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

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

Oceanic Anoxic Event at the Cenomanian–Turonian boundary interval (OAE-2): ichnological approach from the Betic Cordillera, southern Spain

Ichnological analysis of dark sediments of the Oceanic Anoxic Event at the Cenomanian–Turonian boundary interval (OAE-2) from the Betic Cordillera, southern Spain, reveals the presence of ichnofabrics containing trace fossils Chondrites isp., Palaeophycus heberti , Planolites isp., Thalassinoides isp., Trichichnus linearis , Zoophycos isp., and other bioturbational structures. Analysis of the ichnofabrics and trace fossil diversity allowed the reconstruction of relative oxygenation. Before the OAE-2, oxygenation of sediments was generally good, but it was punctuated by short anoxic events. During the OAE-2i, several longer anoxic intervals were interrupted by shorter dysaerobic and oxic periods. After the OAE-2, oxygenation improved and almost all trace fossils known before OAE-2 reappeared, although oxygenation dropped a few times to anoxia. Generally, the short oxygenation changes reveal a periodicity, which may suggest a Milankovitch control.     

Oceanic redox evolution across the end-Permian mass extinction at Penglaitan section,South China

The end-Permian mass extinction (EPME) was known as the most severe biocrisis of the past 600 Ma. In order to explore the redox state of deep water environments, and the causal relationship between anoxia/euxinia and the EPME, this study selected the Penglaitan section in Guangxi, China, and measured the iron speciation and concentrations of trace elements and major elements. Several redox proxy approaches, including both iron speciation proxies and the excess fractions and enrichment factors of redox sensitive elements (RSEs), indicate that anoxia in the Penglaitan depositional environment was sporadic, with the majority of the section deposited under oxic conditions. This includes the interval immediately prior to and during the EPME; euxinia did not occur at the Penglaitan section around the EPME. Thus, anoxia was not the primary driver for the EPME at the Penglaitan section, and the accumulation of free H₂S in the oceanic water-column could not have been the only proximal kill mechanism for the EPME at the Penglaitan section.     

Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis

A brachiopod fauna comprising nine species in eight genera from three closely spaced stratigraphic horizons of the same stratigraphic section is described for the first time from the Laibin Limestone in the uppermost part of the Maokou Formation in the Guadalupian/Lopingian (G/L) GSSP section at Penglaitan, Guangxi Autonomous Region, South China. The brachiopod assemblages are bracketed between two conodont zones: Jinogondolella xuanhanensis Zone below and Jinogondolella granti Zone above and, therefore, they can be safely assigned to the latest Capitanian in age. However, all but one of the nine brachiopod species from the Laibin Limestone carry strong early Lopingian (Wuchiapingian) aspect. Thus, the discovery of this brachiopod fauna not only suggests that some Lopingian brachiopod species had already appeared in the late Guadalupian (Capitanian); more importantly, it has also highlighted the fact that both the previously noted pre-Lopingian life crisis (or end-Guadalupian or Middle Permian mass extinction) and Lopingian recovery/radiation actually occurred in late Capitanian times, sometime before the G/L chronostratigraphic boundary. So far, the Penglaitan GSSP section provides the highest-resolution disappearance patterns of different fossil groups around the G/L boundary.     

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.     

山西兴县中奥陶统马家沟组五5亚段遗迹化石组合及其古环境意义

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

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

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

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.     

Superoxide Dismutase as an Anaerobic Polypeptide : A Key Factor in Recovery from Oxygen Deprivation in Iris pseudacorus?

The perennating organ, the rhizome, was chosen for examination of response to anoxia in the species Iris pseudacorus L., Iris germanica L. var Quechei, and Glyceria maxima (Hartm.) Holmberg. These monocots are known to differ in their tolerance of anoxia. Intact rhizomes were subjected to periods of prolonged anoxia of up to 28 days and superoxide dismutase (SOD) activity was determined in a 48 hour postanoxic recovery phase. Tests were performed to ensure the accuracy of the measured enzyme activities. In the most anoxia tolerant species, I. pseudacorus, SOD activity rose continuously during the period of imposed anoxia, and levels were maintained in the postanoxic recovery phases: 28 days brought about a 13-fold increase to 1576 U SOD per milligram protein. Small increases were found in the less anoxia tolerant I. germanica during anoxic/postanoxic phases, while a drop in activity was recorded in the least anoxia tolerant G. maxima. However, initial levels in G. maxima were more than twice as high as in the other two species. Experiments applying cycloheximide to anoxic rhizome slices of I. pseudacorus inhibited the increase in SOD activity. This indicates that SOD is, paradoxically, induced under anoxia and we suggest that in this species SOD is one of the enzymes identified as anaerobic polypeptides. The significance of the induction of an `oxygen-protecting' enzyme during complete oxygen deprivation is discussed with regard to a possible critical role during recovery from anoxic stress.     

Palaeocupressinoxylon uniseriale n. gen. n. sp., a gymnospermous wood from the upper Permian of Central Taodonggou,southern Bogda Mountains,northwestern China

A silicified wood, Palaeocupressinoxylon uniseriale n. gen. n. sp., is described from the upper Permian of the Central Taodonggou section, Turpan–Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. Multidisciplinary data including U–Pb ID–TIMS zircon dating, vertebrate and invertebrate biostratigraphic, and cyclostratigraphic correlation from current and previous studies indicate that the fossil bearing interval is Wuchiapingian (late Permian) in age. The pycnoxylic wood consists of thick-walled tracheids and parenchymatous rays. It is characterized by separated uniseriate radial tracheidal pits, uniseriate ray cells, and cupressoid cross-field pitting. The absence of growth rings in the wood, together with the occurrence of Argillisols, Gleysols, and Histosols above and below the fossil interval, suggests that a stable landscape and a perennially humid climate prevailed in the Taodonggou area during the Wuchiapingian.     

Potassium retention in crab nerve

Spectrophotometric determinations of the potassium content of small volumes of sea water brought into contact with crab nerves over half-hour intervals demonstrate (a) a continuous loss of potassium from the fibers while at rest in oxygen; (b) an appreciable increase in the leakage during anoxia; (c) a reabsorption of potassium during postanoxic recovery; and (d) a reduction in the amplitude of the anoxic and postanoxic changes in the presence of glucose.     

Upper Triassic (Keuper) non-marine trace fossils from the Haßberge area (Franconia, south-eastern Germany)

Various trace fossils from the Hassberge Formation and the Löwenstein Formation (Middle Keuper, Upper Triassic) of the Haßberge region are described. Twenty-three different forms have been identified, 17 of which are named, includingCruziana pascens n. isp.,Lockeia cunctator n. isp., andRusophycus versans n. isp.Lockeia siliquaria James, 1879,L. amygdaloides (Seilacher, 1953),L. triangulichnus Kim, 1994, andL. elongata (Yang, 1984) are revised and synonymized under the oldest available name,L. siliquaria James, 1879.Rusophycus eutendorfensis (Linck, 1942) andR. carbonarius Dawson, 1864 are revised. The diagnosis ofPolykladichnus Fürsich, 1981 is emended, and a diagnosis forHelminthoidichnites Fitch, 1850 is given for the first time. Among the described ichnotaxa,Skolithos ispp.,Rusophycus carbonarius, andTaenidium barretti are the most common forms. The trace fossil association is typical of theScoyenia ichnofacies, which indicates non-marine, periodically or completely inundated environments, such as floodplains and lake margins. Two palaeoichnocoenoses are identified. One ichnocoenosis, dominated byCruziana problematica, cf.Polykladichnus isp., andSkolithos isp. B characterizes margins of trough cross-bedded sandstones. Another ichnocoenosis, dominated byRusophycus versans n. isp.,Taenidium barretti,Scoyenia gracilis andSkolithos isp. A is related to ephemeral lake deposits. Taxonomic recommendations for the use of hitherto described and figured invertebrate Keuper trace fossils from Germany are given.     

Stratigraphic application of Thalassinoides ichnofabric in delineating sequence stratigraphic surfaces (Mid-Cretaceous), Kopet-Dagh Basin,northeastern Iran

This study integrates ichnological and sedimentological data to refine depositional sequences and interpretations of sea-level dynamics for the shallow marine, Albian–Cenomanian Aitamir Formation in northeastern Iran. Three ichnofabrics are present in a succession of glauconitic mudstone and sandstone. This is a sequence that grades upward from a lower glauconitic sandstone unit with trough cross-stratification, hummocky and ripple cross-lamination into a fining-up unit of mudstone with intercalated sandstone beds. The lower unit contains an ichoassemblage of the Ophiomorpha–Palaeophycus ichnofabric (upper shoreface), whereas the upper unit bears ichnoassemblages of the Thalassinoides ichnofabric (in a distinctive level at the top cycle which demarcates the base of the next cycle) (middle shoreface) and the Chondrites–Planolites ichnofabric (lower shoreface). An upper shoreface–lower shoreface trend from the Ophiomorpha–Palaeophycus ichnofabric to the Chondrites–Planolites ichnofabric represents a deepening-upward sequence. An integrated sedimentological and ichnological approach has allowed the recognition of the internal organization of the sequence and the characterization of significant discontinuity surfaces at sequence scales. Thalassinoides ichnofabric reveals colonization of firmgrounds during prolonged times between erosion and deposition related to transgressive surfaces. Transgressive surfaces (sequence boundaries) are generally well-cemented and marked by increased glauconite content, and densely crowded, predominantly vertical or oblique, relatively large, very distinct, unlined, and uncompacted burrows (omission suite) and are associated with rare highly abraded and fragmented shell remains.     

Anaerobiosis-induced differentiation of Acanthamoeba castellanii

Acanthamoeba castellanii is a free living amoeba ubiquitous in soil and also commonly found in aquatic environments. In waterlogged soils, anoxia is quickly established as the dissolved oxygen is consumed by the organisms present. We were interested in the effects of anoxic conditions upon this organism. Batch cultures degassed with N₂ during mid-exponential growth, induced encystation within 12 h of anoxia, and mature cysts were formed within 2–3 days. Excystation (99%) was achieved by subsequent aeration of these cultures after 3–6 days. Anoxia-induced cysts, maintained in anoxic conditions for up to four months, remained viable. Difference spectra, during anaerobiosis, revealed that cytochromes were not lost, suggesting that the organism retains its respiratory components. The growth rate of trophozoites, grown in a chemostat, was dependent on the concentration of O₂ in the head space and glucose uptake increased at lower dissolved O₂ tensions. The results obtained suggest that A. castellanii has a complex adaptive strategy enabling it to cope with microaerobic and anoxic conditions which may be experienced in the environment.     

Anoxia-Reoxygenation Regulates Mitochondrial Dynamics through the Hypoxia Response Pathway,SKN-1/Nrf,and Stomatin-Like Protein STL-1/SLP-2

Many aerobic organisms encounter oxygen-deprived environments and thus must have adaptive mechanisms to survive such stress. It is important to understand how mitochondria respond to oxygen deprivation given the critical role they play in using oxygen to generate cellular energy. Here we examine mitochondrial stress response in C. elegans, which adapt to extreme oxygen deprivation (anoxia, less than 0.1% oxygen) by entering into a reversible suspended animation state of locomotory arrest. We show that neuronal mitochondria undergo DRP-1-dependent fission in response to anoxia and undergo refusion upon reoxygenation. The hypoxia response pathway, including EGL-9 and HIF-1, is not required for anoxia-induced fission, but does regulate mitochondrial reconstitution during reoxygenation. Mutants for egl-9 exhibit a rapid refusion of mitochondria and a rapid behavioral recovery from suspended animation during reoxygenation; both phenotypes require HIF-1. Mitochondria are significantly larger in egl-9 mutants after reoxygenation, a phenotype similar to stress-induced mitochondria hyperfusion (SIMH). Anoxia results in mitochondrial oxidative stress, and the oxidative response factor SKN-1/Nrf is required for both rapid mitochondrial refusion and rapid behavioral recovery during reoxygenation. In response to anoxia, SKN-1 promotes the expression of the mitochondrial resident protein Stomatin-like 1 (STL-1), which helps facilitate mitochondrial dynamics following anoxia. Our results suggest the existence of a conserved anoxic stress response involving changes in mitochondrial fission and fusion.     

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.     

Palaeoredox geochemistry and bioturbation levels of the exceptionally preserved early Cambrian Indian Springs biota,Nevada, USA

The early Cambrian Indian Springs biota of western Nevada, USA, exhibits Burgess Shale‐type (BST) preservation of a diverse array of animal phyla, including the earliest definitive echinoderms. It therefore provides an important window on animal life during the Cambrian radiation. The objective of this study was to analyse the trace metal palaeoredox geochemistry and bioturbation levels of this BST deposit in order to characterize the palaeoenvironmental conditions in which these animals lived and their remains preserved. A total of 28 rock samples were collected from outcrops at three previously reported intervals of exceptional preservation at the Indian Springs locality, as well as from one interval not exhibiting such preservation. An additional 20 random samples were collected from talus for comparison. In the laboratory, the samples were analysed for trace metal palaeoredox indices (V/Cr and V/(V + Ni) ratios). Bioturbation levels were assessed through X‐radiography and petrographic thin sections using the ichnofabric index (ii) method. Additional samples from coeval strata of the Poleta Formation in the White‐Inyo Mountains, CA, that lack BST preservation were also analysed with the same methodology. Results indicate that oxic bottom water conditions dominated during deposition of these strata, despite consistently low bioturbation levels. This pattern holds for intervals with BST preservation and those without. Although ephemeral incursions of low‐oxygen waters may have taken place, there is no evidence for persistent oxygen restriction in these palaeoenvironments. The low levels of bioturbation indicate limited mixed layer development and a redox boundary near the sediment–water interface, likely allowing post‐burial BST preservation to occur even in this setting dominated by oxic bottom waters. Palaeoecological reconstructions and taphonomic hypotheses relating to the Indian Springs Lagerstätte must consider the palaeoredox conditions revealed in this study. With the dispensing of anoxic bottom waters as a requirement for BST preservation, other models proposing a role for clay minerals, the presence of hypersaline brines and the actions of Fe‐reducing bacteria as mechanisms for exceptional preservation warrant renewed consideration.     

Effect of anoxia and hypoxia on brain lipid metabolism

Lipid metabolism in rat brain was investigated in mild hypoxia (5–7% O₂ in nitrogen), which is associated with no apparent change in energy metabolism, and in severe anoxic conditions (ischemic anoxia), which are associated with a rapid decrease in ATP and oxygen content in brain. When brain slices were incubated with labeled glucose or acetate, the amount of labeled CO₂ produced was no different in experimental and control conditions, but the incorporation of radioactivity into brain lipids was decreased in all hypoxic and anoxic conditions. Interestingly, the incorporation of label from [¹⁴C]glucose into phosphatidylinositols was specifically inhibited by both hypoxic conditions but not by conditions associated with anoxia. The incorporation of the same labeled precursor, i.e., [¹⁴C]glucose, into fatty acids was elevated in ischemic anoxia but reduced after mild hypoxia. Because of the obvious differences in oxygen utilization in brain in anoxic and hypoxic conditions, we believe that the observed disturbances in lipid metabolism may be due to factors other than those that arise from oxygen deficiency alone.