Caedichnus,a New Ichnogenus Representing Predatory Attack on the Gastropod Shell Aperture

Predatory traces, in which the tracemaker has damaged the prey animal's skeleton to kill and consume it, have a deep fossil history and have received much scientific attention. Several types of predatory traces have been assigned to ichnotaxa, but one of the most studied predatory traces, the wedge-shaped excision produced as a result of attacks mainly by crustaceans on the apertures of gastropod shells, has yet to be described as an ichnotaxon. We propose the ichnogenus Caedichnus to describe the shell damage produced by aperture peeling behavior. Caedichnus is produced by predators that are unable to crush their prey's shells outright. Depending on the predator's peeling ability and the prey's withdrawal depth within the shell, the trace can extend through several whorls of the shell. Aperture peel attacks may fail, allowing such damage to be repaired by surviving gastropods. Thus, the types of attacks that produce Caedichnus may exert selective pressure on prey to evolve better-defended shells (in the case of gastropods) or to inhabit better-defended shells (in the case of hermit crabs). The identification of these trace fossils will enhance our understanding of how predation influences the morphological, and even behavioral, evolution of prey organisms.     

Microendoliths in Lower Pliocene Oysters from the Alt Empordà Basin,NW Mediterranean: Paleoenvironmental Inferences

The study of traces made by microendoliths on Neopycnodonte cochlear shells from a coquina in the Vilacolum area (Alt Empordà basin) has permitted us to identify for the first time 18 ichnotaxa in the Lower Pliocene sediments of the NW Mediterranean. A shallow, euphotic environment with local shaded zones (corresponding to deep shallow conditions) is proposed for this shell bed, owing to the presence of abundant heterotroph microborings and rare cyanobacterial traces as well as to the high number of Rhopalia catenata and Ichnoreticulina elegans. This refines a former interpretation, mainly based on body fossils and macroborings, which suggested simply a shallow depositional environment. The ichnospecies Ichnoreticulina elegans, Rhopalia catenata, ?R. clavigera, Saccomorpha clava, S. terminalis, Polyactina araneola, Entobia mikra, Orthogonum lineare and Aurimorpha varia are identified for the first time in marine Pliocene materials, thus furnishing a more complete picture of the temporal span of some microborers.     

The ichnofamily Celliformidae for Celliforma and allied ichnogenera

This contribution undertakes a comprehensive revision, lacking until now, of all the ichnotaxa attributed to fossil bee cells, one of the most common traces in paleosols. These ichnotaxa are morphologically related to Celliforma and consequently grouped herein in the new ichnofamily Celliformidae. Two new ichnogenera are formulated, Cellical‐ichnns igen. nov., for several ichnospecies previously included in Celliforma, and a new ichnospecies, C. chubutensis isp. nov., belonging to this ichnofamily, and Brownichnus igen. nov., for the ichnospecies favosites, which is not morphologically related to Celliforma. The ichnogenera included in this ichnofamily involve traces in which Celliforma, the most simple trace of the group, is part of their structure. Paltniraichnus is akin to Celliforma with antechambers and discrete walls. Uruguay, Corim‐batichnus and Rosellichnus are clusters composed of adjacent rows of Celliforma, or Palmiraichnus‐like cells. Ellipsoideichnus and Cellicalichnus are different arrangements of Celliforma‐like cells attached to tunnels.

Celliformidae are based on the morphology of the traces, all of them comprising cells, groups of cells and cells attached to tunnels. As such, its component ichnotaxa are based exclusively on morphological ichnotaxobases, which, in turn, were evaluated and selected with respect to the nest architecture of the trace makers, the bees. This paper analyzes some procedures and clues (i.e. behavioral homologies) that may be used to select the proper taxobases to erect ichnotaxa when the identity of the trace makers is known, as in this study case.     

The distribution of microborings in molluscan shells from recent reef environments at Lee Stocking Island,Bahamas

Summary The distribution of microbial borings in bivalve shells was assessed between five research sites in windward and leeward environments at the Lee Stocking Island, Bahamas. The research sites are on windward coral reefs (sites B at 2 m, Fat 12 m, and C at 30 m), a tidal channel stromatolite reef (site A at 5 m), and a leeward hard ground reef (site D at 3 m). A total of 22 ichnotaxa have been recognized within 100 samples. Each site contained between 14 and 17 ichnotaxa. Although the diversity of microborings, expressed both as number of taxa and number of individuals per taxon, is similar between sites, the following ranking (in descending order) was found: B-F-A-C-D.Rhopalia catenata was found to dominate at site D, whileReticulina elegans dominated at site C, andFasciculus dactylus dominated at site F. The results are in accordance with the known bathymetric distribution of the boring microorganisms.     

Bite traces on dicynodont bones and the early evolution of large terrestrial predators

Nied?wiedzki, G., Gorzelak, P. & Sulej, T. 2010: Bite traces on dicynodont bones and the early evolution of large terrestrial predators. Lethaia, Vol. 44, pp. 87–92. Dicynodont (Synapsida: Anomodontia) bones from the Late Triassic (late Norian/early Rhaetian) of Poland yield characteristic tooth marks that can be attributed to three ichnotaxa (Linichnus serratus, Knethichnus parallelum and Nihilichnus nihilicus). The general shape and dimension of these traces perfectly match the dental morphology of a co‐occurring carnivorous dinosaur. It is therefore concluded that early carnivorous dinosaurs were feeding on dicynodonts. This discovery constitutes one of the oldest evidence of dinosaur predator–prey interaction. It is suggested that an evolutionary increase in the size of dicynodonts across the Late Triassic may have been driven by selection pressure to reach a size refuge from early dinosaur predators. □Bite traces, dicynodonts, dinosaurs, predation, Triassic.     

Bioerosion along a bathymetric gradient in a cold-temperate setting (Kosterfjord, SW Sweden): an experimental study

In the cold-temperate setting of the Swedish Kosterfjord, a 2-year experiment was launched in order to assess bioerosion rates and to investigate the endolithic borer communities in relation to light availability (relative bathymetry), hydrography and exposure time. The inventory of microendolithic traces, studied by SEM analysis of epoxy resin casts of planted bivalve shells, yields diverse ichnocoenoses comprising a total of 21 traces produced by boring cyanobacteria (7), chlorophytes (4), fungi (6) and traces of uncertain affinity (4). The link between the endoliths (biotaxa) and the traces they leave (ichnotaxa) is evaluated by the study of the boring organisms in situ by transmission light microscopy of planted Iceland spar and bivalve shells. Additionally, the activity of various macroborers (foraminiferans, polychaetes, echinoids, gastropods and sponges) is documented, adding to a distinct diversity maximum at 7 m water depth. A highly condensed photic zonation, due to the high latitude (59°) and eutrophic conditions, is recorded by the measurement of the Photosynthetically Active Radiation (PAR) and is confirmed by the bathymetric range of the photic related ichnocoenoses. At 1 m water depth, a mature shallow euphotic ichnocoenosis dominated by cyanobacteria and at 7 m, a deep euphotic ichnocoenosis dominated by chlorophytes, respectively, is developed after as little as 12 months exposure. With the vanishing light availability from 15 m downwards, the ichnocoenoses development is significantly slowed and only immature dysphotic and aphotic borer communities (dominated by fungi) are encountered. Strong fluctuations of salinity (down to 8%) and temperature (0–20°C) in the euphotic zone indicate most phototrophs present to be considerably euryhaline and eurytherm, while most endolithic fungi appear preferentially in the deeper, more stable marine waters.     

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.     

Middle Pennsylvanian Ichnofauna from Eastern Oklahoma,USA

The Keota site, in the Middle Pennsylvanian (Desmoinesian) Keota Sandstone Member, McAlester Formation (Krebs Group) of Haskell County, Oklahoma, yields an extensive ichnofossil assemblage of arthropod trackways, insect resting traces, arthropod feeding and grazing traces, fish swimming traces, and tetrapod footprints. This ichnofossil assemblage occurs in a 0.8 to 1.5-m-thick unit of thinly laminated and ripple-laminated sandstone at the base of the Keota Sandstone Member that is interpreted as a tidal flat sandstone. The arthropod traces are assigned to the ichnotaxa Diplichnites gouldi Gevers types A and B, Diplopodichnus biformis Brady, Paleohelcura tridactyla Gilmore, Tonganoxichnus buildexensis Mángano, Buatois, Maples and Lanier, Gordia marina Emmons, cf. Cochlichnus sp. and Treptichnus bifurcus Miller. The fish swimming traces are assigned to Undichna britannica Higgs, and the tetrapod tracks to Notalacerta ichnosp. and Pseudobradypus ichnosp. The Keota ichnofossil assemblage thus documents the first North American Pennsylvanian record of Paleohelcura tridactyla and the first record of Pseudobradypus from the western United States. This ichnofossil assemblage also provides compelling evidence of a freshwater habitat in a tidal flat setting during deposition of the fossil-bearing sandstone at the Keota site. Indeed, the Keota ichnofossil assemblage is characteristic of a Carboniferous-Permian Tonganoxichnus assemblage that identifies nonmarine tidal flat settings.     

Deux nouveaux sites à traces de pas dinosauroïdes dans la Formation Dolomitique de l’Hettangien de Lozère,Languedoc-Roussillon,France

Recent ichnological studies revealed two new localities showing dinosaur footprints in the Hettangian Dolomitic Formation from the Causses Basin, southern France. The traces are reported from the northern part of the basin, in Lozère, an area where Lower Jurassic dinosaur ichnites were poorly documented. The surfaces bearing footprints are characterized by a large number of traces. The ichnites are ascribed to Grallator (Hitchcock, 1858), Dilophosauripus (Welles, 1971) and Eubrontes (Hitchcock, 1845) whose trackmakers were theropods. Although numerous tracksites were previously described in the southern part of the Causses Basin, these three ichnotaxa are for the first time observed in association into the Dolomitic Formation.     

Vagorichnus,a new ichnogenus for feeding burrow systems and its occurrence as discrete and compound ichnotaxa in Jurassic lacustrine turbidites of Central China

A new ichnogenus, Vagorichnus, is figured and described from Jurassic lacustrine turbidites of the Anyao Formation, Henan Province, Central China. Vagorichnus consists of complex burrow systems composed of discontinuous curved to meandering segments in which irregular networks are formed. The ichnospecies V. anyao covers burrow systems characterized by constricted and irregularly annulated segments, and ridge‐like knobs. Although traces are preserved as hypichnial ridges on the sole of turbidite sandstones, several lines of evidence indicate that these ichnofossils record the activity of a post‐event infauna. Vagorichnus occurs as discrete and compound ichnotaxa. In the latter case, it grades into Gordia and Tuberculichnus. Vagorichnus is interpreted as invertebrate feeding structures (Fodinichnia).     

Bioerosion of oyster shells in brackish modern mangrove swamps,Nigeria

Studies on shells of the intertidal oyster Crassostrea tulipa cemented to aerial prop roots and stems of the mangroves Rhizo‐phora racemosa, R. harrisonii, and R. mangle from the Cross River and Qua Iboe River estuaries show that photosynthetic en‐dolithic cyanobacteria (blue‐greens) are the major bioerosional agent, affecting about 94% of the shells examined. Pblychaetes attacked less than 10% of these materials and thus have low bioerosional impact.

Herbivorous gastropods play a secondary role because, by grazing, they clear the shell surface of encrusting cyanobacteria and thus enhance the activity of boring forms. Where epilithic cyanobacteria have been removed, the gastropods sometimes leave faint grazing traces of low fossilization potential.

Microfloral boring activity is high at all stations, but the assemblage is of markedly low diversity compared with those of littoral and shallow sublittoral marine communities and may thus be useful as a paleoenvironmental tool. Bioerosional agents are directly or indirectly responsible for the disintegration of oyster shells, whose fragments are incorporated in muddy intertidal sediments.     

Belichnus Traces Produced on Shells of the Bivalve Lutraria lutraria by Gulls

In February 2011, after a storm, thousands of adult, articulated, and still-living common otter shells Lutraria lutraria (L. 1758) were stranded on the North Sea beach of the Island Texel (NL). These 9 to12 cm long bivalves were rapidly found and consumed by both herring- and lesser black-backed gulls. Holes, irregular in outline, were observed in some 10% of the articulated shells of these bivalves. These holes were always smaller on the outside of the valves than on the inside and varied in size from 1 to 20 mm (outside) to 4 to 22 mm (inside). Often the other valve was crushed indicating consumption by gulls. We concluded that these holes were made by the gulls probing the shells; in a few cases, we observed that valves were broken starting from such a hole. Such traces are described in the literature as the ichnogenus Belichnus and were until now attributed to Stomatopoda only. We also suggest that a separate ichnospecies name for two Belichnus holes in one shell should not be used, as we see them simply as a double injury due to two blows. Our findings stress once more the importance of avoiding premature phylogenetic interpretation of traces and the use of a separate ichnotaxonomy.     

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.     

Borings in mobile clasts from Eocene conglomerates of Northern Dalmatia (Coastal Dinarides, Croatia)

Summary Bored clasts occur in Eocene conglomerates deposited in the upper shoreface and beachface settings of the Dinaric foreland basin. The trace fossil assemblage consists ofGastrochaenolites, Trypanites, and possibly some other ichnotaxa and may be compared to theTrypanites Ichnofacies. The preservation characteristics of the borings reflect many stages of colonisation/boring and abrasion. The removal of shells of the boring bivalves, the different depths of the abrasional truncation of borings, and the predominant preservation of the largest excavations (Gastrochaenolites) in the ichnocoenosis are related to repeated phases of abrasion, caused by the mobility of clasts. Coastal gravel is a specific variant of hard substrates, whose mobility controls the colonisation of borers, the type of assemblage and its preservation potential.     

Bioerosional structures and pseudoborings from Late Jurassic and Late Cretaceous-Paleocene shallow-water carbonates (Northern Calcareous Alps, Austria and SE France) with special reference to cryptobiotic foraminifera

Examples of bioerosional processes (boring patterns) are described from shallow-water limestones of the Late Jurassic Plassen Carbonate Platform (PCP) and the Late Cretaceous to Paleocene Gosau Group of the Northern Calcareous Alps, Austria. Some micro-/macro-borings can be related to distinct ichnotaxa, others are classified in open nomenclature. In the Alpine Late Jurassic, bioerosional structures recorded from clasts in mass-flows allow palaeogeographical conclusions concerning the source areas. In particular, these are borings of the Trypanites-ichnofacies detected from clasts (Barmstein limestones) of the PCP or special type of bored ooids of unknown source areas or restricted autochthonous occurrences. In the Lower Gosau Subgroup, Gastrochaenolites macroborings occur in mobile carbonate clast substrates of shore zone deposits (“Untersberg Marmor”). Different types of borings are recorded from rudist shells and coral skeleton, some of which are referable to the ichnotaxon Entobia produced by endolithic sponges. In the present study, special attention is paid to the occurrences of the cryptobiotic foraminifera Troglotella incrustans Wernli and Fookes in the Late Jurassic and Tauchella endolithica Cherchi and Schroeder in the Late Cretaceous. The latter is so far only known to be from the Early Cenomanian of France and is reported here for the first time from the Late Turonian-Early Coniacian stratigraphic interval where it was found in turbulent carbonate deposits within borings penetrating bivalve shells or coralline algae. The records of cryptobiotic foraminifera from the Northern Calcareous Alps are supplemented by a single finding from the Middle Cenomanian of SE France. A palaeoenvironmental interpretation of the occurrences of the cryptobiotic foraminifera is provided.     

Review of Japanese Dinosaur Track Localities: Implications for Ichnotaxonomy,Paleogeography and Stratigraphic Correlation

Nine dinosaur ichnospecies from the Lower Jurassic to Upper Cretaceous of Japan, including two that are new, are described herein. The new ichnotaxa are Asianopodus pulvinicalx ichnogen. et ichnosp. nov. and Schizograllator otariensis ichnosp. nov. The Japanese ichnotaxa are allied to Lower Jurassic ichnospecies in South China, North America, Western Europe and South Africa, and Upper Jurassic to Lower Cretaceous ichnospecies from Southeast and East Asia. This suggests they were part of a global ichnofauna before continental drift began in the Middle Jurassic, leading to the development of a more endemic dinosaur fauna in the Cretaceous. At least two assemblages, an ornithopod-gracile-toed theropod-dominated community, in northeastern Asia, and a robust theropod- and sauropod-dominated community in the southern part of the continent, existed in the Cretaceous. This parallels North American dinosaur distribution patterns in the Cretaceous and seems to be a reflection of paleolatitudinal controls.     

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.     

Grallator-Dominated Fossil Footprint Assemblages and Associated Enigmatic Footprints from the Chinle Group (Upper Triassic), Gateway Area,Colorado

The upper part of the Chinle Group (Late Triassic) of the Gateway area in western Colorado is extraordinarily rich in fossil footprint assemblages. Dominant track types include small Grallator tracks, generally attributed to Coelophysis-like theropods, which often occur in high densities of 50 to 100 per m². Other abundant ichnotaxa that are attributable to dinosaurs or dinosaur-like archosaurs include Pseudotetrasauropus and Tetrasauropus, attributed to prosauropods and sauropods, respectively. Several Pseudotetrasauropus-like tracks appear to be functionally didactyl and may indicate a new ichnotaxon that represents an animal that shows certain unusual features that are convergent with dromeosaurs and certain birds. Such convergence may reflect inherent growth programs as much as functional adaptations. Non-dinosaurian ichnotaxa include Brachychirotherium (probably of aetosaur affinity) and Rhynchosauroides, attributed to a sphenodontid/lizard-like form. Other ichnotaxa include probable therapsid (dicynodont) tracks labeled Pentasauropus sp., mammaloid (non-therian mammal and/or mammal-like reptile) tracks, and the trails of arthropods. Excellent preservation and high track densities mark the Gateway assemblages in a thin stratigraphic interval in the upper part of the Chinle Group (Rock Point Formation). The track assemblages are similar to those reported from the Chinle Group in other parts of the Colorado Plateau and Rocky Mountain region, extending over most of Colorado, Utah, northern Arizona and northern and eastern New Mexico. Some of the Chinle ichnotaxa (Grallator and Brachychirotherium) are found in the overlying Wingate Formation, indicating that it is also Late Triassic in age, at least in the lower part. However, overall the Chinle and Wingate assemblages are quite different, most notably in the rarity of mammaloid/mammal-like tracks in the Chinle Group.     

New Hettangian tracks from the Causses Basin (Lozère,southern France) complement the poor fossil record of earliest Jurassic crocodylomorphs in Europe

A new vertebrate ichnological assemblage is described from the Hettangian Dolomitic Formation of the Causses Basin, at Le Serre (Lozère, southern France). We report tracks that complement the poor fossil record of lowermost Jurassic crocodylomorphs in Europe. Tetradactyl pes and pentadactyl manus imprints form a unique trackway. Traces are ascribed to Batrachopus isp. and they represent the third occurrence of this ichnogenus in European Hettangian geological formations. This is the first report of Batrachopus in Hettangian deposits of the Causses Basin. The tracks share some similarities with Batrachopus deweyi previously described from the Sinemurian of the Causses Basin. Crocodylomorph tracks co-occur with tridactyl dinosaur traces assigned to Dilophosauripus williamsi and Grallator isp. The sedimentology and palaeoichnology of the tracksite indicate that the depositional environment was a tidal to supratidal flat marsh that was emerged periodically. The ichnological assemblage from Le Serre and a synthesis of ichnotaxa co-occurring with Batrachopus in the European lowermost Jurassic tracksites confirm that crocodylomorphs living in marginal-littoral palaeoenvironments were part of theropod-dominated faunas, together with ornithopods but apparently without sauropods.