Short-term colonization trace-fossil assemblages in a carbonate lacustrineKonservat-Lagerstätte (Las Hoyas fossil site, Lower Cretaceous, Cuenca, Central Spain)

Summary Rhythmically laminated fine-grained lacustrine carbonatcs of the Cretaceous La Huérguina Limestone Formation at Las Hoyas fossil site, central Spain, contain not only a spectacular body fossil fauna, but also vertebrate and invertebrate trace fossils. The study of the Las Hoyas invertebrate ichnofauna provides valuable taphonomic, paleoecological and paleoenvironmental information and represents one of the first attempts to study in detail a lacustrine trace-fossil assemblage in carbonate rocks. The Las Hoyas ichnofauna includesCruziana problematica, Helminthoidichnites tenuis, Lockeia isp.,Palaeophycus tubularis, andTreptichnus pollardi. This assemblage is characterized by low ichnodiversity and small burrow size, and it is dominated by surface trails and extremely shallow burrows produced mostly by detritus feeders. The Las Hoyas ichnofauna indicates low to moderately low energy, permanently submerged, shallow-lacustrine areas. The ichnofauna is suggestive of environmental stress, most likely oxygen-depleted conditions in interstitial waters. Occurrence of traces in event beds indicates opportunistic colonization by epifauna and very shallow infauna during brief periods of improved oxygenation related to the passage of density underflows and dilute turbidity currents. The Las Hoyas ichnofauna resembles other shallow lacustrine trace-fossil assemblages of the archetypalMermia ichnofacies and differs significantly from the archetypalScoyenia ichnofacies. It is therefore best regarded as a proximal, depauperate example of theMermia ichnofacies. The Las Hoyas assemblage is similar to the slightly older lacustrine Montsec ichnofauna, but it is remarkably different from marine ichnocoenoses of other Mesozoickonservat-lagerst?tten in lithographic limestones such as those of Cerin, Solnhofen and Nusplingen.     

Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove,Victoria

Two thin‐toed tridactyl tracks in a fluvial sandstone bed of the Eumeralla Formation (Albian) at Dinosaur Cove (Victoria, Australia) were likely made by avian trackmakers, making these the oldest known fossil bird tracks in Australia and the only Early Cretaceous ones from Gondwana. These tracks, which co‐occur on the same surface with a slightly larger nonavian theropod track, are distinguishable by their anisodactyl form, hallux impressions and wide digit II–IV divarication angles. A lengthy hallux impression and other deformational structures associated with one track indicate foot movement consistent with an abrupt stop, suggesting its tracemaker landed after either flight or a hop. The single nonavian theropod track is similar to other tracks described from the Eumeralla Formation at another locality. The avian footprints are larger than most Early Cretaceous avian tracks recorded worldwide, indicating sizeable enantiornithine or ornithurine species in formerly polar environments of Australia. The avian tracks further supplement scant body fossil evidence of Early Cretaceous birds in southern Australia, which includes a furcula from the Wonthaggi Formation. Because of this discovery, Dinosaur Cove, previously known for its vertebrate body fossils, is added to a growing list of Early Cretaceous vertebrate tracksites in southern Australia.     

Enigmatic ichnites in the middle triassic of Southern Spain

In southern Spain, within the Chiclana de Segura Formation of red detrital fades, enigmatic structures were found whose origins are discussed here. Each structure is composed of the union of elongated traces, two laterals and a shorter central one. All are concave epireliefs at the top of a sandstone bed whose age is assumed to be Ladinian (Triassic). The sediments formed within a fluvial environment. We describe these imprints, their structures and measurements, and discuss their orientations and associations. The traces probably have an organic origin, and are most likely tracks made by a vertebrate, possibly an archosaur, although no osseous remains have been found to confirm this interpretation.     

The Lower Cretaceous Patuxent Formation Ichnofauna of Virginia

The vertebrate fauna from the Lower Cretaceous Patuxent Formation of Virginia is composed of a single partial fish impression from the James River at Dutch Gap and a diverse tetrapod ichnofauna from near Fredericksburg that includes trace fossils made by frogs, turtles, theropods, sauropods, ankylosaurs, and ornithopods. The footprints occur on overbank deposits preserved locally within a fluvial braided-stream sequence that formed near the western border of the Early Cretaceous Atlantic Coastal Plain.     

Modern and fossil traces in terrestrial lithic substrates

A number of biogenic processes leads to the formation of distinctive traces in terrestrial lithic substrates. These include: burrowing by vertebrates in moderately lithified rocks; scraping by mammals; smoothing and polishing of limestone surfaces by the locomotion of mammals; excavation by bees, wasps, and ants producing nesting and dwelling tunnels; dissolution of limestone surfaces by terrestrial snails; endolithic activity of fungi, algae, and lichens on subaerial rock surfaces; root corrosion; etc. Processes of biochemical weathering, biophysical erosion, and enlargement of cracks and fissures by the pressure of plant roots do not leave distinctive traces and therefore lie outside the ichnological realm. The fossil preservation of terrestrial bioerosional traces is expected to be uncommon. Nevertheless, various possible means of preservation must be considered, such as by rapid burial by volcanic material, by fluvial sediments, by travertine or tufa, by loess, “conservation”; in caves, case hardening of surfaces of porous rocks, and preservation of subsoil traces below fossil soils.     

Trace fossil evidence of predation upon bone‐eating worms on a baleen whale skeleton from the Oligocene of New Zealand

The osteophagous worm Osedax (Annelida: Siboglinidae) colonizes vertebrate bones in deep‐sea environments globally. Osedax bioerosion of modern bones suggests a potentially destructive agent in the marine vertebrate fossil record, but the dearth of published reports of abundant Osedax traces suggests an uncertain taphonomic influence of this organism. This study reports Osedax traces (Osspecus boreholes, pockmarks and collapsed galleries) in an Oligocene baleen whale (Cetacea: Eomysticetidae) from New Zealand, which constitute the first record of fossil Osedax traces from the southern hemisphere. Some Osedax traces are cross‐cut by linear biogenic scrape marks, implying that sharks or bony fish fed upon Osedax worms, a process which compounds or potentially accelerates worm‐inflicted damage to vertebrate bones in marine environments.     

A Valanginian crustacean microcoprolite ichnofauna from the shallow-marine hydrothermal vent site of Zengővárkony (Mecsek Mts., Hungary)

A rich and diverse crustacean microcoprolite ichnofauna is reported from the Valanginian of the Mecsek Mts., Hungary. Localities from the Bolondút and Dezső Rezső valleys near Zengővárkony (Eastern Mecsek Mts.) provided Favreina aff. dispentochetarius, Palaxius darjaensis, P. triochetarius, P. salataensis, P. tetraochetarius, and P. decaochetarius. They occur in limonitic and partly metasomatized limestones, limestone olistoliths, and ammonite body-chambers. The remarkable richness of this ichnofauna plus the previous records from the same environment (altogether 11 ichnospecies of four ichnogenera from a single stratigraphic level) make it the most diverse crustacean ichnofauna of the Mesozoic. The former ecosystem of the ichnofauna is considered a deeper shallow-marine (water depth <300 m) hydrothermal vent on which the producers of the microcoprolites lived. The related brachiopod fauna shows a remarkable size and is dominated by Lacunosella. This fauna is not similar to typical fossil deep-sea vent faunas; however, it offers a thorough understanding of other fossil shallow-marine hydrothermal vent faunas and the role of crustaceans in these ecosystems. This is the first record of crustacean coprolites from such a fossil site and documents that crustaceans were diverse and played an important role in shallow-marine, non photosynthesis-based ecosystems in the geological past.     

A Tetrapod Ichnoassociation from the Middle Triassic (Anisian,Pelsonian) of Northern Italy

A new Middle Triassic ichnofauna recovered nearby the town of Tisens (Bozen) in the Adige Valley (Trentino, Northern Italy) is described. A relatively large ichnoassociation, in which various ichnogenera can be recognized, comes from the basal inter-supratidal layers of a mixed carbonate siliciclastic unit of Pelsonian (Middle Anisian) age. Most of the isolated footprints and trackways pertain to lizard-like reptiles referable to Rhyncosauroides and, subordinately, to archosaurian reptiles. In particular, the ichnogenera Rotodactylus, Synaptichnium and Chirotherium have been recognized. Many tracks are at present unidentified; among them a lizard-like morphotype (morphotype A). The site also yelded numerous fossil plant horizons and some scattered vertebrate remains referable to Placodontidae (cf. Placodus gigas). The site shows the continuity in the Pelsonian of the “chirothere ichnofauna” and the dominance of Rhynchosauroides in carbonate, tidal flat environments.     

Origin of trace fossils in Polish Carpathian flysch

Fifteen features of trace fossils on flysch sandstone soles have been found useful in distinguishing between pre-depositional traces of mud-inhabiting animals and post-depositional traces of sand-dwellers. The Polish Carpathian flysch ichnofauna of 55 genera and 145 species, including 41 ichno-genera that occur at least in part on sandstone soles, is dominated both in number of taxa and number of specimens by traces of mud inhabitants, and most traces, both of mud- and sand-dwellers, were produced by infaunal deposit-feeders. Because mud was the temporally dominant substrate, the mud-dwellers were much more abundant over long periods of time, but the fauna of turbidites and similar sands may have been nearly as diverse and abundant at any given moment as the mud fauna at another. The currents depositing these sands eroded surficial layers of mud and excavated any burrow fillings in almost every case. The flysch ichnofauna differs from observed trace assemblages of modern deep-sea substrates in that the former consists almost exclusively of burrows, while surface traces dominate the latter.     

Pliocene facies and fossil contents of Qaret El-Muluk formation at Wadi El-Natrun depression, Western Desert, Egypt

Summary A sequence of late Pliocene sediments (Qaret El-Muluk Formation) covers large areas within Wadi El-Natrun depression; at the western periphery of the Nile Delta. The exposed sequence is composed of friable sands. mudstones, shales and minor limestones with combined total thickness of approximately 50 m. The fossil content consists of fresh and brackish water elements; charophytes, ostracods, gastropods, oysters and benthonic foraminifera. Planktonic forams which suggest marine influence have been also recorded. Terrestrial and aquatic continental vertebrate fauna includes mammals, reptiles, fish and aves. Several subtypes of ichthyoliths have been discovered in this study. Both lithological and paleontological data testify to a depositional environment which belongs to a marginal region where fluvial and marine deposition meet in shallow brackish lagoons. The sediments display changes from channelled fluvial sands to deltaic interbedded sands and mudstones to lagoonal shales and limestones to lacustrine chara limestones. Vertical stacking of facies is indicative of a series of transgressive-regressive events. The transgression brought shallow lagoonal conditions to the area. The transgressions were incomplete due to high rate of fluvial input. The lagoon dried up intermittently and calcretes and red brown soils were formed. Each regression is indicated by river generated clastic influx. During periods of active fluvial input, fluvial sediments prograded lagoonward. The clastic material was probably a mixture of reworked detritus from older sediments. An admixture of first cycle material from the basement rocks of the Red Sea hills cannot be ruled out. The diversity of provenance argues for several channels of a large river which existed during the late Pliocene.     

Cretaceous small scavengers: feeding traces in tetrapod bones from Patagonia, Argentina

Ecological relationships among fossil vertebrate groups are interpreted based on evidence of modification features and paleopathologies on fossil bones. Here we describe an ichnological assemblage composed of trace fossils on reptile bones, mainly sphenodontids, crocodyliforms and maniraptoran theropods. They all come from La Buitrera, an early Late Cretaceous locality in the Candeleros Formation of northwestern Patagonia, Argentina. This locality is significant because of the abundance of small to medium-sized vertebrates. The abundant ichnological record includes traces on bones, most of them attributable to tetrapods. These latter traces include tooth marks that provde evidence of feeding activities made during the sub-aerial exposure of tetrapod carcasses. Other traces are attributable to arthropods or roots. The totality of evidence provides an uncommon insight into paleoecological aspects of a Late Cretaceous southern ecosystem.     

Microbially induced sedimentary structures indicating climatological, Hydrological and depositional conditions within recent and pleistocene coastal facies zones (Southern Tunisia)

Summary Extensive tidal areas of the Recent coast of southern Tunisia are overgrown by microbial mats. Different mat types of which each are dominated by distinct and well adapted cyanobacterial species develop. Ecological response of the mat-forming microorganisms to climatological hydrological and sedimentological factors produce characteristic sedimentary structures (=microbially induced sedimentary structures). A suecession of Pleistocene rocks crops out near the lagoon El Bibane, southern Tunisia. The stratigraphic section comprises structures that we regard as fossil equivalents to those microbially induced structures we observe in the Recent coastal area. Preservation of the structures is result of lithification of the microbial mats. This we conclude from fossil filaments of cyanobacteria visible within the rock matrix. The Recent microbially induced sedimentary structures indicate facies zones within the modern tidal environment. Comparison of the Recent structures with the fossil analogues recorded in the stratigraphic section aids to identify the same distinct facies zones within the Pleistocene coastal environment also. Erosion by water currents forms step-like cliffs, and the microbial mat is undermined and ripped off piece by piece. shallows within the supratidal area are overgrown by copious microbial mats comprising structures like biolaminites and—varvites, as well as polygons of cracks. The features originate from effects triggered by seasonal variations of climate. Tufts and reticulate pattern of bulges indicate supernatant water films covering the mat surfaces. Morphologically higher parts of the Recent tidal area are overgrown by single-layered mats forming petees, induced by microbial mat growth and evaporitive pumping. The study demonstrates that microbially induced sedimentary structures can be used to reconstruct small-scaled facies zones within coastal environments. The also include hints on paleoclimatological, hydrological and sedimentological conditions.     

Evidence of Osedax worm borings in Pliocene (∼3 Ma) whale bone from the Mediterranean

Osedax worms subsist entirely on vertebrate skeletons on the seafloor, using root-like tissues to bore into and degrade the bones. Paleontologists have only recently begun to appreciate the possible destructive effect that these worms may have had on the marine vertebrate fossil record and little is known of their evolutionary history. Using microcomputed tomography, we document Osedax-like borings in a fossil whale bone from the Pliocene of Italy and present new data on the borings of extant Osedax worms. The fossil borings are distinguished from those of other known borers and identified as traces of Osedax activity based on diagnostic features. Our results suggest that it is necessary to isolate individual borings for the confident identification of Osedax traces. This is only the second paleogeographic occurrence of Osedax in the fossil record and indicates that by the Pliocene these worms had colonised a large portion of the world's oceans. This is the first evidence for Osedax in the Mediterranean, past or present, and suggests that more species await discovery in this region.     

Trace Fossils from the Brioverian (Ediacaran–Fortunian) in Brittany (NW France)

The Ediacaran–Fortunian ichnofauna from Central Brittany (NW France) is revised for the first time since the pioneering work by Lebesconte at the end of the 19th century. The study is based on fossils from the type-localities of the historical Brioverian taxa Montfortia (traces from Montfort-sur-Meu) and Neantia (wrinkle surfaces from Néant-sur-Yvel), and on two new outcrops from Saint-Gonlay. The ichnofossil assemblage includes Helminthoidichnites tenuis, Helminthopsis tenuis, Gordia marina, Palaeophycus tubularis, ?Neonereites uniserialis, and ?Spirodesmos archimedeus. Locally, the grazing traces are associated with wrinkle surfaces considered as microbially induced sedimentary structures (MISS). The sedimentological characteristics of these deposits correspond to a shelf marine environment. Both U-Pb datings on detrital zircon and ichnostratigraphic criteria tend toward an Ediacaran age of the fossiliferous deposits, but we cannot exclude the possibility of a Fortunian age at the present state of knowledge.     

Soils,time, and primate paleoenvironments

Soils are the skin of the earth. From both poles to the equator, wherever rocks or sediment are exposed at the surface, soils are forming through the physical and chemical action of climate and living organisms. The physical attributes (color, texture, thickness) and chemical makeup of soils vary considerably, depending on the composition of the parent material and other variables: temperature, rainfall and soil moisture, vegetation, soil fauna, and the length of time that soil–forming processes have been at work. United States soil scientists¹ have classified modern soils into ten major groups and numerous subgroups, each reflecting the composition and architecture of the soils and, to some extent, the processes that led to their formation. The physical and chemical processes of soil formation have been active throughout geologic time; the organic processes have been active at least since the Ordovician.² Consequently, nearly all sedimentary rocks that were deposited in nonmarine settings and exposed to the elements contain a record of ancient, buried soils or paleosols. A sequence of these rocks, such as most ancient fluvial (stream) deposits, provides a record of soil paleoenvironments through time. Paleosols are also repositories of the fossils of organisms (body fossils) and the traces of those organisms burrowing, food–seeking, and dwelling activities (ichnofossils). Indeed, most fossil primates are found in paleosols. Careful study of ancient soils gives new, valuable insights into the correct temporal reconstruction of the primate fossil record and the nature of primate paleoenvironments.     

A NEW GENUS OF HIODONTIDAE FROM LUOZIGOU BASIN, EAST JILIN

<正> Manchurichthys (Saito, 1936) has been, for a long time, the only representative of fossil vertebrate known from the Oil-bearing member of Dalazi Formation, Luozigou Basin, East Jilin. Since 1978, some new materials have been collected there by the Regional Geological Brigade of Jilin Province and the present author. Two more forms of fossil fishes are not known to occur in the rocks of this section. One is Sinarnia luozigouensis (Li, 1984) which is referred to Amiiformes; the other, Yanbiania (gen. nov.), is described in this paper and included in the osteoglossomorphan family Hiodontidae.     

Names for trace fossils: a uniform approach

The taxonomic treatment of trace fossils needs a uniform approach, independent of the ethologic groups concerned. To this aim, trace fossils are rigorously defined with regard to biological taxa and physical sedimentary structures. Potential ichnotaxobases are evaluated, with morphology resulting as the most important criterion. For trace fossils related to bioerosion and herbivory, substrate plays a key role, as well as composition for coprolites. Size, producer, age, facies and preservation are rejected as ichnotaxobases. Separate names for undertracks and other poorly preserved material should gradually be replaced by ichnotaxa based on well-preserved specimens. Recent traces may be identified using established trace fossil taxa but new names can only be based on fossil material, even if the distinction between recent and fossil may frequently remain arbitrary. It is stressed that ichnotaxa must not be incorporated into biological taxa in systematics. Composite trace fossil structures (complex structures made by the combined activity of two or more species) have no ichnotaxonomic standing but compound traces (complex structures made by one individual tracemaker) may be named separately under certain provisions. The following emendations are proposed to the International Code of Zoological Nomenclature: The term 'work of an animal' should be deleted from the code, and ichnotaxa should be based solely on trace fossils as defined herein.     

Paleobiology of a Neoproterozoic tidal flat/lagoonal complex: the Draken Conglomerate Formation, Spitsbergen

Carbonates and rare shales of the ca 700-800 Ma old Draken Conglomerate Formation, northeastern Spitsbergen, preserve a record of environmental variation within a Neoproterozoic tidal flat/lagoon complex. Forty-two microfossil taxa have been recognized in Draken rocks, and of these, 39 can be characterized in terms of their paleoenvironmental distributions along a gradient from the supratidal zone to permanently submerged lagoons. Supratidal to subtidal trends include: increasing microbenthic diversity, increasing abundance and diversity of included allochthonous (presumably planktonic) elements, decreasing sheath thickness of mat-building organisms (with significant taphonomic consequences), and an increasing sediment/fossil ratio in fossiliferous rocks. Five principal and several minor biofacies can be distinguished. The paleoecological resolution obtainable in the Draken Conglomerate Formation rivals that achieved for most Phanerozoic fossil deposits. It documents the complexity and diversity of Proterozoic coastal ecosystems and indicates that both environment and taphonomy need to be taken into explicit consideration in attempts to understand evolutionary trends in early fossil record. Three species, Coniunctiophycus majorinum, Myxococcoides distola, and M. chlorelloidea, are described as new; Siphonophycus robustum, Siphonophycus septatum, and Gorgonisphaeridium maximum are proposed as new combinations.     

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

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

Humans Running at Stadiums and Beaches and the Accuracy of Speed Estimations from Fossil Trackways

The concept of dynamic similarity between mechanical properties of vertebrates and engineered structures has served in previous work to suggest that there is a power law relationship between vertebrate speeds and stride length. This relationship, with some additional assumptions about hind limb height, has permitted the calculation of speeds from fossil trackways of dinosaurs. However, there are claims that uncertainties are large. In this work we analyze the accuracy of speed calculations for fossil vertebrates based on fossil trackways by using data derived from both athletic competitions and an experiment with humans walking and running on a beach. Our results show that although there are somewhat different running regimes, in general terms human speed can be described in a simple way, and differences between observed and predicted speeds usually are no more than 10–15%. Thus, while recognizing that some uncertainty remains in the estimation of hind limb height, we conclude that reliable speed calculations can be obtained from vertebrate fossil trackways. Our results also show that very reliable speed estimates can be obtained from human fossil trackways directly from stride length measurements.