Precambrian trace fossils and the rise of bilaterian animals

In this attempt to synthesize present knowledge into a coherent story, the Cambrian explosion is interpreted to represent a true adaptive radiation, an event similar to Phanerozoic radiation events in principle but unique in its possibilities. A model of bilaterian evolution helps explain how this particular event involved the sudden initiation of major coelomate phyla. In many of these groups, preservable skeletons are part of the basic body plan. Biochemical‐sequence analyses indicate that acoelomates and pseudocoelomates branched off long before the coelomate radiation. The great differences between Vendian and Cambrian ichnofaunas, therefore, may be the result of a major shift in composition from Vendian acoelomate‐pseudocoelomate‐procoelomate faunas to Phanerozoic faunas dominated by new coelomate groups.     

Dictyodora and associated trace fossils from the Palaeozoic of Thuringia

Dictyodora occurs in the Hauptquarzit (Late Ordovician; D. zimmermanni) and Bordenschiefer (Early Carboniferous; D. liebeana) of Thuringia, East Germany. It is absent in the Early Devonian Nereitenquarzit, and analysis of the trace fossil assemblages points to environmental partitioning within the ‘deep-sea’Nereites Association. The Carboniferous Dictyodora was much larger than the Early Palaeozoic forms and had a long respiratory (?) wall organ. This may have been an adaptation to feeding deeper in anoxic sediments, and the animal developed large ‘parapodia’ to effect its progression through the sediment.     

Diverse tiering patterns in Paleogene flysch trace fossils,Magura nappe,Carpathian Mountains,Poland

Diversified tiering patterns of trace fossils were observed in three Paleogene lithostratigraphic units of the Magura nappe: the Szczawnica Formation (Paleocene‐early Eocene); the Belove?a beds (middle Eocene); and the Piwniczna Sandstone Member of the Magura Formation (middle‐upper Eocene). Sediments of the first unit were deposited under oxygen‐poor conditions, and those of the second and third units under well‐aerated conditions. Thick‐bedded turbidites and fluxoturbidites of the third unit accumulated quickly, in contrast to ‘normal flysch’ of the first and second units. Analyses of tiering patterns show a thickening of tiers following the improvement of sediment oxygenation. Slowly accumulated deposits of ‘normal’ flysch exhibit a more mature, well‐developed tiering pattern than those which accumulated quickly.     

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.     

Environmental distribution of trace fossils in the Jurassic of Kachchh (Western India)

Summary Trace fossils occur abundantly in Middle Jurassic rocks of the Kachchh Basin. They are found in environments ranging from beach sequences down to central parts of the basin. For stratinomic reasons, they are particularly well preserved in storm deposits. Their distribution pattern exhibits a clear relationship to the hydrodynamic conditions and, secondarily, to bathymetry, and follows the classic ichnofacies concept ofSeilacher (1967). High energy nearshore areas and submarine shoals are represented by members of the Skolithos ichnofacies such asOphiomorpha nodosa, Arenicolites, Diplocraterion parallelum, andRhizocorallium jenense. The storm-influenced ramp contains both members of the Cruziana ichnofacies (e.g.Rhizocorallium irregulare, Thalassinoides suevicus, Taenidium serpentinum, Chondrites) and the Skolithos ichnofacies (in particularOphiomorpha). The former were produced during interstorm phases, the latter are of post-storm origin. Carbonate ramp environments of low to intermediate energy also contain members of the Cruziana ichnofacies, whilst equivalent siliciclastic environments are characterized by a low-diversity Zoophycos ichnofacies. Low energy basinal environments of fine-grained substrates contain an impoverished Cruziana ichnofacies consisting ofChondrites, Trichichnus andThalassinoides suevicus. 32 ichnotaxa are briefly described, among themSphaerichnus lobatus ichnogen. et ichnosp. nov.     

Trace fossils from the Podhale Flysch Basin, Poland - an example of ecologically-based lithocorrelation

Certain ichnogenera from the Podhale Flysch Basin do not occur randomly, but are grouped in distinct horizons, which are useful in mapping. The position of certain tuffite beds proves an approximate isochronism of those trace fossil horizons. Changes of the life conditions, among them especially oxygenation of the sea bottom, were responsible for that ichnologic differentiation. Profuse Occurrences of Taphrhelminthopsis are mainly connected with the presence of lighter, bluish shales, which indicates a well-oxygenated bottom. More distal and frequent sedimentation of turbidites could positively influence the preser-vational potential of the surface forms (very good state of preservation of Taphrhelminthopsis suggests that preservation of surface forms was frequent in the flysch basin), but the O₂ condition was the main controlling factor for the occurrences of ichnofauna. Most discriminating were trace makers of pre-depositional (mud-dwelling) Taphrhelminthopsis , and they were most probably irregular echinoids, while soft-bodied trace makers of pre- and post-depositional Planolites (post-depositional means sand-dwellers) were the most tolerant ones. An increased level of biological activity connected with a shift in the OMZ could have been the reason why Taphrhelminthopsis became so profuse in one period. Concerning preservational problems, some complications in determining pre- and post-depositional forms in the case of composite beds are presented.     

Gonad pouches in a new clam-like monoplacophoran from the Silurian of Sweden

Beyrichidiscus fecundatus n. gen. n. sp. is described from the Upper Silurian of Gotland, Sweden. The minute, isostrophically coiled mollusc exhibits profound allometry and is considered to have been a monoplacophoran on the basis of functional interpretation of apertural structures. A clam-like, semi-infaunal mode of life is suggested. A pair of prominent lateral swellings in the adult shell are interpreted as reflecting the location of the gonads.     

Palaeozoic calcareous plankton: evidence from the Silurian of Gotland

In order to investigate whether or not equivalents to modern calcareous plankton existed in Palaeozoic times, extremely well-preserved successions have to be investigated. The Silurian strata on Gotland (Sweden) are exceptionally well preserved because they have not experienced deep burial conditions and tectonic stress, due to their position on the stable Baltic Shield. Scanning electron microscope investigations of polished, slightly etched rock surfaces revealed the presence of a variety of calcareous micro- and nannofossils. Among these organisms, many can be termed 'calcispheres' (60–100 µm in diameter), whereas others due to their size range (nannofossils) are informally termed herein as 'nannospheres' (10–25 µm in diameter). The systematic attribution of these fossils is unknown ( incertae sedis ). Mesozoic calcispheres are usually attributed to calcareous cysts of dinoflagellates following comparisons of ultrastructure with modern species. The abundance of different calcispheres in the Silurian sediments of Gotland and the observation that many of the calcareous microfossils occur in distinctly different facies as well as their spherical shape indicate that they probably belong to calcareous micro- and nannoplankton. We therefore conclude that calcareous plankton most probably existed already during the Palaeozoic, but can only be observed under conditions of exceptional preservation.     

New species of stromatoporoidea from the Upper Ordovician and Silurian of the Subpolar Urals

Stromatoporoidea are widespread in the Upper Ordovician and Silurian beds of the Kozhim River (western slope of the Subpolar Urals). Five new species Cystostroma prodigiosum sp. nov., Stylostroma flabellatum sp. nov., Labechiina arguta sp. nov., Ecclimadictyon faveolatum sp. nov., and Araneosustroma astroplexum sp. nov. are described.     

Pigmentation of the Early Silurian shallow marine red beds in South China as exemplified by the Rongxi Formation of Xiushan,southeastern Chongqing,central China

The origin and pigmentation of red beds have long been investigated, but mainly on continental settings and carbonate facies (including deep marine red beds). This paper focuses on the shallow marine clastic red beds developed in South China during the Silurian. Based on the sedimentological and geochemical analyses on red layers, green layers and laminae of the Rongxi Formation (Llandovery, Lower Silurian) of Xiushan, southeastern Chongqing, known as the ‘lower red beds’ (LRBs), the following observations are made: (1) the LRBs are primary red beds whereas the green layers are secondary (i.e., formed during early diagenesis); (2) the color differentiation of sediments shows little relevance to the bottom redox potential shift but has a close relationship to grain size and sediment sorting which may be caused by sea level changes and tides. The controlling factors of LRBs pigmentation are significantly different from those of deep marine red beds, and some coeval red beds in North America and Europe. It is noted that, among the shallow marine clastic red beds, the sedimentological and geochemical signatures from the laminae seem to better indicate the environmental conditions than those derived from the layers.     

The appearance of Baragwanathia (Lycophytina) in the Silurian

The evidence mainly involving graptolites is restated which supports an Upper Silurian (Ludlow) dating for a vascular flora containing Baragwanathia (Lycophytina) in Victoria, Australia.     

Heterochrony in the Silurian radiation of encrinurine trilobites

Divergence of Silurian encrinurine trilobite clades from common ancestry may be modelled as hetero-chronic pattern. Comparing ontogenies of ancestral late Ordovician Encrinuroides and the descendant punctarus and uariolaris plexi of Silurian Encrinurus provides a test of this hypothesis. Heterochrony in the punctarus plexus is dissociated with respect to entire organisms, but regionally global throughout cranidia (paedomorphic states) and pygidia (peramorphic states). Regulatory dissociation may explain the apomorphous enrollment strategy of this group. The uariolaris plexus, for which a growth series of Balizoma dimitroui (Perry & Chatterton) provides ontogenetic data, shows a complex of peramorphic cephalic character states. Changes in developmental timing thus provide a mechanism by which morphologic and ecologic divergence of closely-related clades was catalyzed in the Llandovery encrinurine radiation. Paedomorphic and dissociated Lineages of the odontopleurid Leonaspis provide a comparable example of divergent heterochronic pattern associated with coexisting (in the same biofacies) species of comparatively recent common ancestry.     

Radial perforations in crinoid stems from the Silurian of Gotland

Radiating intracolumnal canals are a characteristic feature of large (diameter 10 mm or more) crinoid stems from the Silurian of Gotland. They are found in nodals as well as in internodals where the columnal height exceeds one millimetre. They were formed secondarily in the median and distal portions of crinoid stems with pseudocirriferous holdfasts. Intercolumnal canals are found in the distal parts of stems with true cirri regardless of the size of the stem. It is suggested that these canals played an important role in crinoid physiology. The crinoids are believed to have sustained a large proportion of their tissues through cutaneous digestion and uptake of dissolved substances from the surrounding sea water. The intra- and intercolumnal canals increased the surface of the axial canal in relation to volume. They provided a connection between the axial canal and the surrounding sea water, thus facilitating nutrient transport to the tissues.     

A new type of Precambrian megascopic fossils: the Jinxian biota from northeastern China

Precambrian fossils are crucial for our understanding of the evolution of early organisms. Megascopic body fossils are more important because they potentially represent macroorganisms. However, the Precambrian fossil record is sparse and dominated by microfossils and microbial structures. Here we show a new type of megascopic fossils recovered from the Xingmincun Formation (probably Neoproterozoic age), northeastern China. The specimens are flat, flexible (easily corrugated) and discoidal in outline. Concentric or spiral ridges are preserved on both sides. Petrographical thin section examination indicates that the specimen consist of a thin layer of microcrystalline quartz grains (about 20–30 μm thick) wrapped by an outer sheath, composed primarily of chlorites. Field Emission Scanning Electron Microscopy (FE-SEM) coupled with an x-ray energy dispersive spectrometer system (EDX) analysis shows microstructures and relative element abundance of the fossils, but contributes little in solving their biological affinities. The fossils have previously been linked to discoidal impressions of the Ediacara biota. Close examination on new materials indicates that they are radically different from either the Ediacara impressions or any other Precambrian megascopic remains. Concentric or spiral ridges may result from rhythmic growth and the presence of twin specimens may suggest that the organisms undergo asexual reproduction or inhibition of growth in one direction. Referring them to any known fossil or living group has proved to be difficult. We conclude that they represent a distinct group of Precambrian megascopic organisms regardless of their affinities remaining problematic.     

The flux of CO2 and CH4 from lakes and rivers in arctic Alaska

Partial pressures of CO₂ and CH₄ were measured directly or calculated from pH and alkalinity or DIC measurements for 25 lakes and 4 rivers on the North Slope of Alaska. Nearly all waters were super-saturated with respect to atmospheric pressures of CO₂ and CH₄. Gas fluxes to the atmosphere ranged from −6.5 to 59.8 mmol m⁻² d⁻¹ for CO₂ and from 0.08 to 1.02 mmol m⁻² d⁻¹ for CH₄, and were uncorrelated with latitude or lake morphology. Seasonal trends include a buildup of CO₂ and CH₄ under ice during winter, and often an increased CO₂ flux rate in August due to partial lake turnover. Nutrient fertilization experiments resulted in decreased CO₂ release from a lake due to photosynthetic uptake, but no change in CO₂ release from a river due to the much faster water renewal time. In lakes and rivers the groundwater input of dissolved CO₂ and CH₄ is supplemented by in-lake respiration of dissolved and particulate carbon washed in from land. The release of carbon from aquatic systems to the atmosphere averaged 24 g C m⁻² y⁻¹, and in coastal areas where up to 50% of the surface area is water, this loss equals frac 1/5 to 1/2 of the net carbon accumulation rates estimated for tundra.     

The ichnogenus Tremichnus in the Lower Silurian of western New York

Discovery of Tremichnus cysticus Brett on a crinoid pluricolumnal from the Willowvale Shale of western New York extends the range of this ichnogenus into the Lower Silurian. Pits attributable to Tremichnus apparently represent embedment sites of parasites or commensalistic, filter-feeding organisms of unknown affinities. Embedment of T. cysticus may have been accomplished by localized inhibition of stereom growth within the pits aided by secretion of substances that stimulated accretion of stereom by the host pelmatozoan in areas immediately surrounding the pits.     

The root of the problem: palaeoecology of distinctive crinoid attachment structures from the Silurian (Wenlock) of Gotland

The holdfast (attachment structure) is the most understudied aspect of the palaeoecology of the endoskeleton of fossil crinoids. A new collection of well-preserved holdfasts from a recently reopened quarry at Hunninge, Gotland, in Homerian (upper Wenlock) strata includes several morphologies. The most common are terminal dendritic radicular holdfasts (TDRHs) that may be derived from the cladid Ennallocrinus d'Orbigny. These have a consistent morphology of five, equally spaced, long radices that spread across the sea floor. These crinoids were gregarious, and TDRHs in a group commonly show the same radice orientations. The radices have a large axial canal compared with those of modern crinoids; each included, at least, nervous tissues. Taken together, these features suggest that, apart from attachment, these distinctive TDRHs may have served a sensory function. Other holdfasts in this assemblage also show monospecific aggregations, perhaps suggesting biochemical attraction such as that shown by certain other sessile invertebrates such as barnacles.     

Mode of life of the Silurian uncoiled gastropod Semitubina sakoi n. sp. from Japan

Semitubina sakoi n. sp. from the late Silurian of Japan represents the second species of this genus and also the first record of a Silurian gastropod in Japan. The gastropod shells occur in a thin mudstone bed and were found to be encrusted exclusively by corallites of ? Favosites sp. These corallites reveal that encrustation proceeded as the gastropod shells grew. The ecological relationship between the two organisms is considered to be symbiotic. This mode of life allowed the coral to live on a muddy substrate because clear sea water passed over the colony as the gastropod moved along. The gastropod benefited from this relationship by being protected from shell-boring or shell-crushing predators by the encrusting corallite. In Semitubina sakoi the body whorl is separated from the penultimate one by a considerable gap in a later growth stage and S. sakoi has been cited as one of the uncoiled gastropods. The uncoiling of this gastropod results primarily from rapidly increasing whorl translation rate in the latest growth stage. Taking the symbiotic relationship with ? Favosites sp. into consideration, a deposit feeding or benthic scavenging mode of life is suggested for this gastropod.     

The fourth series of the Silurian System in Podolia

The Skala horizon of Podolia has been under discussion as a possible candidate for the stratotype of the fourth Silurian series (primary subdivision) in the international standard chronostratigraphical scale. After a brief survey of the history of investigation, the section through the basal boundary of the Skala horizon is fully documented. The lithostratigraphy, biostratigraphy, and correlation of the Skala horizon are then discussed.     

Trace fossils of the arthropod Camptophyllia from the Westphalian (Carboniferous) rocks of Lancashire, UK and their palaeoenvironmental context

Two arthropod trace fossils are described and analysed from the Carboniferous Lower Westphalian (C. communis and basal A. modiolaris chronozones) coal-bearing strata of Lancashire. The biserial trackway Diplichnites triassicus consists of five overlapping en echelon sets of 7–9 tracks preserved as epichnia and hypichnia in lacustrine siltstones. The trackway suggests subaqueous in-phase walking by a multi-segmented producer with a body length of 35–40 mm, width 17–22 mm, and 7–9 appendages. Curved, clustered, or laterally repeated, hypichnial lobes with transverse striations on the base of ripple cross-laminated sandstone are identified as Rusophycus versans. This trace fossil is interpreted as shallow resting or furrowing burrows of a homopodous arthropod, 30–60 mm long, 15–30 mm wide, and probably the same kind of arthropod as produced D. triassicus.A review of contemporary arthropod body fossils from Lagerstätten in Lancashire favours the onisciform, or Arthropleura like arthropod Camptophyllia as a potential producer of both of these trace fossils in a lacustrine palaeoenvironment.This study integrates the analysis of sediments, trace fossils and body fossils for reconstructing the arthropod biota and ecology in Westphalian lacustrine and crevasse splay fluvial palaeoenvironments.