Hierarchical random walks in trace fossils and the origin of optimal search behavior

Efficient searching is crucial for timely location of food and other resources. Recent studies show diverse living animals employ a theoretically optimal scale-free random search for sparse resources known as a Lévy walk, but little is known of the origins and evolution of foraging behaviour and the search strategies of extinct organisms. Here we show using simulations of self-avoiding trace fossil trails that randomly introduced strophotaxis (U-turns) – initiated by obstructions such as ¬¬¬self-trail avoidance or innate cueing – leads to random looping patterns with clustering across increasing scales that is consistent with the presence of Lévy walks. This predicts optimal Lévy searches can emerge from simple behaviours observed in fossil trails. We then analysed fossilized trails of benthic marine organisms using a novel path analysis technique and find the first evidence of Lévy-like search strategies in extinct animals. Our results show that simple search behaviours of extinct animals in heterogeneous environments give rise to hierarchically nested Brownian walk clusters that converge to optimal Lévy patterns. Primary productivity collapse and large-scale food scarcity characterising mass extinctions evident in the fossil record may have triggered adaptation of optimal Lévy-like searches. The findings suggest Lévy-like behaviour has been employed by foragers since at least the Eocene but may have a more ancient origin, which could explain recent widespread observations of such patterns among modern taxa.

Investigating Microbial Trace-fossils and Abiotic Alteration in Hydrovolcanic Tuffs of the Fort Rock Volcanic Field, Oregon

Microbial ichnofossils in volcanic rocks provide a significant record of subsurface microbes and potentially extraterrestrial biosignatures. Here, the textures, mineralogy, and geochemistry of two continental basaltic hydrovolcanic deposits - Reed Rocks and Black Hills - in the Fort Rock Volcanic Field (FRVF) are investigated. Methods include petrographic microscopy, micro and powder X-ray diffraction, SEM/BSE/EDF imaging, energy dispersive spectroscopy, stable isotopes, and X-ray fluorescence. Petrographic analysis revealed granular and tubular textures with biogenic morphologies that include terminal enlargements, septate divisions, branching forms, spiral filaments, and ovoid bodies resembling endolithic microborings described in ocean basalts. They display evidence of behaviour and a geologic context expressing their relative age and syngenicity. Differences in abiotic alteration and the abundance/morphotype assemblage of putative microborings between the sites indicate that water/rock ratio, fluid composition and flux, temperature and secondary phase formation are influences on microboring formation. This study is the first report of reputed endolithic microborings in basalts erupted in a continental lacustrine setting.

Conical sedimentary structures, trace fossils or not? Observations, experiments, and review

Conical sedimentary structures are widespread in the geological column. Those that are mediated by organisms (or organic matter) can be attributed to seven principal processes, which are investigated by experiment and/or field observations: (1) sand collapse into a cavity (decomposed body, open shaft, or gallery), (2) upward (escape) or downward locomotion by an organism through the sediment, (3) upward adjustment (equilibration), (4) casting of coelenterates' excavations, (5) organism-mediated soft-sediment deformation in heterolithic sediment, (6) biodeformational small and large excavations by organisms, and (7) fluid (gas or liquid) escape structures. Footprint loading may also be included. Criteria are given to distinguish between these processes. Dewatering pipes are best recognized by a zone of deformed and fluidized sediment at the base, and association with non-life (lethal) facies. Care must be used in assigning specimens to ichnotaxa, and it is generally necessary to slab, and also to carry out stratinomic investigation in the field.

Spatangoid-produced ichnofabrics (Bateig Limestone, Miocene, Spain) and the preservation of spatangoid trace fossils

A spatangoid-produced ichnofabric is described from the Miocene Bateig Limestone, SE Spain. This ichnofabric is characterized by the dominant presence of large meniscate burrows (Bichordites) produced by irregular echinoids. This constitutes an unusual mode of occurrence for spatangoid bioturbation, as their traces are most typically preserved in bases and tops of sandstone event beds. In fact, despite their important role as burrowers in modern settings (that can be extended back to the Early Cretaceous based on their body fossil record), spatangoid trace fossils (Scolicia and Bichordites) are comparatively rare. Several factors play an important role in their preservation: mechanism of burrowing, sediment characteristics, early diagenesis and presence/absence of deep-tier burrowers. Spatangoid-produced ichnofabrics, such as those from the Bateig Limestone, characterize depositional settings with intermittent deposition of event beds where there is an absence of deeper-tier bioturbation. (C) 2008 Elsevier B.V. All rights reserved

Trace fossils and pseudofossils from the Wealden strata (non-marine Lower Cretaceous) of southern England

The trace fossils of the Wealden (non-marine Lower Cretaceous) of southern England are described. Sixteen invertebrate ichnotaxa include Agrichnium fimbriatus, Beaconites antarcticus, B. barretti, Cochlichnus anguineus, Diplichnites triassicus, Diplocraterion parallelum, Lockeia siliquaria, L. serialis, Monocraterion cf. tentaculum, Palaeophycus striatus, P. tubularis, Planolites montanus, Protovirgularia rugosa, Rhizocorallium isp., Scoyenia cf. gracilis, Unisulcus minutus, insect and root traces. Tetrapod tracks and trackways include tridactyl Iguanodontipus burreyi and other ornithopods, theropod, and tetradactyl sauropod (or possibly ankylosaur), together with extensive dinosaur tramplings. Coprolites are referred to two broad types: spiral, with or without included fish scales (attributable to sharks), and elongate and irregular (possibly produced by reptiles). A skinprint and two types of pseudofossil are also included. Five environmental associations are recognised: (1) lacustrine/lagoonal; (2) brackish incursions (flooding events) into the lacustrine/lagoonal environment; (3) a marginal lacustrine association with fluvial input; (4) a fluvial (lacustrine delta) association; (5) floodplain sediments (seasonal wetlands). These associations are assigned to the fluvial-lacustrine Scoyenia Ichnofacies and the incursions to Glossifungites lchnofacies. (c) 2005 Elsevier Ltd. All rights reserved.

Hiatal surfaces from the Miocene Globigerina Limestone Formation of Malta: Biostratigraphy, sedimentology, trace fossils and early diagenesis

The Miocene Globigerina Limestone of the Maltese islands contains widespread omission surfaces with very different characteristics and origins. The terminal Lower Globigerina Limestone hardground (TLGLHg) formed during a period of falling sea level. Coccolith assemblages suggest shallowness. Sedimentary structures and trace fossil assemblages, indicate increasing frequency of storm events and erosional episodes, towards the surface. Calcite cementation which took place around Thalassinoides burrows and formed irregular nodules was followed by dissolution of aragonite. It is suggested that lithification was linked to microbial reactions involving organic matter. In contrast two later surfaces, the terminal Middle Globigerina Limestone omissionground (TMGLOg), which marks the Lower to Middle Miocene boundary, and the Fomm-ir-Rih local hardground (FiRLHg) both contain early diagenetic dolomite. Lithification took place in two phases. The dolomite is interpreted to have formed beneath the sea floor: it was subsequently exhumed and partially corroded as the precipitation of calcitic and phosphatic cements took place around burrows open to the circulation of sea water. (C) 2008 Elsevier B.V. All rights reserved.

Late Silurian trace fossils from the Melbourne Formation, Studley Park, Victoria, southeastern Australia

An ichnoassemblage of 10 ichnospecies is described for the first time from the Late Silurian Melbourne Formation at Studley Park, Victoria, southeastern Australia. The ichnofauna is preserved in a typical deep-water turbidite succession of alternating thin- to thick-bedded sandstone and thin- to medium-bedded mudrocks. Trace fossils observed within the study site have been assigned to three main ichnofacies. Ichnofacies 1 is best developed on the linguoid-rippled upper surface of thin sandstone beds and includes Laevicyclus, Aulichnites, Nereites, Helminthoidichnites, small Chondrites and possible Zoophycos. Ichnofacies 2 is very similar to Ichnofacies 1 in ichnospecies composition but instead contains large forms of Chondrites together with other thin burrow types usually poorly preserved and in very low abundance compared with Ichnofacies 1. Ichnofacies 3 is preserved mainly as casts on the underside of medium- to thick-bedded turbiditic sandstones, and has a very low diversity, with Planolites being the most common trace. A detailed analysis of the ichnofabrics and tiering structures of these ichnofacies suggest that Ichnofacies 1 and 3 represent "simple tiering’, in contrast to Ichnofacies 2, which is more characteristic of 'complex tiering’. Despite the differences in ichnospecies composition and ichnofabrics between the three recognized ichnofacies, the collective ichnoassemblage from the study site can be assigned confidently to the Nereites ichnofacies and is, therefore, interpreted to have formed in a distal submarine fan environment of lower bathyal to abyssal depth. Further, it is possible to recognize two main subenvironments within this deep-sea setting to account for the differences between the ichnofacies. Ichnofacies 1 and 2 are interpreted to represent a typical Nereites ichnofacies located on a level basin floor subenvironment of relatively low energy conditions at the distal end of a submarine fan deposit. In comparison, Ichnofacies 3 is dominated by Planolites with rare other facies-crossing trace fossil forms, and lacks Nereites. It is, therefore, best interpreted as representing a relatively high-energy environment, possibly a distributary channel near the distal end of the submarine fan system.

Distribution of trace fossils and ichnofabric indices in ODP holes of Leg 119

Trace fossils and ichnofabric were examined from cores of Late Cretaceous to Quaternary age recovered from the Kerguelen Plateau, Indian Ocean. Nearly all of the strata are completely bioturbated, with ichnofabric index 6 most commonly recorded. Preserved discrete trace fossils include Chondrites, Planolites, Zoophycos, and Thalassinoides. A continuous Cretaceous/Tertiary boundary section preserved at ODP Site 738 occurs within a 15-cm-thick interval of laminated sediments. The lack of bioturbation indicates the disappearance of bioturbating organisms from the seafloor, possibly as a result of the same factors that caused the mass extinction or changes in other environmental conditions - most probably, bottom-water oxygen concentrations.

(Table 1) Typical trace-fossil occurrences in DSDP Holes 56-436 and 56-435A

Trace fossils are in places abundant in cores from DSDP Leg 56 sites. They are particularly rich in the pelagic-clay sequence at Site 436. Some significant trace fossils, including Zoophycos, Teichichnus, Chondrites, rind and solid burrows, and pellet-armored rods, are described. The ichnofauna, except for pellet-armored rods in diatomaceous mudstone of the landward trench slope, is characterized by cosmopolitan bathyal to abyssal forms.

Quantitative estimation of bioturbation based on digital image analysis

Quantitative determination of modification of primary sediment features, by the activity of organisms (i.e., bioturbation) is essential in geosciences. Some methods proposed since the 1960s are mainly based on visual or subjective determinations. The first semiquantitative evaluations of the Bioturbation Index, Ichnofabric Index, or the amount of bioturbation were attempted, in the best cases using a series of flashcards designed in different situations. Recently, more effective methods involve the use of analytical and computational methods such as X-rays, magnetic resonance imaging or computed tomography; these methods are complex and often expensive. This paper presents a compilation of different methods, using Adobe® Photoshop® software CS6, for digital estimation that are a part of the IDIAP (Ichnological Digital Analysis Images Package), which is an inexpensive alternative to recently proposed methods, easy to use, and especially recommended for core samples. The different methods — “Similar Pixel Selection Method (SPSM)”, “Magic Wand Method (MWM)” and the “Color Range Selection Method (CRSM)” — entail advantages and disadvantages depending on the sediment (e.g., composition, color, texture, porosity, etc.) and ichnological features (size of traces, infilling material, burrow wall, etc.). The IDIAP provides an estimation of the amount of trace fossils produced by a particular ichnotaxon, by a whole ichnocoenosis or even for a complete ichnofabric. We recommend the application of the complete IDIAP to a given case study, followed by selection of the most appropriate method. The IDIAP was applied to core material recovered from the IODP Expedition 339, enabling us, for the first time, to arrive at a quantitative estimation of the discrete trace fossil assemblage in core samples.

西南极南设得兰群岛中新生代火山岩基底沉积学及其大地构造制约

Livingston Island, the second island of South Shetland Island, constains Mesozoic-Cenozoic basement, Mesozoic-Cenozoic volcanic sequences, plutonic intrusions and post-subduction volcanic rocks, which document the history and evolution of an important part of the South Shetland Islands magmatic arc. The sedimentary sequence is named the Miers Bluff Formation (MBF) and is interpreted as turbidite since the first geological study on South Shetland Islands, and is interpreted as turbidite. It base and top are not exposed, but a thickness of more than 3000m has been suggested and seems plausible. The turbidite is overlain by Mid - Cretaceous volcanic rocks and intruded by Eocene tonalites. The age of the Miers Bluff Formation is poorly constrained Late Carboniferous -Early Triassic. Sedimentary Environment, tectonic setting and forming age of sedimentary rocks of the Miers Bluff Formation were discussed by means of the methods of sedimentology, petrography and geochemistry, combinig with the study of trace fossils and microfossil plants. The following conclusions are obstained. A sedimentary geological section of Johnsons Dock is made by outside measuring and watching, and then according the section, the geological map near the Spanish Antarctic station was mapped. Four pebbly mudstone layers are first distinguished, which thickness is about 10m. The pebbly mudstone is the typical rock of debris flow, and the depostional environment of pebbly mudstone may be the channel of mid fan of submarine fan. The sedimentsry structural characteristics and size analysis of sandstones show the typical sedimentary feature of turbidity flow and the Miers Bluff Formation is a deep-water turbidite (include some gravity-flow sediments). The materials of palaeocurrents suggest the continental slope dip to southeast, and indicate the provenance of turbidity sediment in the northwest area. By facies analysis, six main facies which include seven subfacies were recognized, which are formed in mid-fan and lower-fan of submarine, meanwhile, the sedimentary features of each facies and subfacies are summarized. The study of clastic composition, major elements, trace elements and rare earth elements indicates the forming setting of the Miers Bluff Formaton is active continental margin and continental island arc and the provenance is dissected magmatic arc which main composition is felsic gneiss. Many trace fossils of the whole succession were found in the turbidites of the Miers Bluff Formation. All these trace fossils are deep sea ichnofossils. There are fifteen ichnogenus, sixteen ichnospecies. Moreover, a new trace fossil was found and a new ichnogenus and new ichnospecies was proposed - Paleaichnus antarctics ichnogen, et ichnosp, nov.. Except the new ichnogenus and ichnospecies, others had been found in deep-sea flysch turbidites. Some are in mudstone and are preserved in the cast convex of overlying sandstone sole, they formed before turbidity flows occurred and belong to the high-different Graphoglyptida of fiysch mudstone. Others as Fucusopsis and Neonereites are preserved in sandstones and stand for trace assemblages after turbidity sedimentation. These trace fossils are typical members of abyssal "Nereites" ichnofacies, and provide for the depositional environment of the Miers Bluff Formation. Fairly diverse microfossil plants have been recovered from the Miers Bluff Formation, Livingston Island, including spores, pollen, acritarchs, wood fragments and cuticles. Containing a total of about 45 species (forms) of miospores, the palynofiora is quantitatively characterized by the dominance of non-striate bisaccate pollen, but spores of pteridophytes and pollen of gymnosperms are proportionate in diversity. It is somewhat comparable to the subzone C+D of the Alisporites zone of Antarctica, and the upper Craterisporites rotundus zone and the lower Polycingulatisporites crenulatus zone of Australia, suggesting a Late Triassic (possibly Norian-Rhaetian) age, as also evidenced by the sporadic occurrence of Aratrisporites and probable Classopollis as well as the complete absence of bisaccate Striatiti. The parent vegetation and paleoclimate are preliminarily deduced. At last, the paper prooses the provenance of sedimentary rocks of the Miers Bluff Formation locates in the east part to the southern Chile(or Southern South American). In the Triassic period, contrasting with New Zealand, Australia and South American of the Pacific margin of Gondwanaland, the Miers Bluff Formation is deposited in the fore-arc basin or back-arc basin of magmatic arc.