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.     

Tripartichnus n. igen. — a new trace fossil from the Buntsandstein (Lower Triassic) and from the Solnhofen Lithographic Limestones (Upper Jurassic), Germany

Two species of a new trace fossil genus (cubichnia) are described:Tripartichnus triassicus n. igen., n. isp. from the “Plattensandstein“ (Upper Buntsandstein, Röt-Folge, Lower Triassic) of Keltern-Dietlingen (Enzkreis, Baden-Württemberg, SW-Germany) andTripartichnus imbergi n. igen., n. isp. from the Solnhofen Lithographic Limestones (Upper Jurassic, Tithonian) from Waiting and Pfalzpaint (district of Eichstätt, Bavaria, SE Germany). Due to the characteristic shape of the new ichnotaxa the tracemakers can be named in all probability:Tripartichnus triassicus was most likely made byEuthycarcinus kessleri Handlirsch, 1914 andTripartichnus imbergi byPalinurina sp.     

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.     

Scolicia shirahamensis isp. nov.: a triple-corded scolicia and its ichnological implications

Abstract

Peculiar meniscate burrows with three sediment cords occur in early to middle Miocene tidal-flat deposits of southwestern Japan. Two of the cords are situated at the bottom and the other is at its center. Detailed observations of the burrow structures and comparative neoichnological studies of modern spatangoid burrows in a tidal flat revealed that the former two were true drainage tubes and the latter was fecal in origin. The trace fossil was thus assigned to the ichnogenus Scolicia. Based on these findings, a new ichnospecies Scolicia shirahamensis isp. nov. has been described here. The central sediment cord is seemingly identical to the drainage tube of the ichnogenus Bichordites, another ichnogenus that has been commonly ascribed to a fossil spatangoid burrow, similar to Scolicia. Careless ichnogeneric identification of a spatangoid burrow, based only on the central sediment cord, therefore, may produce an incorrect identification.     

Lithostratigraphy of the Lower Cambrian metaclastics and their age based on trace fossils in the Sand?kl? region, southwestern Turkey

In the Sand?kl? region of the Taurus Range of Turkey, greater than 3000 m in thickness metamorphosed siliciclastics and volcanics (Kocayayla Group) underlies the trilobite-and conodont-bearing Middle-Late Cambrian Hudai Quartzite and Çaltepe Formation.The Kocayayla Group, previously regarded as Infracambrian or Precambrian, is dated for the first time as Early Cambrian on the basis of trace fossils. Cruziana ?fasciculata, C. ?salomonis, ?Cruziana isp., ?Diplichnites isp., Monomorphichnus isp., Petalichnus isp., Rusophycus ?avalonensis, R. ?latus, Arenicolites isp., cf. Altichnus foeyni, Planolites isp., Skolithos isp., and ?Treptichnus isp. have been recognised. These trace fossils are considered Tommotian or younger in age but older than the overlying, trilobite and conodont bearing Middle Cambrian limestones of the Çaltepe Formation. The trace fossils were likely produced by trilobites, suspension feeding annelids and deposit feeding “worms”, probably polychaetes. Sections bearing abundant Skolithos represent the Skolithos ichnofacies, which is typical of high energy environments with loose sandy, well sorted to slightly muddy substrates in intertidal to shallow subtidal zones. The other trace fossils represent the Cruziana ichnofacies, which is typical of subtidal, poorly sorted and soft substrates, from moderate energy to low energy environments between the fairweather and storm wave base.The Kocayayla Group was deposited at an early stage in a shallow marine stable shelf condition. The shelf subsided in a later stage and was affected by normal faults along which mafic and felsic volcanic rocks erupted. The volcanic activity had ceased and a shallow marine clastic sedimentation took place in the final stage of the shelf development. The Kocayayla Group was deformed and metamorphosed before the deposition of the trilobite-bearing Middle-Upper Cambrian succession.     

Scolicia and its producer in shallow-marine deposits of the Miocene Chenque Formation (Patagonia,Argentina): functional morphology and implications for understanding burrowing behavior

Abstract

Scolicia is one of the most conspicuous trace fossils in lower shoreface deposits of the Miocene Chenque Formation of Patagonia, Argentina. This ichnotaxon consists of horizontal, sinuous or meandering trace fossils with a laminated backfill and two parallel strings located at the base. Abundant body fossils attributed to Brisaster iheringi occur in close association to these trace fossils. The echinoids are very well preserved, and most of the specimens have some areas with their spines attached in life position. In particular, preservation of the subanal tufts of spines, associated to isopores (which are interpreted as being associated to funnel building tube feet), supports the interpretation that these organisms are the producers of Scolicia isp. A shaft connecting the burrow with the sediment-water interface was not observed in vertical sections of the excavations, although on bedding plane surfaces, some small circular tube outlets are recognized. However, the great diversity and abundance of ichnofossils co-occurring in these deposits may preclude adequate identification of vertical structures. These deposits provide an excellent opportunity to integrate trace fossil data, body fossil information and observations from modern analogues in order to perform a morpho-functional analysis of Scolicia.     

Lopingian (Upper Permian) trace fossils from the northern Penglaitan Section,Laibin, Guangxi,South China and their environmental implications

The end-Permian mass extinction has been widely documented to be accompanied with oceanic anoxia, which was considered as one of the most plausible killing mechanisms. However, it is still unclear when anoxia began to occur and how widely affected during the pre-extinction interval. In this study, Lopingian bottom-water oxygen level changes around the Wuchiapingian–Changhsingian boundary (WCB) at the northern Penglaitan section in Guangxi are analysed based on trace fossil assemblages and ichnofabric indices. Detailed bed-by-bed ichnological analyses confirm the presence of trace fossils Chondrites intricatus, C. isp., Palaeophycus isp., Planolites isp. A, P. isp. B, Thalassinoides isp. A, T. isp. B, Trichichnus linearis, Zoophycos isp., and other bioturbational structures. The ichnofabric indices around the WCB are mostly 1–2, but rapidly increase to 4–5 in some short intervals. The distinct fluctuations of ichnofabric indices suggest that the oxygen level of sediments was generally deficient, but the dysoxic or anoxic periods were punctuated by several short oxic intervals. Thus, it can be inferred that the benthic organisms were under secularly environmental stresses around the WCB at the northern Penglaitan section. The anoxic conditions unfavourable for aerobiosis in the deep basin may start in the late Wuchiapingian. However, more sections need to be studied to determine whether this is a local or regional phenomenon.     

Determination of a Late Miocene rocky palaeoshore by bioerosion trace fossils from the Bozcaada Island, Çanakkale,Turkey

Bioerosion is a common process in hard substrates. This study introduces an example from the rocky palaeoshore cropping out at a sea cliff on the Bozcaada Island. It includes bioerosion trace fossils preserved in limestone boulders of the shallow marine and lacustrine Alcitepe Formation of Late Miocene age. The ichnotaxa include borings produced by duraphagous drillers (Oichnus isp.), phonorids (cf. Conchotrema isp.), clionid sponges (Entobia cf. goniodes, Entobia geometrica, Entobia laquea, Entobia ovula, E. cf. solaris, Entobia isp.), endolithic bivalves (Gastrochaenolites torpedo, Gastrochaenolites lapidicus, Gastrochaenolites isp., Phrixichnus isp.), polychaete annelids (Maeandropolydora isp., Maeandropolydora sulcans, Maeandropolydora decipiens, Caulostrepsis taeniola, Caulostrepsis isp.), echinoids (cf. Circolites isp.) and spinculid worms (cf. Trypanites isp.). Barnacles are also common as encrusters. The borings can be ascribed to the Gastrochaenolites-Entobia assemblage, which is typical of Neogene rocky-shores. They belong to the Entobia ichnofacies indicating various conditions of light, energy, and depth. Therefore they can reveal environmental changes and play an important role in forming palaeo-rocky shores and wave-cut platforms during marine trangressive events.     

Stable deep-sea macrobenthic trace maker associations in disturbed environments from the Eocene Lefkara Formation,Cyprus

Ichnological analysis of Eocene deep-pelagic whitish chalky calcilutites interstratified with high-energy calcarenite beds in the lower part of the Petra Tou Romiou section (southern Cyprus) was conducted to interpret the development and evolution of the trace maker associations during calcilutite deposition after high-energy episodes. The trace fossil assemblage from the chalky calcilutites consists of Chondrites isp. (Chondrites intricatus and Chondrites targionii), Planolites isp., Taenidium isp., Thalassinoides isp., and Zoophycos isp., typical of the Zoophycos ichnofacies. A composite chalky ichnofabric reveals a multi-tiered association of burrowing animals: the uppermost tier determines a mottled background, the upper tier shows the highest trace fossil abundance and diversity (Planolites, Taenidium and Thalassinoides), the middle tier features Zoophycos and large Chondrites, and the deepest tier consists mainly/exclusively of small Chondrites. This ichnofabric has an autocomposite character, associated with bioturbation by a single ichnocoenosis and gradual upward migration of the tiered macrobenthic community as the pelagic calcilutite sedimentation slowly progresses. There are no changes in the trace fossil assemblage between or within calcilutite intervals, regardless of the associated calcarenite beds. This supports a stable, mature, background calcilutite trace maker association, which recovers shortly after the deposition of high-energy calcarenites. In turn, there would have been a rapid re-establishment of paleoenvironmental conditions during pelagic calcilutite accumulation after episodic deposition of any calcarenitic material.     

Taphonomic signatures on modern molluscs and corals from Red Sea coast,southern Saudi Arabia

Bioeroded gastropod, bivalve and coral specimens (n = 570) were collected from the Jazan area, Saudi Red Sea coast, from which 22 ichnospecies of 8 ichnogenera were identified and illustrated. These traces were produced by clionid sponges (Entobia geometrica, E. ovula, E. cf. goniodes, E. cretacea, E. laquea, E. cf. paradoxa and E. isp.), duraphagous drillers (Oichnus ovalis, O. paraboloides, O. simplex and O. isp.), endolithic bivalves (Gastrochaenolites cf. dijugus, G. lapidicus, G. torpedo and G. isp.), polychaete annelids (Caulostrepsis taeniola, C. isp., Maeandropolydora sulcans, M. isp. and ?Trypanites isp.), acorn barnacles (Rogerella isp.), and vermetid gastropods (Renichnus isp.). The seashells act as hard substrate for colonization by serpulid worm, bivalves, bryozoans, and barnacles. Ichnogenus Entobia was most abundant (56.1%), followed by Gastrochaenolites (25.4%), Caulostrepsis (5.3%), Trypanites (4.2%), Maeandropolydora (3.2%), Oichnus (2.8%), Renichnus (1.9%), and Rogerella (1.0%). Oichnus occurred on the thin-shelled and smooth molluscs, while most Gastrochaenolites borings were found in the larger and thicker seashells as a suitable substrate for the settlement of polychaetes, lithophages, naticids, mytilids, and vermetids. Presence of annelid traces among radial ribs and at the siphonal areas of bivalves is indicative of nutrient capturing from water flow during the lifetime of these bivalves, within a shallow, high energy marine environment, where disarticulation, fragmentation, and abrasion of the seashells were abundantly observed.     

The sclerobionts of the Bajocian Oolithe ferrugineuse de Bayeux Formation from Calvados (Paris Basin,Normandy, France)

Silicified fossils collected in ploughed fields at Gavrus (Calvados, France), mainly mollusc shells, are Bajocian in age, and come from the Oolithe ferrugineuse de Bayeux Formation. The entire formation is highly condensed and most fossils are reworked. Their silicification allowed treatment with dilute hydrochloric acid. This treatment brought to light numerous encrusting sclerobionts: Porifera (3 taxa), Bryozoa (n), Polychaeta (9), Brachiopoda (n), Bivalvia (5). The borings and bioerosional traces are described using the “categories of architectural design” as defined by Buatois et al. (2017). Among the 28 ichnotaxa described (corresponding to an ichnodisparity of 14), Planavolites wisshaki isp. nov., Kleithrichnus belemnophilus igen. nov., isp. nov. and Foggara foggara igen. nov., isp. nov. are new. Some Gastrochaenolites dijugus Kelly and Bromley, 1984 borings accommodate the shell of the presumed borer Lithophaga fabella J.-A. Eudes-Deslongchamps, 1838. A group of Nododendrina europaea (Fischer, 1875) on a belemnite rostrum provides a good instance of an ichnogenetic series. Encrusting sclerobionts and macroscopic boring and bioerosional trace-makers are dominantly suspension feeders. The ichnofacies is a peculiar Entobia-ichnofacies, found in deeper, lower-energy environments than the “classical” coastal Entobia-ichnofacies, on deep shell-grounds far from the coast, in the deep euphotic zone.     

Three New Ichnogenera of Biting and Gnawing Traces on Reptilian and Mammalian Bones: A Case Study from the Miocene of the Czech Republic

The Ahníkov (Miocene, Czech Republic) site represents a concentration of vertebrate skeletal remains in a swamp setting. Autochthonous bone deposits were strongly altered by sedimentary processes and early diagenesis. Biting and gnawing traces recognized on hard animal tissues (bones, teeth, antlers, turtle thoraces) represent seven recurring morphotypes. The following ichnotaxa are erected: Nihilichnus nihilicus n. igen. et n. isp., Nihilichnus mortalis n. isp., Machichnus regularis n. igen. et n. isp., Machichnus multilineatus n. isp., Machichnus bohemicus n. isp., and Brutalichnus brutalis n. igen. et n. isp. Each kind of bones or similar substrates bears a specific proportion of various bite traces but no observed morphotype is specific for a single substrate. Ethologically, traces of sharpening of teeth are principally different from predation traces. The beavers Steneofiber eseri and Steneofiber depereti, the carnivore Amphicyon sp. and the crocodiles are presumed as the tracemakers.     

Early cretaceous bird-dominated and dinosaur footprint assemblages from the northwestern margin of the Junggar Basin, Xinjiang, China

Here we describe a number of mostly isolated bird and non-avian dinosaur footprints from the Huangyangquan track site in the Lower Cretaceous Lower Layer of the Tugulu Group near Huangyangquan Reservoir in the Wuerhe District, Karamay, Xinjiang Uyghur Autonomous Region, China. Bird footprints at the site pertain to Koreanaornis dodsoni n. ichnosp., Goseongornipes isp., Aquatilavipes isp., and Moguiornipes robusta n. ichnogen. n. ichnosp. Moguiornipes tracks may be the products of a bird with lobed feet. These shorebird-morph tracks are the first solid evidence of birds in the Tugulu Group, and greatly enrich the known faunal diversity of the Lower Layer of the Tugulu Group. The Huangyangquan avian ichnofauna is comparable to those of the Jindong Formation, Korea and the Dakota Formation, USA. Non-avian dinosaur footprints at the site pertain to cf. Jialingpus isp., Asianopodus isp., and Kayentapus isp. This is the first report of Jialingpus from Lower Cretaceous strata. The discoveries of Asianopodus isp. and Kayentapus isp. increase the known ranges of these two ichnogenera.     

A Lower Cambrian shallow-water occurrence of the branching ‘deep-water’ type trace fossilDendrorhaphe from the Lontova Formation, eastern Latvia

We describe a unique Lower Cambrian (Lontova Formation, Latvia) occurrence of a branching trace fossil assigned toDendrorhaphe isp. In the Mesozoic and Cenozoic this type occurs exclusively in deep-water flysch deposits.Dendrorhaphe may, therefore, be an additional example of a flysch-type trace fossil with an origin in early Palaeozoic shallow-water environments and subsequent onshore-offshore displacement, or perhaps, expansion. Recently,Dendrorhaphe Seilacher 1977, has been synonymized withDendrotichnium Farrés 1967 andUbinia Grosshejm 1961. This needs further consideration, especially because of possible relation of these ichnogenera toChondrites patulus Fischer-Ooster 1858.     

Upper Pliocene bivalve shell concentrations from the Lower Chelif basin (NW Algeria): Systematics,sedimentologic and taphonomic framework

Sedimentologic and palaeontological investigation of the Upper Pliocene Slama Formation in the Lower Chelif Basin (NW Algeria) led us to collect important bivalve assemblages for taxonomic and taphonomic purposes. A rather comprehensive inventory list of Upper Pliocene bivalves from northwestern Algeria is now available and consists of 30 species, 17 of which are extinct ones. Four principal taphonomic attributes were analysed: bioerosion, encrustation, fragmentation, and abrasion. Physical and biogenic sedimentary structures are used for palaeoenvironmental interpretations. The taphonomic, sedimentologic and ichnological characteristics of most of the deposits suggests they originated from discontinuous processes of winnowing and bypassing of sediments, probably due to the action of storms in shallow waters, mainly in the shoreface depositional environment. The bivalve assemblage is dominated by disarticulated valves and displays significant taphonomic alteration in the shells. Sclerobionts traces in shells particulary affect the oyster shells. Bioerosion traces are predominately those of clionid sponges (Entobia isp.), polychaetes (Maeandropolydora isp. and Caulostrepsis isp.), bivalves (Gastrochaenolites isp.), and of predatory gastropods (Oichnus isp.). Among the sclerobionts, the identified encrusters were juvenile oyster recruits, barnacles, polychaetes (serpulid tubeworms), bryozoans (Microporella sp. and Acanthodesia sp.), and vermetid gastropods (Petaloconchus intortus).     

Crustacean microcoprolites from Jurassic (Oxfordian) hydrocarbon-seep deposits of Beauvoisin,southeastern France

Decapod crustaceans are the most conspicuous predators and scavengers of modern chemosynthesis-based ecosystems at hydrothermal vents and hydrocarbon seeps; however, very little is known about decapod crustaceans at ancient seeps or vents. Some decapods, including galatheids, are known to produce highly specific, internally structured microcoprolites. Late Jurassic (Oxfordian) hydrocarbon-seep limestones from Beauvoisin (southeastern France) were found to contain such microcoprolites including Favreina fontana n. isp., F. cf. martellensis Brönnimann and Zaninetti, Palaxius salataensis Brönnimann, Cross and Zaninetti and Palaxius isp. These diverse ichnofossils indicate that crustaceans were also a part of ancient chemosynthesis-based ecosystems. This is the first report of crustacean coprolites from a deep-water environment.     

山西兴县中奥陶统马家沟组五5亚段遗迹化石组合及其古环境意义

鄂尔多斯盆地东缘关家崖剖面奥陶系马家沟组五5亚段地层中保存大量无脊椎动物遗迹化石,可识别出居住迹、牧食迹、觅食迹三大类,共7个遗迹属、14个遗迹种,分别是Arenicolites isp.,Cylindricumisp.,Helminthopsis abeli,Helminthopsis isp.1,Helminthopsis isp.2,Lorenziniaisp.,Planolites isp.,Planolites beverleyensis,Planolites montanus,Teichichnus rectus,Thalassinoides isp.1,Thalassinoides isp.2,Thalassinoides isp.3,Thalassinoides isp.4。根据遗迹化石特征及其沉积环境,可划分3个遗迹组合:1、Helminthopsis-Planolites组合,该组合以觅食迹为主,部分为居住迹,出现在潮间带和潮下带;2、Thalassinoides-Teichichnus组合,该组合遗迹化石种类较单一,主要为觅食迹,分布于潮间带和潮上带;3、Thalassinoides-Helminthopsis组合,该组合出现在潮间带。基于遗迹化石组合的分布特点,提出山西兴县奥陶系马家沟组五5亚段的沉积环境模式。     

An Updated Review of the Avian Footprint Record from the Yacoraite Formation (Maastrichtian-Danian), Northwestern Argentina

Abstract

The Yacoraite Formation (Maastrichtian–Danian; Balbuena Subgroup; Salta Group), from Northwestern Argentina, represents a shallow epeiric unit which is the result of transgressions in the Andean basin of South America. Herein, we study the avian footprints from the Maimará locality, Jujuy province, and Quebrada del Tapón ichnosite, Salta province. The avian footprints from the Maimará locality is less diverse, made up: cf. Alaripeda isp., Avipeda isp., cf. Gruipeda filiportatis, and cf. Gruipeda isp. The avian track record from the Quebrada del Tapón ichnosite is composed of: cf. Alaripeda isp., Ardeipeda cf. egretta., Gruipeda filiportatis (=Yacoraitichnus avis) and others Gruipeda or compared to this ichnogenus, cf. Uhangrichnus isp., and several indetermined avian footprints. Both ichnological assemblages have tracks assigned to shorebirds or Charadriiformes. The facies distribution of trace fossils reinforces the overall model of a shorebird tracks dominance in moderate to low energy settings. This distribution can be divided into three sub-environments: (1) a moderate to high energy shoreline under wave action, (2) a supratidal body of ephemeral ponds, far away from direct wave influence and (3) a landward position, beyond the ephemeral ponds system. We have included all the ichnoassemblages within the shorebird ichnosubfacies as a subset of the Scoyenia Ichnofacies.