A New Ichnospecies of Arthrophycus from the Late Carboniferous (Pennsylvanian) of Michigan,USA

A unique ichnofossil assemblage from Pennsylvanian-age sandstones near Eaton Rapids, Michigan, USA, comprises straight-to-curved traces preserved in convex hyporelief, with the transverse ridges and median grove associated with the ichnogenus Arthrophycus. The Michigan traces show some branching or pseudo-branching (also known from other Arthrophycus specimens) and are among the smallest structures (millimeters in diameter) attributed to this ichnogenus. The orientation of the Michigan Arthrophycus burrows is distinct from other ichnospecies of this taxon in the pronounced co-planar alignment of the burrows, as opposed to the multiplanar, interweaving, “bundled” nature typical of the ichnogenus. On this basis we assign the Michigan specimens to a new ichnospecies of Arthrophycus. The “paralleling behavior” of the new taxon may reflect a strategy of the tracemaker to avoid previous burrows and reflect differences in resource availability or current energy. This new taxon supports previous records of the occurrence of this ichnogenus in Upper Carboniferous strata.     

Silurian trace fossils in carbonate turbidites from the Alexander arc of southeastern Alaska

Early to Late Silurian (Llandovery‐Ludlow) body and trace fossils from the Heceta Formation of southeastern Alaska are preserved in the oldest widespread carbonates in the Alexander terrane. These fossils represent the earliest benthos to inhabit diverse shallow and deep subtidal environments in the region and are important indicators of early stages in benthic community development within the evolving Alexander arc. The ichnofossils are particularly significant because they add to a small but growing body of knowledge about trace fossils in deep‐water carbonates of Paleozoic age.

Carbonate turbidites that originated along a deep marine slope within the arc yield a low‐diversity suite of trace fossils consisting of five distinct biogenic forms. Simple burrows (Planolites, two forms), ramifying tunnels (Chondrites), and tiny cylindrical burrows (?Chondrites) represent the feeding activities (fodinichnia) of pre‐turbidite animals that burrowed in the lime mud before the influx of coarser sediment deposited by turbidity currents. These trace fossils are associated locally with cross‐cutting burrows created as domichnia (Palaeophycus). Rarer hypichnial burrows and endichnial traces were created by post‐turbidite animals that fed soon after the deposition of coarse detritus from turbidity flows.

Trace fossils in these deposits reflect much lower diversity levels than in Paleozoic siliciclastic turbidites. This difference may represent unfavorable environmental conditions for infaunas, differential preservation, or significant paleogeographic isolation of the Alexander terrane during the Silurian. Greater utilization of trace fossils in terrane analysis may help to resolve this issue and provide new data for reconstructing the paleogeography of circum‐Pacific terranes.     

Furrows and firmgrounds: evidence for predation and implications for Palaeozoic substrate evolution in Rusophycus burrows from the Silurian of New York

Tarhan, L.G., Jensen, S. & Droser, M.L. 2011: Furrows and firmgrounds: evidence for predation and implications for Palaeozoic substrate evolution in Rusophycus burrows from the Silurian of New York. Lethaia, Vol. 45, pp. 329–341. The Silurian Herkimer Formation of east‐central New York contains abundant, exceptionally preserved composite Rusophycus‐Teichichnus burrows. We suggest that the most likely interpretation of these composite trace fossils is as structures formed by trilobites entering the sediment in search of prey. Parallel alignment of the paired traces, asymmetrical configuration of the Teichichnus along the longitudinal axis of the associated Rusophycus, depth correlation and deformation of the Teichichnus all suggest that this relationship was predatory. In addition, sectioned material indicates that these Rusophycus may have been open at the sediment‐water interface, while the crisp preservation of both Rusophycus and Teichichnus, along with the preservation of such delicate morphological details as scratch marks, suggests that the sediment must have been relatively firm at the time the traces were formed. The formation and preservation of Rusophycus in cohesive sediments located very close to the sediment‐water interface hold important implications for the manner in which we consider Palaeozoic substrates and their temporal and spatial evolution. Moreover, these findings demonstrate that the morphology and taphonomy of ichnological associations may, in the context of sedimentological relationships, prove a powerful proxy for tracking substrate conditions through both space and time. □firmgrounds, New York, predation, Rusophycus, substrate, taphonomy.     

Problematic megafossils in Cambrian palaeosols of South Australia

Abstract: Red calcareous Middle Cambrian palaeosols from the upper Moodlatana Formation in the eastern Flinders Ranges of South Australia formed in well‐drained subhumid floodplains and include a variety of problematic fossils. The fossils are preserved like trace fossil endichnia but do not appear to be traces of burrows or other animal movement. They are here regarded as remains of sessile organisms, comparable with fungi or plants living in place, and are formally named as palaeobotanical form genera under provisions of the International Code of Botanical Nomenclature. Most common are slender (0.5–2 mm) branching filaments flanked by green‐grey reduction haloes within the red matrix of palaeosol surface horizons (Prasinema gracile gen. et sp. nov.). Other axial structures (Prasinema nodosum and P. adunatum gen. et spp. nov.) are larger and show distinctive surface irregularities (short protuberances and irregular striations, respectively). The size and form of these filaments are most like rhizines of soil‐crust lichens. Other evidence of life on land includes quilted spheroids (Erytholus globosus gen. et sp. nov.) and thallose impressions (Farghera sp. indet.), which may have been slime moulds and lichens, respectively. These distinctive fossils in Cambrian palaeosols represent communities comparable with modern biological soil crusts.     

Complex behavioural patterns and ethological analysis of the trace fossil Zoophycos: evidence from the Lower Devonian of South China

The trace fossil Zoophycos is abundant in the shallow‐marine deposits (tempestites) of the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by deposit‐feeding behaviour, where the animal constructed the trace upwards without leaving faeces in the spreiten. The width of the marginal tube in different whorls is almost constant. The scratches are observed on the wall of the marginal tubes. The Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos‐maker might have been a vermiform polychaete instead of a predator such as a decapod crustacean (Spongeliomorpha producer). Based on stratigraphical and ichnological features, the complex trace fossils resulted from the complex activity of different opportunistic organisms (r‐strategist) that quickly occupied and thrived within the quiet, nutrient‐rich environment after storm events.     

Revision of the disparid Stylocrinus (Crinoidea) from the Devonian of Europe,Asia and Australia

Abstract: The discovery of new specimens and the restudy of known collections result in revision of the diagnosis and the stratigraphic distribution of the disparid crinoid genus Stylocrinus, from the Middle and Upper Devonian of Europe, Asia and Australia. The consistent development of three basal plates, the atomous arms with internally inclined edges adjoining laterally with adjacent brachials in an interlocking network and an apparently rudimentary pinnulation is recognised. The high ecophenotypic plasticity of the common species S. tabulatus negates the validity of several former subspecies and demonstrates the general morphologic variability of the aboral cup proportions. This contrasts with the low morphological spectrum of rarer stylocrinid species. With exclusion of ‘S. elimatus’ (Silurian) from Stylocrinus, the genus is limited to the Devonian. A neotype is proposed for the lost holotype of S. tabulatus. Stylocrinus prescheri sp. nov. is described from the Eifelian to Givetian of Europe and Asia. The first evidence of the gastropod grazing trace fossil Radulichnus on a crinoid aboral cup (S. tabulatus), the post‐mortem incurred ossicular‐boring of radial and basal plates as well as the post‐mortem encrusting by a rugose coral are further observations on Stylocrinus aboral cups.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

History of Ichnology: The Origins of Trace Fossil Taxonomy and the Contributions of Joseph F. James and Walter H. Häntzschel

The taxonomy of trace fossils has had a somewhat controversial history because they do not represent the actual animal remains but rather their work on and in the substrate. As such, traditional palaeontologists and zoologists have viewed them with some skepticism. Ichnologists owe a great debt to two geologists: Joseph F. James of Cincinnati and Walter H. Häntzschel of Hamburg, who took it upon themselves to impose some order on the chaos that constituted trace fossil taxonomy at the time. James, working independently and in ignorance of Alfred Nathorst, arrived at and utilized many of the same criteria his Swedish counterpart employed to criticize the fucoid origins of many trace fossils in the late 19th century. With his restudy of the systematics of Fucoides, Skolithos, and Arthrophycus, James brought to light many of the taxonomical nightmares that faced—and are still facing—the fledging science and can be rightfully considered the first trace fossil taxonomist. During the 1940s and 1950s, Häntzschel collected the widely scattered pertinent data from the literature, an immense task that, when published in 1962 (and later revised and expanded in 1975), made trace fossils accessible to further research and started a worldwide boom in trace fossil research.     

Reassessment of sediments and trace fossils from old red sandstone (Lower devonian) of Dunure,Scotland, described by John Smith (1909)

In 1909 John Smith, a Scottish naturalist and geologist,described 23 «genera and 51 «species of trace fossils from small patches of sediment associated with andesite lava flows at Dunure, Ayrshire. He interpreted the traces as evidence of a diverse invertebrate fauna which inhabited small pools and fissures in the lava surface between eruptions. Smith's collection (c. 300 specimens) was presented to the British Geological Survey, Edinburgh but has remained largely unstudied.Re-examination of the Smith collection shows thatit came from 3 separate localities which differ in composition of the ichnofaunas, associated sediments and sedimentary structures. Arthropod trackways dominate the ichnofauna occurring in laminated siltstone frequently with ripple marks and foam marks suggesting formation in shallow lacustrine conditions. Preservational and behavioural analysis of trace fossils reveals about 10 valid ichnogenera of locomotion and swimming trackways, resting traces, feeding trails but few burrows. Presence of early terrestrial arthropod traces is uncertain. Palaeocological interpretation is of ephemeral lakes in distal braid-plain situation subject to subsurface invasion of andesite lava producing fluidization and deformation of wet sediment. The Dunure ichnocoenosis shows unique diversity of Devonian arthropod trace fossils.     

Enigmatic occurrence of Permian plant roots in Lower Silurian rocks,Guizhou Province,China

Pinnatiramosus qianensis Geng, 1986, is a plant with a complex, extensive pinnate branching system and pitted tracheids, collected from marine Lower Silurian (Llandovery; c. 430 Ma) rocks in Guizhou Province, China. It challenges long‐held theories on the origin and early evolution of vascular plants in the Silurian and Devonian. However, there is a hypothesis that the fossils were not syngenetic with the entombing rock, but were the rooting systems of much younger plants, probably of Permian age. New sections and collections of P. qianensis have been subjected to detailed analyses, which indicate that P. qianensis belongs to an early Permian (c. 285 Ma) rooting system growing down into lower Silurian rocks.     

Evidence for a fungal affinity for Nematasketum,a close ally of Prototaxites

As part of a project investigating diversity in early terrestrial vegetation, new specimens of the enigmatic, non‐embryophyte Nematasketum diversiforme have been described from Lower Devonian strata in the Welsh Borderland to elucidate its affinities and role in the ecosystem. Charcoalified fragments were investigated using scanning electron microscopy. They show for the first time that at least part of the organism possessed an axial organization with a peripheral zone forming a rind. The specimens differ from the closely‐allied Prototaxites by the presence of unevenly thickened tubes, both in well‐defined areas (medullary spots), hypothesized as being the sites of hyphal generation, and in the matrix of wide, smooth‐walled and narrow hyphae. Two specimens that show branching but no unequivocal thickened tubes are best assigned to the prototaxalean complex. The fragments, with longitudinally aligned wide tubes in a matrix of much branched, narrower examples, resemble rhizomorphs and cords of extant basidiomycote fungi and are interpreted as exploratory and translocating organs. Postulated affinities of Prototaxites include algae, fungi and hepatics, with the latter interpreting the fossils as rolled‐up liverwort mats. Our studies on a close ally of Prototaxites show no evidence for hepatic affinity, although similarities with fungal linear mycelial structures. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 1–18.     

Host-specific pit-forming epizoans on Silurian crinoids

Brett, Carlton E. 197807 15: Host-specific pit-forming epizoans on Silurian crinoids. Lethaia , Vol. 11. pp. 217–232. Oslo. ISSN 0024–1164.
Circular-parabolic pits occur commonly on the endoskeletal remains of certain Paleozoic crinoids. Detailed study of several hundred specimens, representing about 30 pelmatozoan species from the Upper Silurian Rochester Shale of New York and Ontario, reveals that such pits occur exclusively in seven species of crinoids. Furthermore, there are consistent differences in the morphology and orientation of holes occurring on the different crinoid species. This suggests that distinct epizoan species settled selectively on given hosts. The relationship between the hole-producing epizoans and crinoid hosts is inferred to have been a form of dependent commensalism. Preliminary surveys of other Paleozoic crinoid assemblages reveal similar host-selectivity by pit-producing epizoans. Crinoidepizoan pairs apparently co-evolved through considerable spans of geologic time as related genera and species of different ages, from Silurian to Pennsylvanian, exhibit similar pits.     

An integrative ichnological and taphonomic approach in a transgressive–regressive cycle: a case study from Devonian of Paraná Basin,Brazil

The palaeoenvironmental context of a section of the Devonian Ponta Grossa Formation (Paraná Basin) was examined using an integrated ichnological and taphonomic approach. Three taphofacies (T‐A, T‐B and T‐C) and six ichnofabrics are recognized. T‐A is mainly composed of disarticulated organisms and is associated with Chondrites, Planolites‐Palaeophycus, Asterosoma‐Zoophycos and Asterosoma‐Chondrites ichnofabrics, representing the Cruziana ichnofacies. The upward increase in diversity and abundance of body fossils for Taphofacies A can be explained by changes in sedimentation rates during aggradational to progradational phases. T‐B beds, represented by articulated brachiopods, occur in distal tempestites associated with the Skolithos ichnofabric (Skolithos ichnofacies) and represent higher hydrodynamic energy and oxygenation than in T‐A. Taphofacies C, characterized by a mix of articulated and disarticulated organisms, generally lacks bioturbation except for Chondrites ichnofabrics occurring only at the bed tops, suggesting low substrate oxygenation and low‐energy conditions. T‐C indicates the maximum transgression in the T‐R cycle, characterizing it as a good indicator of maximum flooding surfaces. The composite Asterosoma‐Teichichnus ichnofabric (bioturbation degree 5) occurs in layers without body fossils and represents suites of the Cruziana ichnofacies. This likely reflects intense intrastratal activity at all substrate levels, which facilitated the oxygenation of the substrate disallowing the preservation of organism remains. This observation indicates that the bioturbation is an important factor controlling the loss of taphonomic information within the TAZ, mainly when associated with detritus‐feeding trace fossils.     

New trace fossil evidence for eurypterid swimming behaviour

We describe a recently discovered trace fossil from a eurypterid Konservat‐Lagerstätte in the upper Silurian Tonoloway Formation of Pennsylvania, and formally describe contemporaneous traces from the Williamsville Formation Lagerstätte of Ontario. The traces from both localities are assigned here to Arcuites bertiensis igen. et isp. nov. Based on comparisons with previously described eurypterid trackways, neoichnological experiments, and the co‐occurrence with eurypterid remains, Arcuites is interpreted as having been made by the swimming leg (sixth prosomal appendage) of swimming juvenile to adult eurypteroid eurypterids, and represents the first unambiguous trace fossil evidence for eurypterid swimming behaviour. The morphology of Arcuites indicates that eurypteroid eurypterids swam using drag‐based rowing, whereby the animal propelled itself forward by moving its oar blade‐like swimming paddles in an in‐phase backstroke. Arcuites morphology also indicates that the eurypteroid swimming appendage had a greater degree of movement than was previously suggested, and a revised rowing model is proposed. Differences in the abundance of A. bertiensis in the Tonoloway and Williamsville formations suggest a bathymetric control on eurypterid swimming behaviour and trace production. The association of Arcuites with eurypterid body fossils in both units indicates that these Lagerstätten were autochthonous assemblages and provides additional evidence for eurypterid inhabitation of shallow subtidal marine environments in the late Silurian.     

Reassessment of the Devonian problematicum Protonympha as another post‐Ediacaran vendobiont

Protonympha is an enigmatic fossil represented by two species from the Middle Devonian (Protonympha transversa) and Late Devonian (Protonympha salicifolia) of New York. Although interpreted in the past as a polychaete worm or starfish arm, Protonympha is not found with marine fossils, but with fossil plants. This fossil plant community was a swamp woodland of Lepidosigillaria whitei, with ground cover of Haskinsia colophylla, fringing brackish to freshwater coastal lagoons of the Catskill Delta. Protonympha shares with Ediacaran Vendobionta a quilted body of unskeletonized biopolymer that is unusually resistant to burial compaction. In overall form, Protonympha is most like the Ediacaran genus Spriggina. Protonympha has branching and tapering tubular structures radiating from the bottom. These rhizine‐like structures, thallus stratification and internal chambers revealed by petrographic thin sections suggest affinities with lichenized fungi. As for Cambrian Swartpuntia and Ordovician–Silurian Rutgersella, Protonympha may have been a post‐Ediacaran vendobiont.     

Ichnology of deep‐sea fan overbank deposits of the Ganei slates (Eocene,Switzerland)— a classical flysch trace fossil locality studied first by Oswald Heer

Ganei (Switzerland) is a classical locality for trace fossils. At this site, Heer (1877) described a large number of trace fossils, several of which were new taxa. The trace fossils occur in thin‐bedded turbidites in which the basal divisions of the Bouma sequence are typically absent; the turbidites are assigned to the Ganei Slates and are Eocene in age. They are interpreted to have been deposited in an overbank environment within an upper to middle fan area distal to a channel. Two trace‐fossil associations occur: the first (I) is characterized by bulldozing organisms producing biodeformational structures, Scolica, and Nereites irregularis; the second (II) association shows a distinct tiering pattern with near‐surface graphoglyptids and a mixed layer with simple tubes such as cf. Palaeophycus and Planolites, plus patterned tubes such as Nereites cirrinalis, and Chondrites. Deeper turbidite layers were colonized by Chondrites and Gyro‐phyllites. All trace fossils show a normal size spectrum compared to previously studied trace‐fossil associations, so the degree of oxygenation probably did not influence the composition of either trace‐fossil association. Seafloor sediment was probably soft and did not affect the trace‐fossil associations. Sedimentation rate and event frequency did not change and are estimated to have been in a range of 5–10 cm/1000 years and 2–5 events per 1000 years, respectively. The composition of trace‐fossil associations I and II is therefore interpreted to have been controlled by the benthic food content being higher for trace‐fossil association I than for II.     

Precambrian–Cambrian boundary interval occurrence and form of the enigmatic tubular body fossil Shaanxilithes ningqiangensis from the Lesser Himalaya of India

The affinity of the Ediacaran fossil Shaanxilithes ningqiangensis and putatively related forms has long been enigmatic; over the past few decades, interpretations ranging from trace fossils to algae to metazoans of uncertain phylogenetic placement have been proposed. Combined morphological and geochemical evidence from a new occurrence of S. ningqiangensis in the Krol and Tal groups of the Lesser Himalaya of India indicates that S. ningqiangensis is not a trace fossil, but rather an organic‐walled tubular body fossil of unknown taxonomic affinity. Specimens consist of compressed organic cylindrical structures, characterized by extended, overlapping or fragmented iterated units. Where specimens intersect, overlapping rather than branching or intraplanar crossing is observed. Lithologic comparisons and sequence stratigraphic data all suggest a late Ediacaran age for the uppermost Krol Group and basalmost Tal Group. By extending the biogeographical distribution of S. ningqiangensis, hitherto confined to the Ediacaran of China and potentially Siberia, to the Precambrian–Cambrian boundary interval of India, this new occurrence of S. ningqiangensis expands the biostratigraphic utility of this enigmatic fossil to the inter‐regional and intercontinental scale. Moreover, study of these new and exceptionally preserved samples may help to significantly constrain the long‐debated problem of Shaanxilithes' affinity, elucidating its ‘problematic’ status and shedding new light upon the ecology and taphonomy of one of the most significant intervals in early life history.     

Palaeoenvironmental implications of the ichnology and geochemistry of the Westbury Formation (Rhaetian), Westbury‐on‐Severn,south‐west England

Abstract: The Westbury Formation (Rhaetian) beds of Westbury Garden Cliff, Westbury‐on‐Severn, west of Gloucester, Britain, show an unusual combination of features. Both deep water and emergent characteristics are present within the sediments and the trace fossils. The ichnoassemblage consists of abundant Selenichnites, Planolites beverlyensis and Lockeia with rarer Oniscoidichnus, Chondrites, Rhizocorallium irregulare, Taenidium serpentium, an unusual form of Walcottia and Merostomichnites‐like traces. These trace fossils display an interesting relationship with the sediments: low‐energy Cruziana ichnofacies is found within high‐energy sandstones. The sandstones are interbedded with laminated mudstones, apparently deposited in deep water, but some aspects of the ichnoassemblage, preservation and sedimentation indicate shallower water. One new trace fossil, Radichnus allingtona igen. et isp. nov., closely resembles the traces of modern fiddler crabs and imply emergence, by analogy. This ichnofauna is similar to early stage disaster colonisation in recent experiments in Long Island Sound (south of Connecticut, USA) and with storm‐influenced deposits within the Cardium Formation (Seebe, Alberta, Canada). This indicates a lagoonal environment with influxes of sand and oxygen. Total organic carbon levels were found to fluctuate greatly between stratigraphic layers but remained relatively high. This implies low oxygen conditions. The abundance of sulphur (in pyrite) also supports an interpretation of anoxic conditions, and low sedimentation rates within the shale layers. A restricted shallow basin or lagoonal environment is proposed for the palaeoenvironment, with fluctuating oxygen influencing diversity.