Evidence for seafloor microbial mats and associated metazoan lifestyles in Lower Cambrian phosphorites of Southwest China

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.     

First evidence of Ediacaran–Fortunian elliptical body fossils in the Brioverian series of Brittany,NW France

Body fossils are described for the first time from the Upper Brioverian (Ediacaran–Fortunian) deposits of Central Brittany (NW France). The material consists of a dozen of specimens preserved on two slates, recently collected in a quarry in Saint‐Gonlay. The fossils, centimetric in size, have an elliptical and acuminated outline, with a peripheral bulge but without conspicuous concentric or radial lines. These body fossils are preserved in slates that are locally rich in grazing traces, sometimes associated with microbial mats. The sedimentological characteristics of these deposits correspond to a shallow marine shelf environment, with a low to moderate hydrodynamism. Such findings, in concomitance with previous ichnological discoveries, unravel the potential of the Brioverian strata from Brittany as a new locality to provide answers on Ediacaran and early Cambrian biocenoses.     

AN ORDOVICIAN LOBOPODIAN FROM THE SOOM SHALE LAGERSTÄTTE, SOUTH AFRICA

Abstract:  The first lobopodian known from the Ordovician is described from the Soom Shale Lagerstätte, South Africa. The organism shows features homologous to Palaeozoic marine lobopodians described from the Middle Cambrian Burgess Shale, the Lower Cambrian Chengjiang biota, the Lower Cambrian Sirius Passet Lagerstätte and the Lower Cambrian of the Baltic. The discovery provides a link between marine Cambrian lobopodians and younger forms from the Silurian and Carboniferous. The new fossil preserves an annulated trunk, lobopods with clear annulations, and curved claws. It represents a rare record of a benthic organism from the Soom Shale, and demonstrates intermittent water oxygenation during the deposition of the unit.     

Trace fossils from lower Cambrian Hongjingshao Formation,Yunnan, China: Taxonomy,palaeoecology, palaeoenvironment

The Hongjingshao Formation (Cambrian Series 2, Stage 3) yields abundant, complex trace fossils that have not been systematically investigated up till now. Our detailed ichnological study of the traces from this formation in Malong area, Yunnan, China, recognizes four groups: (1) simple horizontal or sub-horizontal burrows; (2) complex branched burrow systems; (3) arthropod trails and; (4) simple vertical burrows belonging to the Cruziana ichnofacies. A trophic web is reconstructed for the community in Hongjingshao Formation based on both trace and body fossils. The ichnological and sedimentological features of the formation indicate an intertidal setting. The trace fossil assemblage demonstrates that Early Cambrian organisms were able to colonize very shallow marine environments, which further supports the landward expansion of the Early Cambrian ecosystems.     

Trace fossils from arenig flysch sediments of eire and their bearing on the early colonisation of the deep seas

A sequence of Lower Ordovician (Arenig) turbidites in Co. Wexford, Eire, has yielded one of the earliest diverse ichnofaunas yet recorded from deep water sediments comprising: Chondrites, Glockerichnus, Gordia, Helminthopsis, Lorenzinia, Neonereites, Palaeophycus, Paleodictyon, Planolites, Sublorenzinia, Taenidium, Taphrhelminthopsis, Teichichnus and Tomaculum. This ichnofauna is critical in any analysis of the colonisation of the deep seas by trace fossil‐producing animals.

A world‐wide review shows that the earliest trace fossils are mainly from Late Precambrian shelf sea environments, but many more evolved during very rapid diversification in the pre‐trilobite Lower Cambrian.

There was little increase in diversity in shallow water after the Lower Cambrian but a progressive colonisation of the deep ocean took place and this accelerated during the Ordovician, when the main lineages of deep sea trace fossils were established there. Rosetted, patterned, meandering and simple spiral forms evolved in shallow water in the Upper Precambrian and pre‐trilobite Lower Cambrian and only later migrated into the deep sea, whereas complex, closely programmed, spiral traces may have evolved there.     

A twofold role for global energy gradients in marine biodiversity trends

Explanations for major biodiversity patterns have not achieved a consensus, even for the latitudinal diversity gradient (LDG), but most relate to patterns of solar energy influx into Earth systems, and its effects on temperature (as biochemical activity rates are temperature sensitive) and photosynthesis (which drives nearly all of the productivity that fuels ecosystems). Marine systems break some of the confounding correlations among temperature, latitude and biodiversity that typify the terrestrial systems that have dominated theoretical discussions and large‐scale analyses. High marine diversities occur not only in warm shallow seas where productivity may be either low or high, depending on regional features, but also in very cold deep‐sea regions, indicating that diversity is promoted by stability in temperature and in trophic resources (nutrients and food items), and more specifically by their interaction, rather than by high mean values of either variable. The common association of high diversity with stable but low to moderate annual productivity suggests that ecological specialization underlies the similarly high diversities in the shallow tropics and deep sea. Recent work on shallow‐marine bivalves is consistent with this view of decreasing specialization in less stable habitats. Lower diversities in shallow seas are associated with either high thermal seasonality (chiefly in temperate latitudes) or highly seasonal trophic supplies (at any latitude), which exclude species that are adapted to narrow ranges of those variables.     

Palaeoecological significance of calcimicrobial communities during ramp evolution: An example from the Lower Cambrian of Germany

Summary The palaeoecology of calcimicrobial communities from the only Gondwana-related Lower Cambrian in Central Europe (eastern Germany, carbonates and siliciclastics) has been studied. Six morphological groups of calcimicrobes are described. Some of them show a significant ability for sediment stabilization and construction of biohermal reef mounds. Other types of calcimicrobes were more common in biostromal thickets. Some of them were capable of populating different environments, growing in different modes and on different substrates. AnEpiphyton-archaeocyathan reef mound is described, illustrating the importance of calcimicrobes for mound formation. The fossil communities together with a complex of sedimentary features allow a reconstruction of the depositional history of the environment. Based on comparison with similar Gondwanan Lower Cambrian successions (Sardinia/Italy, Spain) and facies development a model is proposed describing the sedimentary history. Three depositional stages are distinguished: (1) deep subtidal ramp, (2) shallow subtidal ramp, (3) shallow subtidal to intertidal mixed siliciclastic-carbonate ramp with migrating oolitic shoals. In spite of similarities with the deposits in southern Europe, some distinct differences exist with respect to the succession of facies, the completeness of the sections, the fossil spectrum, and the nature of the siliciclastic sediments. For the German Lower Cambrian, a facies development from a low energy deep environment to a high energy shallow environment (partly restricted and with some evaporites) can be reconstructed. As compared with Sardinia and Spain, the depositional environment of the eastern German Lower Cambrian successions was predominantly characterized by low-energy conditions.     

Callovian (Middle Jurassic) marine microplankton from southwestern Germany: Biostratigraphy and paleoenvironmental interpretations

Late Middle Jurassic dinoflagellate cyst assemblages are documented from a section in Kandern and in samples from additional localities in southern Germany. The changes in the composition of the marine microfloras through the Callovian sections are related to fluctuations in sea-levels and changes in depositional environments. The Lower Callovian claystones, which represent inner shelf, soft bottom deposits, contain rich, diverse and well preserved dinoflagellate cyst assemblages. In the Kandern section a marked transition from the Lower Callovian claystone facies to the Anceps-oolite occur within the Calloviense zone. This transition was caused by a regression, and coincides with the LZA−3.1/LAZ−3.2 boundary on the cycle chart of Haq et al. (1987). Diverse dinoflagellate cyst assemblages also occur in the Anceps-oolite, but they are not so abundant as the acritarchs which dominate the overall palynofloras. The Anceps-oolite is interpreted as representing an offshore, but shallow marine environment. The transition to the overlying Upper Callovian “Renggeri Clay” is more gradual, but accompanied by marked changes in the overall composition of the marine microfloras. The “Renggeri Clay”, which represent an outer shelf, soft bottom depositional environment, shows a higher species diversity and abundance of dinoflagellate cysts than the underlying sequences. The distribution pattern of selected species through the Kandern section indicates that some species were related to specific environments, i.e. open marine forms, and that others could have been opportunistic forms, tolerating more shallow marine environments.

The distribution of the dinoflagellate cysts from southern Germany is related to the standard Northwest European ammonite zonation. The ranges of selected biostratigraphic key species are compared with ranges previously reported from the British and northwest European Jurassic. There are several discrepancies with respect to the earliest appearance and extinction datums, the only relevant Callovian palynostratigraphic “events” coinciding in southern Germany and Britain being: The extinction of Aldorfia aldorfensis at the top of the Calloviense zone, and the earliest incoming of Belodinium spp. and Liesbergia scarburghensis at the base of the Lamberti zone.     

Occurrence of the Lower Cambrian anemone-style trace fossils in the Zabuk Formation (Mardin–Derik,SE Turkey)

The Zabuk Formation of the Derik Group exposed over much of south-eastern Turkey presents a succession composed of shallow marine and fluvial siliciclastic sedimentary rocks. The Lower Cambrian assemblages containing abundant anemone-style trace fossils are known from most major palaeocontinents such as Laurentia, Baltica, and Gondwana. These have possible affinities with semi-infaunal dwelling anemones on siliciclastic mid-latitude shelves of West Gondwana in early Fortunian deposits. Among them is Bergaueria, characterized by plug-shaped burrows as exemplified by Bergaueria perata, that is, a characteristic trace fossil of the Cambrian globally.     

Early Palaeozoic diversification of chitons (Polyplacophora, Mollusca) based on new data from the Silurian of Gotland, Sweden

Isolated, well-preserved silicified sclerites of an unusually diverse Silurian paleoloricate assemblage from Gotland preserve morphological features that are important in interpreting palaeobiology. The typically granular dorsal ornament is comparable with Recent chitons, and is hence possibly linked functionally with sensory aesthetes. Ventral structures particularly in thickened shells indicate major muscle attachment sites sub-apically or, equivalently, beneath the rim of the apical area, and also marginally. In early chitons such as Cambrian Matthevia, deep ventral cavities represent comparable sub-apical sites. Three Gotland genera with an unusual, holoperipheral shell growth style apparently represent plated aplacophorans (cf. Acaenoplax), which coexisted with paleoloricate chitons in shallow inshore carbonate shelf environments. Sclerite features of all the Gotland genera are discussed together since they share most characteristics. The new family Heloplacidae includes those genera and Acaenoplax, which in a preliminary cladistic analysis form a sister group to other Lower Palaeozoic paleoloricates. Multiplated skeletons in paleoloricates and this group of aplacophorans represent parallel evolution of dorsal armour, which in chitons resulted in overlapping, articulating sclerites. The diversity of the mid-Silurian Gotland assemblage is examined against early evolutionary diversification of polyplacophorans, aplacophorans, and in relation to the overall record of Palaeozoic paleoloricate and neoloricate chitons. Peaks in diversity in early Ordovician, mid-Silurian and early Carboniferous times correspond to periods with widespread development of low latitude carbonate shelves. Neoloricates, apparently with an additional shell layer that contributed articulatory plates, appeared in the Devonian from where the fossil record remains poorly known.     

Nonmarine aspects of the Cambrian Tonto Group of the Grand Canyon, USA, and broader implications

A re-examination of the classic time-transgressive model of the Cambrian Tonto Group of the Grand Canyon challenges some of the basic assumptions regarding strandline migration during the Sauk transgression, the applicability of sequence stratigraphy to epicratonic deposits, and the criteria by which siliciclastic deposits of the North American Cambrian system are categorized as marine. Indicators for high short-term depositional rates, coupled with the complex splay and coalescence of meter-scale parasequence architectures, detract from the shoreline migration and deepening seas model that has held sway for over 60 years. Ostensibly marine signatures, represented by sparse and fragmentary trilobite fossils and abundant bedding plane trace fossils, are closely associated with nonmarine signatures, represented by nonmarine palynoflora and clay mineral assemblages, and highly oxidized paleoweathering profiles. The vast extent of three-dimensional exposure of the Cambrian section in the Grand Canyon shows a paleogeography not of a single discrete shoreline translated laterally with rising and falling sea-level, but of a single mosaic setting of emergent shoals and rocky peaks, broad fluvial and tidal channels, and shallow pools and interchannel flats. Much detail concerning sediment dynamics and paleoenvironment is lost or overlooked when summary terms like “marginal marine” or “nearshore” are applied to such settings. It is suggested here that the Tonto Group, with its mixed marine and nonmarine attributes, and considering its historic role as a teaching model, be viewed not as an exceptional Cambrian deposit, but as an exemplar of epicratonic deposits on a pre-vegetated landscape.     

A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Morphology,ontogeny, and heterochrony in lower and middle Cambrian Gogiids (Eocrinoidea,Echinodermata) from Guizhou Province,China

Gogiid eocrinoids from the Lower Cambrian (Balang Formation — Guizhoueocrinus) and basal Middle Cambrian (Kaili Formation — Sinoeocrinus, Globoeocrinus) in Guizhou Province, China are found in great numbers and are extremely well preserved as high fidelity molds in shale/mud sized siliciclastics. Because of their numbers, complete ontogenetic growth sequences have been observed. Significant differences in growth patterns (heterochrony) are present between Lower and Middle Cambrian genera: thecal plates tend to be paedomorphic in development, while the sutural pores developed between them are peramorphic in their development. Because of the large surface area of theca and brachioles relative to the size of the attachment area, considerable drag in ambient currents would necessitate a strong attachment medium. Anchoring by “biogluing”, possibly by collagen, directly to the substrate or to biodetritus is proposed as the method of attachment.     

New arthropods from the Lower Devonian Hunsru¨ck Slate (Lower Emsian, Rhenish Massif, western Germany)

Four new genera and species of arthropod, Cambronatus brasseli Wingertshellicus backesi Eschenbachiellus wuttkensis and Magnoculus blindi , are described from the Hunsru¨ck Slate (Lower Emsian) of Germany. All four occur in the Wingertshell Member in the vicinity of Bundenbach. They preserve remarkable details of the ventral morphology, including the appendages, as a result of pyritization. In each case the body consists of just two tagmata, a cephalon and a large number of similar trunk somites. Both Cambronatus Wingertshellicus have fluke-like appendages making up a tail fan. The affinities of these arthropods lie with the Crustacea, apart from Magnoculus , which is an arachnomorph, but they do not fall within those clades with modern representatives. They show that morphologies other than those represented by trilobites and modern arthropod groups persisted long after the Cambrian, at least in muddy bottom marine settings.     

Lebensspuren von Meersauriern und ihren Beutetieren im mittleren Jura (Callovien) von Liesberg, Schweiz

Summary Numerous gutter-like furrows, up to 60 cm wide and up to 9 m long are preserved at the interface “Macrocephalus Beds”/“Callovian Marl” over a surface of 20 by 200 m. They are interpreted as feeding traces made by large marine vertebrates, most likely plesiosaurs and ichthyosaurs searching for food in the lime mud of the shallow Middle Jurassic sea floor. Possible prey animals were infaunal invertebrates (crustaceans) which produced an intricate meshwork of burrows (mainlyRhizocorallium irregulare andThalassinoides) in the bottom sediments, as well as infaunal bivalves. Evidence from cololites of predatory pelagic reptiles (ichthyosaurs, plesiosaurs) as well as reptile regurgitalites indicate that these animals fed not only on fast-swimming vertebrates and cephalopods but also on epi- and endobenthic invertebrates. In addition, the cololites show that the predators ingested considerable amounts of bottom sediment. Different sizes and shapes of the traces suggest that the gutters were produced by different reptiles or age groups. Candidates for the widest gutters are pliosaurs. Of the marine vertebrates known from Jurassic time, only the snout of adult pliosaurs of the genusLiopleurodon was broad enough to produce gutters more than 40 cm wide. Smaller, less than 15 cm wide gutters, could have been made by plesiosauroids or by the narrow pointed snouts of ichthyosaurs. Almost identical traces described from the Oxfordian of Spain and similar but smaller traces from the Lower Devonian of Prague are equally interpreted as feeding traces on the sea floor. Feeding traces of vertebrates in bottom sediments may give detailed information on the hunting behaviour of the predators. However, the attribution of the traces to definite vertebrate taxa remains uncertain.

     

A trace fossil assemblage from fluvial Old Red deposits (Wood Bay Formation; Lower to Middle Devonian) of NW-Spitsbergen, Svalbard

From the fluvial Old Red Sandstone (ORS) of the Lower to Middle Devonian Wood Bay Formation (NW-Spitsbergen), a diverse trace fossil assemblage, including two new ichnotaxa, is described: Svalbardichnus trilobus igen. n., isp. n. is interpreted as the three-lobed resting trace of an early phyllocarid crustacean (Rhinocarididae). Cruziana polaris isp. n. yields morphological details that point towards a trilobite origin. This occurence of presumably marine trace makers in a fluvial red bed sequence raises the question of whether we are dealing with marine ingressions that are not sedimentologically expressed, with homeomorphy, or with an adaptation of marine groups to non-marine environments.     

Dinosaurier aus Nordostdeutschland

Remains of dinosaurs in NE Germany The former clay pit of Grimmen (NE Germany) represents a glacially dislocated outcrop of upper Lower Jurassic shallow‐marine strata. Apart from typical marine organisms, the Lower Jurassic of Grimmen has also yielded the remains of the basal thyreophoran dinosaur Emausaurus ernsti. E. ernsti was a comparatively small dinosaur, generally characterized by a dermal armor composed of small isolated scutes. Recently, the remains of a second dinosaur belonging to the sauropod clade have been identified. The specimen represents the second Early Jurassic basal sauropod from Germany ever found. The sauropod was probably washed into the marine realm from nearby coastal land after its death, whereas the compacted pile of skeletal elements of E. ernsti found in a concretion is suspected to be a Speiballen produced by a large‐scaled marine reptile.     

Isopodichnus in a trace fossil assemblage from the Old Red Sandstone

Isopodichnus stromnessi sp. nov. from the Lower Stromness Flags of Orkney is described in detail, and the range of variation in the traces ascribed to different habits of different sized individuals producing the traces. Activity diagrams illustrate scme of the burrowing habits of the organism. Isopodichnus is typical of non-marine strata, particularly those of shallow impermanent lacustrine environments, and can be distinguished from Cruziana on the basis of the shape of the trace ending. AsSociated burrows are briefly described and indicate the presence of a hitherto undescribed invertebrate fauna associated with the Orcadian flagstone sequence of the Middle Old Red Sandstone.     

Expression patterns of hedgehog, wingless, and decapentaplegic during gut formation of Gryllus bimaculatus (cricket).

We observed expression patterns of hedgehog (hh), wingless (wg), and decapentaplegic (dpp) during gut development of Gryllus bimaculatus (the cricket), a typical hemimetabolous insect, and compared with those observed in Drosophila, a typical holometabolous insect. Gryllus hh(Gbhh) and Gbwg are expressed in both foregut and hindgut, while Gbdpp is expressed only in the hindgut: at the boundaries between the small and large intestine, and between the large intestine and rectum. Although the expression patterns of Gbhh and Gbwg are essentially comparable to those observed in Drosophila, the expression pattern of Gbdpp differs from those of the Drosophila dpp.     

Trace fossil analysis of lacustrine facies and basins

Two ichnofacies are typical of lacustrine depositional systems. The Scoyenia ichnofacies characterizes transitional terrestrial/nonmarine aquatic substrates, periodically inundated or desiccated, and therefore is commonly present in lake margin facies. The Mermia ichnofacies is associated with well oxygenated, permanent subaqueous, fine-grained substrates of hydrologically open, perennial lakes. Bathymetric zonations within the Mermia ichnofacies are complicated by the wide variability of lacustrine systems. Detected proximal–distal trends are useful within particular lake basins, but commonly difficult to extrapolate to other lakes. Other potential ichnofacies include the typically marine Skolithos ichnofacies for high-energy zones of lakes and substrate-controlled, still unnamed ichnofacies, associated to lake margin deposits. Trace fossils are useful for sedimentologic analysis of event beds. Lacustrine turbidites are characterized by low-diversity suites, reflecting colonization by opportunistic organisms after the turbidite event. Underflow current beds record animal activity contemporaneous with nearly continuous sedimentation. Ichnologic studies may also help to distinguish between marine and lacustrine turbidites. Deep-marine turbidites host the Nereites ichnofacies that consists of high diversity of ornate grazing traces and graphoglyptids, recording highly specialized feeding strategies developed to solve the problem of the scarcity of food in the deep sea. Deep lacustrine environments contain the Mermia ichnofacies, which is dominated by unspecialized grazing and feeding traces probably related to the abundance and accessibility of food in lacustrine systems. The lower diversity of lacustrine ichnofaunas in comparison with deep-sea assemblages more likely reflects lower species diversity as a consequence of less stable conditions. Increase of depth and extent of bioturbation through geologic time produced a clear signature in the ichnofabric record of lacustrine facies. Paleozoic lacustrine ichnofaunas are typically dominated by surface trails with little associated bioturbation. During the Mesozoic, bioturbation depth was higher in lake margin facies than in fully lacustrine deposits. While significant degrees of bioturbation were attained in lake margin facies during the Triassic, major biogenic disruption of primary bedding in subaqueous lacustrine deposits did not occur until the Cretaceous.