The Tommotian phase of the Early Cambrian Agronomic Revolution in the carbonate mud environment of central Siberia

The profound ecological change of the marine benthos that eventually led to the almost complete destruction of the Precambrian matgrounds by benthic grazers and bioturbators (the agronomic revolution) was largely completed in the Tommotian. At that time, burrows produced by bottom‐dwelling animals as shelters against predators were supplemented by burrowing for food by predators and sediment feeders. The limy mud ichnofauna of that age in Siberia was very different from the roughly coeval sand bottom faunas of Baltica. Although the exact zoological identity of the animals forming the infaunal Tommotian traces remains unknown, they probably mostly represent various kinds of early nemathelminthes. No apparent locomotion traces of mollusc origin have been encountered in the Early Cambrian, despite the abundant occurrence of skeletal fossils attributed to molluscs. Possibly the standard muscular foot, typical of modern molluscs, had not yet developed. Ichnotaxa represented are Teichichnus isp., Rhizocorallium isp., Chondrites isp., possibly the Buren ichnocomplex and others.     

Devonian Trace Fossils of the Horlick Formation,Ohio Range,Antarctica: Systematic Description and Palaeoenvironmental Interpretation

The clastic Horlick Formation contains an ichnofauna of 28 ichnotaxa dominated by burrowers. These are a mixture of simple vertical forms (Skolithos linearis, S. magnus, Bergaueria cf. langi, Rosselia socialis, Monocraterion isp.), U or arc-like forms (Diplocraterion parallelum, Arenicolites types A and B, Catenarichnus antarcticus, C. isp., aff. Lanicoidichna isp.), and complex, vertical spiral structures (Asterosoma isp., Spirophyton isp.). Horizontal burrows include Ancorichnus cf. capronus, Palaeophycus tubularis, and Psammichnites devonicus isp. nov. Surface traces comprise Haplotichnus isp., Cruziana problematica, C. rhenana, Rusophycus aff. carbonarius, R. isp., Protovirgularia rugosa, Lockeia ornata and cubichnia indet., while trackways include Diplichnites gouldi, D. isp., Maculichna? isp. and large imprints. The Horlick Formation (maximum 56 m) records an early Devonian transgression onto a deeply weathered land area that lay in the direction of Marie Byrd Land, spreading from the South Africa sector of Gondwana. Analysis of the trace fossils confirms their formation in near-shore to intertidal environments, with some ichnotaxa living close to the marine/fluvial boundary (e.g., Spirophyton, aff. Lanicoidichna, Cruziana problematica, Rusophycus aff. carbonarius, Maculichna?). The ichnofauna probably comprised a variety of marine suspension and deposit-feeding worms, shallow burrowing molluscs (bivalves and possibly bellerophontids) and several different types of arthropods, including trilobites.     

Tursia—A New Ichnogenus from Pleistocene Shallow Water Settings in Southern Italy

Abstract

Tursia flabelliformisigen. et isp. nov. is described from the lower Pleistocene Argille Subappennine Formation near Tursi, southern Italy. The trace fossil occurs in protected sandy shoreface sediments and is associated with abundantBichordites. Tursia is a vertical fan-shaped spreite structure that is interpreted as the feeding trace of a deep infaunal deposit-feeding organism, either bivalve or “worm.”     

New Ichnospecies of Cardioichnus from the Miocene of the Guadalquivir Basin,Huelva, Spain

A new ichnospecies of Cardioichnus, Cardioichnus reniformis isp. nov., is documented and described from marine strata of the Late Miocene of southwestern Spain. It is a bilobate resting trace with a heart‐shaped outline and is related to the work of a wedge‐shaped irregular echinoid of the spatangoid group.     

The post‐Palaeozoic fossil record of drilling predation on lingulide brachiopods

Research on drilling predation, one of the most studied biological interactions in the fossil record, has been concentrated on prey with calcareous skeletons (e.g. molluscs, echinoids, rhynchonelliform brachiopods). Based on a compilation of literature sources and surveys of paleontological collections of the Florida Museum of Natural History and the National Museum of Natural History, we provide a tentative evaluation of the post‐Palaeozoic history of drilling predation on the organophosphatic brachiopods of the family Lingulidae. Despite temporal, geographical and methodological limitations of the data assembled here, the results indicate that lingulide brachiopods have been subject to drilling predation since at least the Eocene. Variation in drilling frequencies at the locality level suggests that lingulides may occasionally experience somewhat elevated predation pressures from drilling organisms. Overall, however, drilling predation on lingulide brachiopods has been infrequent in the Cenozoic and may have been absent in the Mesozoic. The Mesozoic‐to‐Cenozoic increase in drilling frequencies on lingulides is similar to the trends observed in other marine benthic invertebrates and consistent with the hypothesis that predation pressures increased through time in marine ecosystems.     

Walking on ashes: insect trace fossils from Laetoli indicate poor grass cover associated with early hominin environments

More than 4000 insect trace fossils collected in recent years from Pliocene deposits at Laetoli in northern Tanzania provide new insights on early hominin palaeoenvironments. These trace fossils include: Fictovichnus gobiensis, Coprinisphaera murguiai, C. kheprii, Coprinisphaera ispp., Quirogaichnus isp., Teisseirei linguatus isp. nov., Celliforma ritchiei isp. nov., C. spirifer, C. germanica, C. cfr. curvata, Celliforma ispp., Rosellichnus isp., Vondrichnus planoglobus, Laetolichnus kwekai igen. et isp. nov. and Krausichnidae indet. They reveal that at least one species of moth, three dung beetles and five other coleopterans, nine taxa of solitary bees, and an indeterminate number of taxa of termites inhabited the Lower Laetolil environments. The Upper Laetolil environments, which have yielded a rich diversity of vertebrate fossils, including the early hominin Australopithecus afarensis and its putative footprints, supported several taxa of termites, one dung beetle, five other coleopterans, and eleven taxa of bees. The Upper Ndolanya environments, which have yielded the hominin Paranthropus aethiopicus, record four taxa of dung beetles, four other coleopterans, and two taxa of bees. The record of larval mortality and lack of intruder activity, revealed by the absence of emergence and intruder traces, may be associated with the anoxic/hypoxic conditions caused by the instantaneous burial of soils under thick volcanic ashes. The record of the Celliforma Ichnofacies in the Upper Laetolil environments indicates the dominance of shrubland to woodland with limited grass cover. This is supported also by the absence of Coprinisphaera, which suggests a scarcity of fresh grasses and a low abundance of large mammal grazers.     

Ichnologic Note A New Ichnospecies of Cardioichnus from the Cretaceous (Albian) of New Mexico

A new ichnospecies of Cardioichnus, Cardioichnus biloba isp. nov., is described and documented from shallow marine strata of Cretaceous (Late Albian) age in southern New Mexico, USA. It is a heart-shaped resting trace which, unlike all other ichnospecies of Cardioichnus, is epichnial and increases in width to a bilobate trace, often with a raised anterior terminus. Only the posterior end of this trace is associated with Bichordites burrows, and the anterior end is bilobate and undisturbed. This represents a resting position which differs from other known species of this cubichnium, which are hypichnial and subquadrate in symmetry. The tracemaker is a spatangoid echinoid, most likely Heteraster d'Orbigny, 1853.     

LATE PRECAMBRIAN TRACE FOSSILS FROM NEW SOUTH WALES

Seven trace-fossil species are described from the upper part of the Torrowangee Group (Upper Proterozoic) of western New South Wales, and a variety of other traces are illustrated. A few forms occur in the Fowlers Gap Beds, and a more diverse and abundant fauna is recorded from the stratigraphically higher Lintiss Vale Beds. Virtually all the traces are preserved in the plane of bedding, as semi-reliefs. The named trace fossils all come from the Lintiss Vale Beds, and are as follows: Planolites ballandus sp. nov., Planolites? sp., Cochlichnus serpens sp. nov., Gordia? sp., Torrowangea rosei gen. et sp. nov., Phycodes? antecedent sp. nov., and Curvolithus? davidis sp. nov. Three of these species are regarded as feeding burrows (endogene), and are thought to represent the activity of infaunal, worm-like deposit feeders. Others may be either feeding burrows or crawling trails. There are also a few impressions which seem to be rest marks. A discussion of the significance of the trace-fossil occurrences is presented.     

Asteriacites lumbricalis von schlotheim 1820: Ophiuroid trace fossils from the lower triassic thaynes formation,central Utah

Several trace fossils of burrowing ophiuroids, Asteriacites lumbricalis von Schlotheim 1820, occur on slabs of thinly‐bedded silty limestone in the lower Spathian part of the Lower Triassic Thaynes Formation in central Utah. This is the first record of Asteriacites in the Lower Triassic of North America. Their occurrence adds information to studies of the recovery of marine faunas following the Permian extinctions, including the oxygénation of benthic sediments.     

Upper Triassic (Keuper) non-marine trace fossils from the Haßberge area (Franconia, south-eastern Germany)

Various trace fossils from the Hassberge Formation and the Löwenstein Formation (Middle Keuper, Upper Triassic) of the Haßberge region are described. Twenty-three different forms have been identified, 17 of which are named, includingCruziana pascens n. isp.,Lockeia cunctator n. isp., andRusophycus versans n. isp.Lockeia siliquaria James, 1879,L. amygdaloides (Seilacher, 1953),L. triangulichnus Kim, 1994, andL. elongata (Yang, 1984) are revised and synonymized under the oldest available name,L. siliquaria James, 1879.Rusophycus eutendorfensis (Linck, 1942) andR. carbonarius Dawson, 1864 are revised. The diagnosis ofPolykladichnus Fürsich, 1981 is emended, and a diagnosis forHelminthoidichnites Fitch, 1850 is given for the first time. Among the described ichnotaxa,Skolithos ispp.,Rusophycus carbonarius, andTaenidium barretti are the most common forms. The trace fossil association is typical of theScoyenia ichnofacies, which indicates non-marine, periodically or completely inundated environments, such as floodplains and lake margins. Two palaeoichnocoenoses are identified. One ichnocoenosis, dominated byCruziana problematica, cf.Polykladichnus isp., andSkolithos isp. B characterizes margins of trough cross-bedded sandstones. Another ichnocoenosis, dominated byRusophycus versans n. isp.,Taenidium barretti,Scoyenia gracilis andSkolithos isp. A is related to ephemeral lake deposits. Taxonomic recommendations for the use of hitherto described and figured invertebrate Keuper trace fossils from Germany are given.     

The earliest fossil evidence of bone boring by terrestrial invertebrates,examples from China and South Africa

We report the oldest fossil evidence of osteophagia by terrestrial invertebrates on both the Asian and African continents. Bones attributable to the Middle Jurassic dinosaur Chuanjiesaurus (Dinosauria: Sauropoda) were found with post-mortem insect modification in the Chuanjie Formation, Yunnan Province, China. The morphology of the borings closely matches the ichnogenus Cubiculum. Based on the lack of bioglyphs observed in Cubiculum ornatus, a new ichnospecies is proposed here. The new trace fossil, Cubiculum inornatus isp. nov., is interpreted to have been constructed for pupation by an unknown taxon of insect. Additionally, we report even older borings from Early Jurassic dinosaur bones of the Elliott Formation in the Karoo Basin, which represent the second oldest occurrence of insect traces in bone from continental settings. Both trace fossils sites have palaeogeographic implications for the origins and dispersal of osteophagia amongst terrestrial invertebrates during the Mesozoic. These discoveries push back the antiquity of pupation in animal bones by more than 100 million years to the Middle Jurassic, indicating that this behaviour, and osteophagy more generally, originated early in the Mesozoic, roughly comparable with the origination of insect pupation in woody substrates (Late Triassic).     

Deep Endichnial Cruziana from the Lower-Middle Ordovician of Spain — A Unique Trace Fossil Record of Trilobitomorph Deep Burrowing Behavior

Full reliefs of Cruziana furcifera from the Lower-Middle Ordovician quartzite sandstone beds (Pochico Formation, southern Spain) points to deep, infaunal burrowing of trilobites. Some specimens show an unusual vertical extension with a wider lower part and a narrower upper part in cross section. They are referred to trilobites, which burrowed deeply in the sediment and were oriented obliquely head down and tail up. Deep burrowing seems to be common for other members of the Cruziana rugosa group, foremost C. rugosa and C. furcifera, less for C. goldfussi. The deep burrowing recorded in the discussed trace fossils can be referred to the earliest common infaunalization caused by trilobites and other arthropods during the Ordovician, probably in a response to a food competition on the sea floor, which promoted a behavioral plasticity within the same taxon or closely related taxa of trilobites.     

Morpho-anatomical differences of the Early Cambrian Chengjiang and Recent lingulids and their implications

Since the publication of Darwin's theory of evolution in 1859, lingulids have probably been the most widely quoted examples of arrested evolution. This, to some degree, may be because few anatomical features are impressed either on or inside valves so these may not adequately reflect the extent of change incurred during lingulid evolution. Two lingulid brachiopods, Lingulella chengjiangensis and Lingulellotreta malongensis, from the Early Cambrian Chengjiang Lagerstätte (Yunnan, South China) show preservation of a series of soft parts, notably the lophophores, setae mantle canals and perfectly impressed visceral region, which are vital to understanding the evolution and lifestyle of brachiopods. Analysis of the valve interiors favours the claim that an epifaunal mode of life could be a plesiomorphic state in contrast to the infaunal one of modern lingulids. Based on these fossils, the three‐pseudosiphon formation of setae found in Recent lingulids is proposed to be an apomorphy derived as an adaptation to an infaunal lifestyle. Comparison between the interior of the fossils with that of modern lingulids does little to support the widespread notion that the morphology of this lineage has remained remarkably constant since at least the early Palaeozoic.     

A comparison of crayfish burrow morphologies: Triassic and Holocene fossil,paleo‐ and neo‐ichnological evidence,and the identification of their burrowing signatures

The architectural and surficial morphologies of crayfish burrows from the Upper Triassic Chinle Formation and Holocene sediments were compared in order to determine: 1) if Triassic burrows could truly be attributed to crayfish activity; 2) how comparable the burrowing mechanisms are; and 3) whether or not a common set of burrowing signatures could be identified for both ancient and modern freshwater crayfish. Materials used in this study include burrows from the members of the Upper Triassic Chinle Formation, casts of modern burrows constructed by Procambarus clarkii Hobbs and Procambarus acutus acutus (Girard) in the laboratory, and casts of naturally constructed modern burrows of Cambarus diogenes di‐ogenes (Girard).

Triassic and Holocene crayfish burrow morphologies exhibit simple to complex architectures, varying degrees of branching, chamber, and chimney development. They also exhibit relatively textured surficial morphologies (bioglyphs) such as scrape and scratch marks, mud‐ and lag‐liners, knobby and hummocky surfaces, pleopod striae, and body impressions. Holocene crayfish construct distinctive burrows due to their conservative limb arrangement, functional morphology, and behavior with respect to environmental stimuli. Similarities between Holocene and Triassic crayfish burrows suggest that extant and Triassic crayfish employed identical burrowing mechanisms. Features of the surficial and architectural morphologies impart a distinctive signature to burrows of both ancient and modern freshwater burrowing crayfish.

Burrowing signatures of crayfish can be used to identify new and previously misinterpreted continental trace fossils. These are useful in studies of the paleohydrogeology, paleoclimatology and paleoecol‐ogy of burrow‐bearing deposits.     

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.     

Parataenidium,a new taenidium‐like ichnogenus from the carboniferous of Ireland

The new ichnogenus Parataenidium is erected for backfilled tubular trace fossils that can appear superficially similar to Taenidium, but are divided horizontally into two distinct levels. Two ichnospecies are recognised: Parataenidium mullaghmorensis isp. nov. and Parataenidium moniliformis (Tate 1859). The latter ichnospecies is transferred from Eione Tate 1859, which is a junior homonym of Eione Rafinesque 1814, and therefore unavailable for Tate's ichnotaxon. The ichnogenus is an important component of late Paleozoic shallow‐water siliciclastic sediments, and can be considered as a “guide”; indicator for the Carboniferous.     

A LEAFCUTTER BEE TRACE FOSSIL FROM THE MIDDLE EOCENE OF PATAGONIA, ARGENTINA, AND A REVIEW OF MEGACHILID (HYMENOPTERA) ICHNOLOGY

Abstract:  The ichnospecies Phagophytichnus pseudocircus isp. nov. is described to include trace fossils characterized by leaf-margin excisions showing eccentricity values of 0.35–0.65 and more than 270 degrees of an arc, a non cuspate margin and vein stringers or necrotic flaps of tissue along the margin. A method for determining ellipse eccentricity was performed on leaf discs obtained from the nests of the modern leafcutter bee Megachile rotundata (Hymenoptera: Megachilidae), which provided objectively obtained values comparable to the trace fossil from the middle Eocene of Argentina and other world-wide ichnological records, historically and subjectively considered to be 'circular' trace fossils and attributed to leafcutter bees. The material described herein represents the first evidence for fossil Megachilidae from the Southern Hemisphere.     

Bored Bivalves in Upper Triassic (Norian) Event Beds,Northeastern British Columbia,Canada

Skeletobionts are important components of most shallow marine ecosystems. Prior to the fossils reported herein, evidence of skeletobionts was absent from Upper Triassic successions on the northwestern margin of Pangaea. The boring Talpina ramosa is reported from bivalve body fossils from the Upper Triassic (Lower Norian) lower Pardonet Formation at Pink Mountain in northeastern British Columbia. This Ichnotaxon penetrates through both the outer and inner surface of articulated and disarticulated bivalve shells preserved within sharp-based event beds. The occurrence of these trace fossils underscores the paucity of borings, bioerosional structures, and encrusting taxa from Triassic successions in the western Pangaean realm. Due to erosional removal of shallow water strata by a post-Triassic unconformity, these event beds provide the only available information regarding the ecological health of Late Triassic depositional systems in the study area.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.