek Formation, Queensland, Australia) contains an assemblage of disarticulated echinoderm ossicles that are exquisitely preserved. Amongst this material we recognize pelmatozoan brachials, radials, basals and holomeric columnals. Although we cannot reconstruct the complete animal with precision, these elements represent the oldest known pelmatozoan with crinoid-like appendages. Key elements include isotomously to heterotomously branched uniserial appendage plates with a tripartite adoral food groove, a longitudinal central canal interpreted as housing entoneural nerve, and differentiated articulation facets. There are also epispire-bearing radials bearing one to four arm insertion-facets, each one pierced by a central neural canal. These canals run internal towards the oral area beneath the external food groove. Co-occuring material includes single truncated cone-shaped basals and holomeric columnals, both with a similar articulation pattern, and irregular, epispire-bearing thecal plates. This mosaic of crinozoan (uniserial isotomous to heterotomous arms with neural canal), blastozoan (epispire-bearing thecal plates, appendage leading to oral thecal food groove without direct connection with body cavity) and apomorphic characters (circumoral instead of basal entoneural plexus) is unexpected and demonstrates that crinoid-like pelmatozoans with uniserial, branched arms appeared significantly earlier than the Tremadocian, when the first articulated crinoid skeletons are found. It also raises questions about the polyphyletic appearance of feeding appendages among pelmatozoan echinoderms. [ABSTRACT FROM AUTHOR]

Shells of Oikozetetes and isolated halkieriid sclerites from a section of the lower Cambrian Mernmerna Formation in the Flinders Ranges, South Australia, are tentatively considered as being derived from the same scleritome. Details of shell morphology and the possible combination of biomineralized shell and sclerites suggest that Oikozetetes, if interpreted correctly, is closely related to Halkieria. A new interpretation of Oikozetetes shell morphology, in addition to the first report of paired muscle scars on the interior surface, sheds new light on the possible functional morphology of halkieriid shells and the means of attaching the shell to the body. The occurrence of Oikozetetes in South Australia extends its biostratigraphic range to the lower Cambrian and biogeographic range to East Gondwana. [ABSTRACT FROM AUTHOR]

Pagetia is the most abundant trilobite in the Kaili Formation (lower to middle Cambrian). During the course of studying the museum collections of Kaili trilobites ( n > 1000), a cluster containing 22 pagetiid individuals at various growth stages is noted. Specimens record the growth range from the degree 0 meraspid to late holaspid phases. Based on the relative completeness of the moults except for missing free cheeks, these specimens are interpreted as intact exuviae that had undergone minimal transport prior to burial. If Kaili pagetiids had pelagic or planktonic living habits as previously suggested, it would be difficult to explain the presence of intact exuviae in clusters. Therefore, Kaili pagetiids are interpreted here as having a benthic mode of life after the onset of the meraspid phase. [ABSTRACT FROM AUTHOR]

A core drilling (Andrarum-3), from the classical locality at Andrarum, Scania, southernmost Sweden, penetrated a 28.90-m-thick Cambrian succession. The core comprises dark grey to black, finely laminated mudstones and shales with early concretionary carbonate lenses (stinkstones or orsten) and a few primary carbonate beds. The middle Cambrian (provisional Series 3) part of the core comprises 17.35 m, whereas the Furongian Series (upper Cambrian) part covers the remaining 11.55 m. Nineteen trilobite and two phosphatocopine genera are present in the middle Cambrian, whereas the less diverse Furongian interval yielded four trilobite and three phosphatocopine genera. Other, less frequent, faunal elements include conodonts ( s. l.), brachiopods, sponge spicules, bradoriids, and coprolites. Trilobites and phosphatocopines were used to subdivide the core into seven biozones ranging from the Ptychagnostus atavus Zone to the Parabolina spinulosa Zone ( P. spinulosa Subzone). Carbon isotopic analyses (δ13Corg) through the core show two important excursions, the negative DrumIan Carbon isotope Excursion (DICE) in the Pt. atavus Zone, and the Steptoean Positive Carbon Isotope Excursion (SPICE) beginning near the first appearance of Glyptagnostus reticulatus and extending upward into the Olenus and Agnostus ( Homagnostus) obesus Zone. The DICE displays a peak value, in the samples at hand, of –30.45‰δ13Corg in the lower part of the P. atavus Zone. The δ13Corg values increase through the overlying L. laevigata and A. pisiformis zones and display peak values of c. –28.00‰δ13Corg in the lowermost Furongian Olenus wahlenbergi and O. attenuatus subzones. Thereafter the values decrease significantly through the O. scanicus Subzone. Both isotopic excursions have been documented from several palaeocontinents, but never before from Baltica. Moreover, for the first time these excursions are recorded from organic matter in an alum shale setting. The recorded shift of +1.50–2.00‰δ13Corg is approximately half the magnitude of the SPICE documented from other regions. This discrepancy may be related to temporal variations in the type, origin, or diagenesis of the organic fraction analysed. [ABSTRACT FROM AUTHOR]

The Lower Cambrian Eriboll Formation of northwest Scotland is renowned for the high density, low diversity trace fossils ( Skolithos ichnofacies) found in its upper Pipe Rock Member. Ichnofabric analysis of the member indicates that relatively small examples of Skolithos terminating at the same foreset boundary were formed during a brief colonization window after a single depositional event, that particularly long Skolithos specimens are equilibrichnia, and that palimpsests of Skolithos represent the marginal, slightly deeper water fringes of the Pipe Rock Member depositional environment. Nearest neighbour analysis, however, suggests that such palimpsests were uncommon. A much more diverse trace fossil assemblage is present in the overlying Fucoid Member (An t-Sròn Formation), comprising Cruziana barbata, Dactylophycus, Didymaulichnus, Halopoa imbricata, ? Margaritichnus, Monocraterion, Monomorphichnus, Palaeophycus striatus, P. tubularis, ? Phycodes, Planolites montanus, ? Polarichnus, Rusophycus ramellensis, ? Psammichnites, Skolithos and various unidentified traces, and represents the Cruziana ichnofacies. Above the Fucoid Member, the Salterella Grit Member ichnofauna is more impoverished, yielding only Cruziana, Monocraterion, Rusophycus, Skolithos and ? Spirophyton. The ichnological variations between the Pipe Rock, Fucoid and Salterella Grit members are interpreted as being driven by changes in sea level. The low trace fossil diversity in the Pipe Rock Member indicates opportunistic colonization of laterally extensive, shoreface sediments deposited by regular influxes of terrigenous material, which were overlain by more distal, ichnologically diverse sediments (Fucoid Member) as sea level rose. A minor regression then caused an increase in terrigenous sediment input, producing an impoverished, proximal Cruziana ichnofacies (Salterella Grit Member). [ABSTRACT FROM AUTHOR]

One hundred and twenty-five linguliformean brachiopods of late Marjuman (Cambrian) age with shell perforations, presumably caused by predation, were recovered from shallow-water limestones at two localities of the Deadwood Formation in the Black Hills of South Dakota, USA. Three-quarters of the perforated valves belonged to taxa in the order Acrotretida, while one-quarter of the specimens belonged to those of the order Lingulida. This is the first report of predation of fossil lingulids. In both orders there was a marked selection for valve type. Ninety-five per cent of all perforations of acrotretids were in the ventral valve, while 87% of all perforations of lingulids were in the dorsal valve. The highest rate of predation of collected acrotretids, at any stratigraphic horizon, was 22%, while the rate of predation of collected lingulids, at any given stratigraphic horizon, was as high as 9%. Half of the perforated valves had round holes with a sharp, non-beveled exterior edge, and half had irregularly shaped holes with chipped interior edges. The former type is attributed to either boring or a swift, piercing percussive strike, while the latter is attributed to a smashing percussive strike with a blunt appendage. A third type of perforation consisting of minute, roughly circular holes is thought to be too small to be the work of predators, and is assumed to be the result of an encrusting organism or parasite. The different types of perforation seen in the brachiopod valves indicate that there was more than one type of predator attacking them, including possibly one of the earliest durophages. Various hypothetical predators are suggested as potential candidates for causing the shell perforations. The criteria for their selection as possible linguliformean predators include possessing the ability to produce one of the two types of shell perforation, and being small enough to warrant preying on small (< 2 mm) brachiopods. [ABSTRACT FROM AUTHOR]

The Furongian (upper Cambrian) Leptoplastus Zone marks a time of critical changes in the evolution of olenid trilobites. This zone, unexposed at Andrarum in Skåne, southern Sweden, has been re-excavated and the sequence of faunas and sediments logged in detail. The faunal succession accords with that previously described from borehole cores by Westergård, and the subzones of L. paucisegmentatus , L. raphidophorus , L. crassicornis , L. ovatus , L. angustatus , and L. stenotus have been recognized. In the first two subzones the olenid assemblages are monospecific. At the base of the L. crassicornis Subzone more than one species is present and morphotypes with long genal spines appear for the first time. Faunal turnover is rapid, but the incoming of new species is invariably linked to an abrupt change in sedimentation, or follows an unfossiliferous interval; species either arose or migrated in after a time of environmental perturbation. Particular faunal associations are often confined to discrete sedimentary packages though some species may range through a succession of sedimentary changes. Leptoplastus crassicornis has very long genal spines, adapted for resting on the sea floor; it may have competed with the coeval, and very similar, L. angustatus . Subsequently, L. angustatus is accompanied by the stout-bodied, short-spined L. ovatus , which presumably occupied a different niche within the same environment. Leptoplastus stenotus is convergent on the much earlier L. paucisegmentatus , and likewise is found as a monospecific assemblage, presumably being adapted to a similar niche. [ABSTRACT FROM AUTHOR]

The increase in the depth and intensity of bioturbation through the Proterozoic-­Phanerozoic transition changed the substrates on which marine benthos lived from being relatively firm with a sharp sediment-water interface to having a high water content and blurry sediment-water interface. Additionally, 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', and its impact on benthic metazoans has been termed the 'Cambrian substrate revolution'. The shallow marine phosphorites of the Lower Cambrian Meishucun Formation of southwest China contain evidence suggestive of the presence of seafloor microbial mats. This evidence includes abundant and distinctive red-colored bedding planes enriched in heavy iron minerals and mica, interpreted as resulting from mat-decay mineralization and mica trapping by microbial mats. The radular grazing trace fossil Radulichnus is also found in this formation, indicating a firm, microbial mat-bound substrate. These radular scratches are always preserved with circular impressions around 10 cm in diameter, possibly the fossils of soft-bodied organisms. The first relatively intense bioturbation in this region is found in this formation and is dominated by horizontal Thalassinoides burrows, which could represent undermat mining behavior. The evidence for the presence of microbial mats in the Lower Cambrian Meishucun Formation, and for metazoan lifestyles associated with such mat-bound seafloors, reveals that normal marine environments dominated by typical Proterozoic-style soft substrates still existed during the Cambrian substrate revolution. [ABSTRACT FROM AUTHOR]

Kullingia is considered a key taxon in demonstrating the presence of terminal Proterozoic-early Cambrian chondrophorine hydrozoans. However, Kullingia concentrica from the Lower Cambrian of northern Sweden possesses several features that show that it is not a body fossil but that it was formed by current or wave-induced rotation of an anchored tubular organism, possibly a sabelliditid. A scratch circle interpretation applies also to several other reports of Lower Cambrian Kullingia, including Kullingia delicata from the Chapel Island Formation of Newfoundland. Given the considerable number of problematic fossils that have been interpreted as chondrophorines, it is difficult to put an age on the oldest fossil chondrophorines, but it may be as late as the Devonian. Overall, scratch circles appear to be rarely preserved. The occurrence of these scratch circles in Lower Paleozoic storm deposits is probably related to low levels of bioturbation that enhanced both the likelihood of formation and preservation of these structures. [ABSTRACT FROM AUTHOR]

The newly examined Lower Cambrian strata in the Laisvall-Storuman area, central Swedish Caledonides, yield diverse and stratigraphically significant phytoplanktic organic-walled microfossils (acritarchs) associated with the olenellid trilobites of Holmia sp. that appear to be at the lowermost horizon ever recorded in the Caledonides. The acritarchs, recovered throughout the Grammajukku Formation, are taxonomically reviewed in the context of regional and global trends of phytoplankton diversity. The assemblages are assigned to acritarch zones Skiagia ornata-Fimbriaglomerella membranacea and Heliosphaeridium dissimilare-Skiagia ciliosa, corresponding to the Schmidtiellus mickwitzi and Holmia kjerulfi trilobite zones. The records of acritarch radiations and appearances of trilobites and other faunas in Baltica, and on a more interregional scale, are biochronologically correlated, revealing tightly coupled evolutionary events among primary producers and consumers. The timing of early diversifications of trilobites in various faunal provinces is discussed and the relative age of the oldest known non-mineralized arthropods from the Zawiszyn Formation in Poland is estimated. [ABSTRACT FROM AUTHOR]

The function of the digestive system of naraoiid arthropods is interpreted in the light of new observations on Early Cambrian specimens from China and detailed comparisons with Recent crustaceans and other arthropods. In naraoiids, paired tubular diverticulae ending as blind caeca are present along the entire midgut, and are interpreted as sites for the secretion of digestive enzymes. Naraoia bears one pair of long, ramifying, distensible diverticulae, possibly used for both food storage and digestion as suggested by Recent analogues (e.g. branchiuran and isopod crustaceans and limulids). Naraoiids were probably epibenthic scavengers/predators rather than mud-eaters. They were either opportunistic intermittent feeders (Naraoia) or more regular feeders (Misszhouia). The mud-fills of the alimentary canals are likely to be artefacts due to taphonomic and weathering processes or, less likely, to sediment ingestion by animals trapped alive in turbiditic flows. The case study of naraoiid arthropods adds to other fossil evidence supporting the idea that predation played a key role in the Early Cambrian food-webs and that organs adapted for this purpose had already reached a high level of diversity and anatomical sophistication. [ABSTRACT FROM AUTHOR]

A phosphatized bilaterally symmetrical, spirally coiled conch from the Lower Cambrian phosphates of the Meishucunian Stage of eastern Yunnan, China, is composed of two well-preserved layers. The outer one is built of longitudinal mineralized fibres not strictly parallel to each other, but may anastomose or branch. The fibres form comarginal ribs with transversal grooves between them, corresponding to growth stages. In the ribs, the fibres are arranged closely and are steeply inclined towards the apex, while in the grooves the fibres are subparallel to the conch surface and more loosely packed. Small elliptic and triangular holes are situated within and between the fibres of the outer layer. The inner layer consists of transversal fibres running around the conch. They are almost parallel to each other but partly separated by narrow discontinuous slits. Despite overall morphological similarity of the conch to shells of some early Cambrian molluscs, the observed orthogonal configuration of longitudinally oriented fibrous structures of the outer layer and transversal fibres of the inner layer is found in orthothecid hyoliths, phosphatized microstuctures of which have been described from the Lower Cambrian of the Siberian Platform. The new material from the Yangtze Platform supports a conclusion of a distinctive type of microstructure available in hyoliths different from molluscan microstructures available from the Lower Cambrian. [ABSTRACT FROM AUTHOR]

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]

Tubular agglutinated fossils of Platysolenites antiquissimus Eichwald, 1860, P. cooperi n.sp. and Spirosolenites spiralis Glaessner, 1979 are examined from selected lower Cambrian successions in Avalonia (England; Wales; Newfoundland) and Baltica (Russia and Estonia). Platysolenites cooperi n.sp. is shown to extend below the first appearance of P. antiquissimus in the Neoproterozoic-Cambrian boundary stratotype region, SE Newfoundland, and to occur at higher stratigraphic levels in Wales and Finnmark. Taphonomic, teleological and ultrastructural studies on well-preserved material are consistent with a similar grade of organization and mode of life for P. antiquissimus and the living astrorhizacean foraminiferid Bathysiphon. However, agglutinated proloculi are here described from both rectilinear P. antiquissimus and coiled S. spiralis, which suggests that neither were typical astrorhizaceans. [ABSTRACT FROM AUTHOR]