Evidence of Bioerosive Structures in Quaternary Glaciomarine Sediments from Southwestern Iceland

The Late Weichselian marine and glaciomarine sediments occurring in many places along the coast of Iceland are rich in invertebrate fossils, especially mollusks and barnacles. A diverse ichnofauna, constituted especially by bioerosional traces documenting the activity of predators and animals that use the host shell for attachment or the construction of dwellings, was identified at the localities of Ósmelur, Saurbær, and Brekkubakkar in southwestern Iceland. The ichnotaxa identified include Anellusichnus circularis (Santos, Mayoral, and Muñiz), Caulostrepsis isp., Centrichnus concentricus (Bromley and Martinell), Clionolithes isp., Finichnus peristroma (Taylor, Wilson, and Bromley), Finichnus isp., Sedilichnus asperus (Nielsen and Nielsen), Sedilichnus cf. excavatus (Donovan and Jagt), Sedilichnus gradatus (Nielsen and Nielsen), Sedilichnus smiley isp. nov., Sedilichnus ovalis (Bromley), Sedilichnus paraboloides (Bromley), Sedilichnus simplex (Bromley), and Sedilichnus spongiophilus (Müller).     

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.     

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.     

Shell microstructure of the early bivalve Pojetaia and the independent origin of nacre within the mollusca

Abstract: Pojetaia and Fordilla are the oldest bivalve molluscs, occurring in roughly co‐eval rocks from the Tommotian, and are the only undisputed, well‐known bivalves from the Cambrian. New specimens reveal that Pojetaia had a laminar inner shell microstructure reminiscent of the foliated aragonite of modern monoplacophorans, and the same is true for Fordilla. A similar shell microstructure is seen in Anabarella and Watsonella, providing support for the hypothesis that they are the ancestors of bivalves. Foliated aragonite shares many similarities with nacre, and it may have been the precursor to nacre in bivalves. No cases of undisputed nacre occur in the Cambrian, in spite of much shell microstructure data from molluscs of this time period. Thus, although considered by many to be homologous among molluscs, we conclude that nacre convergently evolved in monoplacophorans, gastropods, bivalves, and cephalopods. This independent origin of nacre appears to have taken place during, or just prior to, the Great Ordovician Biodiversification Event and represents a significant step in the arms race between predators and molluscan prey.     

Lebensspuren von Meersauriern und ihren Beutetieren im mittleren Jura (Callovien) von Liesberg, Schweiz

Summary Numerous gutter-like furrows, up to 60 cm wide and up to 9 m long are preserved at the interface “Macrocephalus Beds”/“Callovian Marl” over a surface of 20 by 200 m. They are interpreted as feeding traces made by large marine vertebrates, most likely plesiosaurs and ichthyosaurs searching for food in the lime mud of the shallow Middle Jurassic sea floor. Possible prey animals were infaunal invertebrates (crustaceans) which produced an intricate meshwork of burrows (mainlyRhizocorallium irregulare andThalassinoides) in the bottom sediments, as well as infaunal bivalves. Evidence from cololites of predatory pelagic reptiles (ichthyosaurs, plesiosaurs) as well as reptile regurgitalites indicate that these animals fed not only on fast-swimming vertebrates and cephalopods but also on epi- and endobenthic invertebrates. In addition, the cololites show that the predators ingested considerable amounts of bottom sediment. Different sizes and shapes of the traces suggest that the gutters were produced by different reptiles or age groups. Candidates for the widest gutters are pliosaurs. Of the marine vertebrates known from Jurassic time, only the snout of adult pliosaurs of the genusLiopleurodon was broad enough to produce gutters more than 40 cm wide. Smaller, less than 15 cm wide gutters, could have been made by plesiosauroids or by the narrow pointed snouts of ichthyosaurs. Almost identical traces described from the Oxfordian of Spain and similar but smaller traces from the Lower Devonian of Prague are equally interpreted as feeding traces on the sea floor. Feeding traces of vertebrates in bottom sediments may give detailed information on the hunting behaviour of the predators. However, the attribution of the traces to definite vertebrate taxa remains uncertain.

     

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.     

The Trace Fossil Diplopodichnus from the Lower Jurassic Lacustrine Sediments of Central China and the Isopod Armadillidium vulgare (Pillbug) Lebensspuren as its Recent Analogue

The trace fossil Diplopodichnus isp. was identified in lacustrine siliciclastics of the Lower Jurassic Anyao Formation in the Henan Province, central China. This is the youngest occurrence of this ichnogenus. It is similar to some variants of recent surface traces of the isopod Armadillidium vulgare (pillbug) crawling on soft mud in temporal puddles. Therefore, isopods are suggested to be producers of the described Diplopodichnus isp. It seems that isopods should be taken into account as important component of deep lacustrine ecosystems by the Early Jurassic.     

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.     

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.     

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.     

Ichnofacies and microbial build-ups on Late Miocene rocky shores from Menorca (Balearic Islands), Spain

Angular unconformites between Jurassic and Miocene strata are exposed in the sea cliffs of Cala Cigonya on the northwest coast of Menorca in the Balearic Islands of Spain. The geological discontinuities represent rocky shores on opposite sides of a former headland with 15 m of topographic relief. On the east flank, Jurassic dolomite is overlain by Upper Miocene (Tortonian to Messinian) breccia and laminated limestone. Here, a partially exhumed dolomite surface records Miocene bivalve borings of the ichnospecies Gastrochaenolites torpedo and G. lapidicus that achieved a density of >1,000 borings/m². Other associated traces include sponge borings (Entobia isp.) and polychaete borings (Caulostrepsis isp.). A breccia deposit 0.8 m thick was derived from the underlying dolomite and angular clasts still retain evidence of bivalve borings. Above follows a succession of laminated limestone beds more than 5 m thick, including some levels with dome-shaped stromatolites and other horizons with reworked dolomite clasts. Thin-section analysis of the laminated limestone reveals dark and light couplets 0.2 mm thick consistent with microbial origins. In contrast, the west flank was buried by coarse sandstone and laminated sediments. Here, dwelling structures of regular echinoids (Circolites kotoncensis) are the dominant traces preserved on the dolomite surface, reaching a maximum density of 66 borings/m². Associated borings include Entobia geometrica as well as rare traces of Gastrochaenolites isp. and Trypanites isp. Notable for the absence of a basal Miocene breccia, the west flank is interpreted as a sheltered rocky shore coeval with an exposed rocky shore on the east flank. Today, heavy surf on the north coast of Menorca is related to the Tramontana winds that blow out of Spain during winter months. Similar atmospheric circulation patterns must have prevailed during the Late Miocene, but the replacement of ichnofacies by microbial build-ups resulted from increased salinity during the Messinian crisis.     

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.     

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.     

Late Triassic Tetrapod-Dominated Ichnoassemblages from the Argana Basin (Western High Atlas,Morocco)

Diverse tetrapod track assemblages with Scoyenia invertebrate traces were discovered in the Triassic Timezgadiouine and Bigoudine formations of the Argana Basin (Western High Atlas, Morocco). The ichnofossils occur in alluvial plain sandstones and mudstones of the Irohalène Member (T5) and Tadart Ouadou Member (T6) considered Carnian-Norian in age by vertebrate remains and palynomorphs. Tetrapod footprints are assigned to Apatopus, Atreipus-Grallator, Eubrontes isp., Parachirotherium, cf. Parachirotherium postchirotherioides, Rhynchosauroides ispp., and Synaptichnium isp. They can be referred to lepidosauromorph/ archosauromorph, basal archosaur, and dinosauromorph trackmakers. Apatopus, represented by 11 tracks of a more than 4 m long trackway, is recorded for the first time outside of North America and Europe. The assemblage concurs with the proposed Late Triassic age of the track-bearing beds by the occurrence of Apatopus, Atreipus-Grallator, and Eubrontes. If this is accepted, the stratigraphic range of Synaptichnium and Parachirotherium, hitherto known only from Early or Middle Triassic deposits, has to be extended to the Carnian-Norian. The occurrence of Eubrontes in the Irohalene Member (T5) provides further evidence for large theropods in pre-Jurassic strata. All assemblages are referred to the Scoyenia ichnofacies indicating continental environments with alternating wet and dry conditions.     

Insights from functional morphology and neoichnology for determining tracemakers: a case study of the reconstruction of an ancient glacial arthropod‐dominated fauna

Apportioning trace fossils to producers is significant for palaeoecological and palaeoenvironmental studies. A tracemaker is usually attributed through the comparison of the morphology of the trace to the body plans of modern and extinct animals. However, hypotheses of tracemaker identifications may be tested with the use of further methods. This contribution aims to evaluate the potential tracemakers of a trackway‐dominated ichnofauna preserved in the siltstone–claystone rhythmites of the Pennsylvanian Rio do Sul Formation in the southern Brazil. These deposits are typical of the deglaciation phase of the Late Palaeozoic Ice Age of Gondwana in the Paraná Basin. Thus, the parameters applied in this work are as follows: (1) functional analyses of the traces to infer the biomechanics of the locomotion of the producer; and (2) neoichnology, whereby modern traces whose producers are known are compared to those of the ichnofauna in question. The conclusions obtained with these methods strongly support millipedes as the producers of Diplichnites gouldi and Diplopodichnus biformis. The malacostracan crustacean‐related ichnogenera are Umfolozia sinuosa and Gluckstadtella elongata, produced by syncarids or peracarids, while isopods are suggested as the tracemakers of Glaciichnium isp. and Protovirgularia dichotoma. In terms of branchiopod crustaceans, Cruziana problematica is attributed to notostracans. Helminthoidichnites tenuis, Mermia carickensis and Treptichnus pollardi are considered traces produced by insect larvae. The palaeobiology of the tracemakers that produced the ichnofauna preserved in the rhythmites of the Rio do Sul Formation permits inferences that terrestrial (millipedes) and aquatic (crustaceans and insect larvae) animals colonized the sediments. This conclusion points to a dynamic depositional environment, with changing physical and chemical parameters conditioning the biota.     

Spirodesmos milanai n. isp.: A Shallow-Water Spiral Trace Fossil from the Cambrian of the Eastern Cordillera,Northwest Argentina

Abstract

A new ichnofossiliferous locality in Salta Province (northwest Argentina) contains an association with numerous irregular spiral traces assigned to Spirodesmos milanai n. isp., in mature sandstones and quartzites with rippled bedding surfaces, rare wavy lamination and cross-bedded stratification. This record of early spiral behavior is interpreted as a primitive grazing method formed on muddy laminae above sand layers, and is related to a feeding strategy of an annelid-type of organism. Associated traces are Cruziana cf. semiplicata, Diplocraterion isp., Monocraterion isp., Rusophycus isp., Skolithos linearis Haldeman and Skolithos magnus Howell. The ichnoassemblage is similar to a shallow-water ichnoassociation from the Permian Ecca Group of South Africa (Mason et al., 1983 Mason, T. R., Stanistreet, I. G. and Tavener-Smith, R. 1983. Spiral trace fossils from the Permian Ecca Group of Zululand. Lethaia, 16: 241–247. [Crossref], [Web of Science ®] , [Google Scholar]).     

Sedimentological implications of an unusual form of the trace fossil <Emphasis Type="Italic">Rhizocorallium</Emphasis> from the Lower Muschelkalk (Middle Triassic), S. Poland

Specimens of the trace fossil Rhizocorallium isp. with retrusive limbs occur in 10–20-cm-thick composite beds: the bottom layers contain abundant Planolites and Thalassinoides trace fossils, and the top layers are built of horizontally bedded pelitic limestone and contain less bioturbational structures. The first specimens of Rhizocorallium isp. occur just 2–3 cm above the lower surfaces of the top layers. The horizontal protrusive sections are 1 cm high and 1 cm wide. At some sites, the horizontal tunnels end, and the limbs become first obliquely retrusive (45°) at a distance of 1.5–2.5 cm, and then vertically retrusive at a distance of 4–6 cm, or at once vertically retrusive at a similar distance. They reach to the upper surfaces of the top layers. These composite beds with unusual Rhizocorallium mark successive stages of distal tempestite sedimentation. The erosional events are recorded as uneven bases and a lack of Rhizocorallium, which suggests erosion to approx. 10 cm deep. The lower parts of the top layers record deposition of lime mud from a powerful flow. The protrusive sections of Rhizocorallium record relatively quiet periods resulting from significantly decreased sedimentation. The successive retrusive sections provide evidence that the quiet conditions suddenly came to an end, and that sedimentation was reactivated or accelerated. The last retrusive limbs of Rhizocorallium isp. mark the border of the storm event and the next erosion event. The composition of the fecal pellets shows that these burrowing animals were deposit-feeders, fast enough to find a sufficient amount of food, and also fast enough to escape being buried.     

Lacertoid Footprints of the Upper Permian Arenaria di Val Gardena Formation (Northern Italy)

Lacertoid footprints are the largest component of the Upper Permian Arenaria di Val Gardena Formation ichnofauna that contains hundreds of specimens mostly referred to the ichnogenus Rhynchosauroides Maidwell 1911. In this paper, we analyzed unpublished material and re-examined the Rhynchosauroides footprints of that ichnofauna, in particular the figured specimens. Analysis of Rhynchosauroides and its type ichnospecies R. rectipes Maidwell 1911 was first necessary. This preliminary investigation highlighted several problems, including ichnospecies named on the basis of poorly preserved material and in some cases significantly different from the type ichnospecies.

The study allowed for recognition of three ichnotaxa referred to Rhynchosauroides: Rhynchosauroides pallinii Conti et al., 1977, Rhynchosauroides isp.1 and Rhynchosauroides isp.2 and Ganasauripus ladinus igen. et isp. nov. Other material previously referred to Rhynchosauroides is herein regarded as unclassifiable, in the light of present ichnological knowledge and procedures.     

Cretaceous Nonmarine Trace Fossils from the Hasandong and Jinju Formations of the Namhae Area,Kyongsangnamdo, Southeast Korea

Nonmarine strata of the Cretaceous Hasandong and Jinju formations of the Sindong Group of the Namhae area, Kyongsangnamdo, southeast Korea, contain relatively diverse and abundant ichnofaunas including domichnia, fodinichnia, pascichnia, and repichnia. These ichnofaunas are systematically described herein and three ichnocoenoses, namely the Skolithos, Palaeophycus, and Chondrites ichnocoenoses, are defined based on their paleoenvironmental occurrence. The Skolithos ichnocoenose characterizes tabular laminated sandstone facies deposited on fluvial channel levées. It is dominated by infaunally produced ichnotaxa, particularly Skolithos magnus Howell, Planolites montanus Richter and Chondrites isp. type A, with reduced numbers of Skolithos verticalis (Hall) Taenidium barretti (Bradshaw) and rare examples of Circulichnus montanus (Vyalov), Cochlichnus anguineus Hitchcock, Helminthopsis abeli Ksi??kiewicz, Helimthopsis hieroglyphica Wetzel and Bromley, Laevicyclus isp., Planolites beverleyensis (Billings), Skolithos isp., Spirodesmos isp., Thalassinoides suevicus (Rieth), and rhizoliths. The Palaeophycus ichnocoenose characterizes floodplain environments and occurs in purple mudstones and associated massive sandstone facies of crevasse-splay deposits and dark gray to green shale facies of abandoned channels. Palaeophycus tubularis Hall and Planolites montanus Richter are the most characteristic ichnotaxa, while Beaconites coronus (Frey, Pemberton and Fagerstrom), Planolites annularius Walcott, Skolithos magnus Howell, Thalassinoides paradoxicus (Woodward), and sauropod and ornithopod tracks also occur but in reduced numbers. The Chondrites ichnocoenose occurs in dark gray to black mudstones deposited in marginal lacustrine environments. It is dominated by Chondrites isp. type B. with associated Beaconites antarcticus Vyalov, Helminthopsis hieroglyphica, Palaeophycus tubularis, Skolithos magnus, Taenidium barretti, and Torrowangea rosei Webby. The three ichnocoenoses recognized herein cannot be satisfactorily assigned to currently defined nonmarine ichnofacies, suggesting that existing models of such ichnofacies are in need of additional refinement.