Ontogeny of the redlichiid trilobite Eoredlichia intermediata from the Chengjiang Lagerstätte,lower Cambrian,southwest China

A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.     

The Sirius Passet Lagerstätte: silica death masking opens the window on the earliest matground community of the Cambrian explosion

The Sirius Passet Lagerstätte (SP), Peary Land, North Greenland, occurs in black slates deposited at or just below storm wave base. It represents the earliest Cambrian microbial mat community with exceptional preservation, predating the Burgess Shale by 10 million years. Trilobites from the SP are preserved as complete, three‐dimensional, concave hyporelief external moulds and convex epirelief casts. External moulds are shown to consist of a thin veneer of authigenic silica. The casts are formed from silicified cyanobacterial mat material. Silicification in both cases occurred shortly after death within benthic cyanobacterial mats. Pore waters were alkali, silica‐saturated, high in ferric iron but low in oxygen and sulphate. Excess silica was likely derived from remobilized biogenic silica. The remarkable siliceous death mask preservation opens a new window on the environment and location of the Cambrian Explosion. This window closed with the appearance of abundant mat grazers later as the Cambrian Explosion intensified.     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

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.     

An exceptional record of Cambrian trilobite moulting behaviour preserved in the Emu Bay Shale,South Australia

Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock 1964 being extremely abundant, and the larger Redlichia takooensis Lu 1950 , 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.     

High‐resolution δ13Corg chemostratigraphy links the Decorah impact structure and Winneshiek Konservat‐Lagerstätte to the Darriwilian (Middle Ordovician) global peak influx of meteorites

The precise age of the Winneshiek Shale, a recently discovered Konservat‐Lagerstätte located in a very unusual depositional setting inside the Decorah impact structure, has remained uncertain in the absence of biostratigraphically highly diagnostic fossils. This chemostratigraphical study, based on δ¹³C_org data from 36 drill core samples through the shale, shows that the age ranges from the upper part of a small unnamed δ¹³C excursion in the Dw1 Stage Slice of the Darriwilian Global Stage to the lower part of the MDICE excursion in Stage Slice Dw2 of the same stage. This Dw1–Dw2 interval has an isotopic age of ~464–467 Ma. The gradational contact between the Winneshiek Shale and the underlying, rapidly deposited, impact breccia indicates minimal time difference between the impact event and the Winneshiek Shale. New age data show that the Decorah impact event was coeval with the early Darriwilian abnormally high influx of micrometeorites and meteorites recorded in sections in Baltoscandia, Russia and China and that the Decorah crater can be included among the unusually large number of meteorite craters formed during Middle and early Late Ordovician time. As is commonly the case in black shale deposits, the partly uniquely preserved Winneshiek Shale crater fauna is impoverished taxonomically and adds numerically relatively little to the conspicuous and much discussed Darriwilian global biodiversification increase.     

The bivalved arthropods Isoxys and Tuzoia with soft‐part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island,Australia)

Abstract: Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft‐bodied features previously unknown from this Burgess Shale‐type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better‐known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.     

A rare non‐trilobite artiopodan from the Guzhangian (Cambrian Series 3) Weeks Formation Konservat‐Lagerstätte in Utah,USA

We describe a weakly biomineralized non‐trilobite artiopodan arthropod from the Guzhangian Weeks Formation of Utah. Falcatamacaris bellua gen. et sp. nov. is typified by a thin calcitic cuticle, broad cephalon without eyes or dorsal ecdysial sutures, an elongate trunk with distinctively sickle‐shaped pleural spines and a long tailspine with a bifurcate termination. The precise affinities of Falcatamacaris gen. nov. are problematic due to the presence of unique features within Artiopoda, such as the peculiar morphology of the pleural and posterior regions of the trunk. Possible affinities with aglaspidid‐like arthropods and concilitergans are discussed based on the possession of 11 trunk tergites, edge‐to‐edge articulations and overall body spinosity. The new taxon highlights the importance of the Weeks Formation Konservat‐Lagerstätte for further understanding the diversity of extinct arthropod groups in the upper Cambrian.     

Plywood‐like shell microstructures in hyoliths from the middle Cambrian (Drumian) Gowers Formation,Georgina Basin,Australia

Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators.     

Multi‐jawed chaetognaths from the Chengjiang Lagerstätte (Cambrian,Series 2, Stage 3) of Yunnan,China

Chaetognaths (arrow‐worms) are enigmatic in terms of their phylogenetic position, while the existence of Protosagitta spinosa from the Chengjiang Lagerstätte suggests minimal change in their unique bodyplan since at least the early Cambrian. Apart from rare (and sometimes controversial) soft‐bodied remains, the fossil record of chaetognaths is otherwise almost entirely dependent on early Palaeozoic phosphatic microfossils, some of which are placed amongst so‐called protoconodonts. Fused spine clusters are strikingly similar to the cephalic grasping apparatus of extant forms and are assumed to have had a comparable configuration. Here we report a new chaetognath, Ankalodous sericus gen. et sp. nov., coeval with Protosagitta but with a complex feeding apparatus consisting of multiple bundles of recurved spines whose principal function appears to have been grasping. Like all other chaetognaths a predatory mode of life is likely, but its position relative to the sediment–water interface is less certain. Reduction of the feeding apparatus, from the multi‐jawed arrangement of A. sericus to the grasping spines and associated smaller teeth seen in other chaetognaths, was probably a subsequent development and conceivably was linked with a shift to a pelagic mode of life. We also report a new specimen of Protosagitta. This confirms earlier observations but it possesses hitherto unrecognized features, including a cephalic tentacle and fin rays.     

Feeding behaviour of a new worm (Priapulida) from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia)

Singuuriqia simoni gen. et sp. nov. represents the first record of a priapulid worm from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia). It is defined by an unusually broad, longitudinally folded, foregut which tapers through the pharynx towards the anterior mouth; posteriorly, the same longitudinal folding is evident in the narrow gut. The slender, smooth, trunk in the unique specimen passes anteriorly into an oval proboscis which culminates in a smooth, extensible, pharynx with pharyngeal teeth. The capacity for substantial expansion of the foregut permitted rapid ingestion of food prior to digestion at leisure. Cololites suggest both carnivorous and deposit feeding behaviour, indicating that Singuuriqia, like the present day Priapulus, was probably omnivorous.     

The La Voulte Lagerstätte (Callovian): Evidence for a deep water setting from sponge and crinoid communities

The siliceous sponges and crinoids from the Chénier Ravine (France, Lower Callovian) are used here as biological markers to characterize the palaeoenvironment of the adjacent and contemporaneous La Voulte Lagerstätte that is remarkable for its unique soft-bodied fauna (e.g. worms, cirrate octopods, vampire squids). Sponges are abundant with dominant hexactinellids (80%) and lithistids (20%). Four lines of fossil evidence, supported by Recent analogues, indicate that this sponge community inhabited deep-water setting under dysphotic or aphotic conditions: (1) the dominance of hexactinellids; (2) the prevalence of cone-shaped and erect morphologies that usually characterize Recent bathyal sponges; (3) the close similarities with Recent hexactinellids from the continental slope and; (4) the lack of encrustation by photophilic organisms. Attachments of sponges on hard substrate (e.g. crystalline basement) and their preservation in soft sediments (e.g. muds) indicate heterogeneous bottom conditions. The Chénier fauna also contains small stalked, asymmetric cyrtocrinid crinoids that are known to live on hard substrates in bathyal environments such as the SW Pacific steep seamounts. Convergent palaeoenvironmental clues obtained from both crinoids and siliceous sponges support the notion that the La Voulte area, including the La Voulte Lagerstätte, was situated in the upper part of the bathyal zone near the slope-basin transition with a water depth most probably exceeding 200 m. Supporting evidence from heterogeneous substrates and complex fault systems indicate a depositional environment along the external part of the slope where steep topographies and blocks usually favour the settlement of cyrtocrinid crinoids and hexactinellid sponges. La Voulte may therefore be one of the rare and extremely precious Mesozoic Lagerstätten to have fossilized a deep marine environment.     

Lower Cambrian (Botomian) polycystine Radiolaria from the Altai Mountains (southern Siberia, Russia)

Relatively well-preserved polycystine Radiolaria are here described from Lower Cambrian (Botomian) strata of the Shashkunar Formation, Altai Mountains in southern Siberia (Russia). These radiolarians display a test formed of a disorderly and three-dimensionally interwoven meshwork of numerous straight and curved bars branching from a five-rayed point-centered spicule located within the inner shell surface. The shell structure allows their assignment to the family Archeoentactiniidae, thus extending the known age range of the family down to the Lower Cambrian. The Botomian age is based essentially on trilobites (Parapagetia-Serrodiscus zone), but also on archaeocyathids identified in earlier publications. The study of the radiolarian-bearing sedimentary sequence confirms the presence of polycystine radiolaria in the external platform environments of Lower Cambrian ecosystems.     

Soft–part preservation in heteromorph ammonites from the Cenomanian–Turonian Boundary Event (OAE 2) in north–west Germany

Abstract: From thinly laminated marlstones of the Hesseltal Formation, representing the Late Cenomanian Oceanic Anoxic Event (OAE) 2, at Lengerich in the Teutoburger Wald (Westfalen, north‐west Germany), 17 sediment‐compacted baculitid ammonites with carbonised and partially phosphatised soft parts are recorded. Some preserve remains of the buccal mass, including jaws (occasionally articulated) and radulae, as well as of the cephalic cartilage, such as eye capsules. Such have not yet been recorded previously for the order Ammonoidea. In addition, originally unmineralised parts found preserved in these specimens include extensive portions of the digestive tract, the siphonal tube, false colour patterns (megastriae), as well as traces of what would appear to be the oviduct. At the same levels, patches with numerous isolated horny upper and rarer lower jaws as well as radulae occur; these may represent regurgitates or faeces of larger predators. The cephalopod remains described were deposited in an epicontinental setting, possibly at palaeodepths between 200 and 600 m. In this particular Late Cretaceous fossil Lagerstätte, upper jaws and anaptychi of ammonites rank among the commonest fossils.     

Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte,lower Cambrian of North Greenland

Stein, M., Peel, J.S., Siveter, D.J. & Williams, M. 2009: Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte, lower Cambrian of North Greenland. Lethaia, Vol. 43, pp. 258–265. Isoxys volucris is the most commonly occurring species in the lower Cambrian Sirius Passet Lagerstätte of North Greenland. Newly identified material allows a first, limited, account of the ventral morphology of this species, hitherto known only by the morphology of its shield. The antennula is large and robust, composed of about seven articles armed with spines, and was probably not sensorial. The postantennular limbs are serially similar, biramous with a large paddle‐shaped exopod fringed with setae. It is possible that the animal possessed a furca. The inner lamella, lining the ventral surface of the shield is recognised in Isoxys for the first time. Comparisons with other congeneric species of which aspects of the ventral morphology are known, show similarities with Isoxys auritus from China, reconsidered here, but indicate differences in antennular morphology with other species as currently understood. □Cambrian, Greenland, Isoxys, soft anatomy, Sirius Passet, palaeoecology.     

Taphonomy and palaeoecology of Late Triassic (Carnian) ammonoid concentrations from the Taurus Mountains,Turkey

The deposits of the Carnian Kas?mlar Formation within the Taurus Platform Units of south‐western Turkey represent an important archive of a Late Triassic ecosystem. New palaeontological information was obtained by analysing the Kasimlarceltites mass occurrence, located within the Kas?mlar Formation and named after the Lower Carnian (Julian) ammonoid genus Kasimlarceltites. This is the dominant taxon (> 94%) within the mass occurrence: nearly 775 million ammonoids and 50 million gastropods were extrapolated for the whole extension (at least 5 km²) of the Kasimlarceltites beds. This calculation is one of the main findings within this study, as it is the first time that such a fossil mass occurrence was quantified. Additionally, orientation measurements of the planispiral ammonoids and the helical gastropods enabled reconstructing the history of the mass occurrence and interpreting the underlying transport mechanisms. Further taphonomic aspects (e.g. biofabric, preservation, bioerosion or genetic classification) as well as comparisons with samples of the same acme zone from different localities near A?a?iyaylabel (AS IV, KA I‐II) point to a two‐phased genetic history. Accordingly, local mass mortality within the Kasimlarceltites fauna due to oxygen fluctuations or methane degassing may have initially led to a primary accumulation. These deposits were then reworked and redeposited basinward by gravity flows to create the present‐day secondary allochthonous concentrations.     

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.     

Anatomy and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil Lagerstätte (lower Cambrian; Southwest China)

An updated reconstruction of the body plan, functional anatomy and life attitude of the bradoriid arthropod Kunmingella is proposed, based on new fossil specimens with preserved soft parts found in the lower Cambrian of Chengjiang and Haikou (Yunnan, SW China) and on previous evidence. The animal has a single pair of short antennae pointing towards the front (a setal pattern indicates a possible sensory function). The following set of seven appendages (each composed of a 5-segmented endopod and a leaf-like exopod fringed with setae) is poorly differentiated, except the first three pairs (with possible rake-like endopodial outgrowths, smaller exopods) and the last pair of appendages (endopod with longer and more slender podomeres). The endopods are interpreted as walking legs with a possible role in handling food particles (marginal outgrowth with setae). The leaf-like exopods may have had a respiratory function. The trunk end is short, pointed, flanked with furcal-like rami and projects beyond the posterior margin of the carapace. The attachment of the body to the exoskeleton is probably cephalic and apparently lacks any well-developed adductor muscle system. The inferred life attitude of Kunmingella (e.g. crawling on the surface of the sediment) was that of a dorsoventrally flattened arthropod capped by a folded dorsal shield (ventral gape at least 120°), thus resembling the living ostracode Manawa. The animal was also probably able to close its carapace as a response to environmental stress or to survive unfavourable conditions (e.g. buried in sediment). The anterior lobes of the valves are likely to have accommodated visual organs (possibly lensless receptors perceiving ambient light through the translucent head shield). Preserved eggs or embryos suggest a possible ventral brood care. The presence of Kunmingella in coprolites and its numerical abundance in Chengjiang sediment indicate that bradoriids constituted an important source of food for larger predators. Kunmingella differs markedly from the representatives of the crown group Crustacea (extant and Cambrian taxa) and from the stem group derivatives of Crustacea (exemplified by phosphatocopids and some ‘Orsten’ taxa) in showing no major sign of limb specialization (e.g. related to feeding strategies). Although it resembles other Chengjiang euarthropods in important aspects of its body plan (e.g. uniramous antennae, endopod/exopod configuration), Kunmingella possesses several features (e.g. antennal morphology, post-antennular appendages with 5-segmented endopods) which support the view that bradoriids may be very early derivatives of the stem line Crustacea.     

Reappraisal of the Eocene whiptail stingrays (Myliobatiformes,Dasyatidae) of the Bolca Lagerstätte,Italy

The Eocene whiptail stingrays of the family Dasyatidae from the Bolca Lagerstätte, NE Italy, are revised herein in detail. The analysis of the anatomical and morphometric features allows us to identify the species “Dasyatis” zigni (Molin, 1861) as a junior synonym of “D.” muricatus (Volta, 1796), and to assign it to the new genus Tethytrygon gen. n. This new taxon exhibits a unique combination of features (e.g., rhombic disc wider than long, elongated tail folds fail to reach the tip of the tail, thorns absent, single serrated tail sting, “caniniform” teeth on upper jaw, tooth crown ornamentation absent, 175–179 vertebrae, 108–117 pectoral radials, 24–27 pelvic radials and other features of clasper anatomy) that clearly support its attribution to the subfamily Neotrygoninae of the stingray family Dasyatidae. The morphological and phylogenetic affinities of Tethytrygon gen. n. with the living neotrygonines (Neotrygon and Taeniura) suggest a close association of this taxon with the tropical shallow‐water habitats hypothesized for the Bolca palaeoenvironment during the early Eocene. Moreover, the analysis of the fossil occurrences of the neotrygonines provides new insights into the role of the Tethys for the origin and evolutionary history of certain whiptail stingrays.