Evidence for seafloor microbial mats and associated metazoan lifestyles in Lower Cambrian phosphorites of Southwest China

The increase in the depth and intensity of bioturbation through the Proterozoic-­Phanerozoic transition changed the substrates on which marine benthos lived from being relatively firm with a sharp sediment-water interface to having a high water content and blurry sediment-water interface. Additionally, microbial mats, once dominant on normal marine Proterozoic seafloors, were relegated to stressed settings lacking intense metazoan activity. This change in substrates has been termed the 'agronomic revolution', and its impact on benthic metazoans has been termed the 'Cambrian substrate revolution'. The shallow marine phosphorites of the Lower Cambrian Meishucun Formation of southwest China contain evidence suggestive of the presence of seafloor microbial mats. This evidence includes abundant and distinctive red-colored bedding planes enriched in heavy iron minerals and mica, interpreted as resulting from mat-decay mineralization and mica trapping by microbial mats. The radular grazing trace fossil Radulichnus is also found in this formation, indicating a firm, microbial mat-bound substrate. These radular scratches are always preserved with circular impressions around 10 cm in diameter, possibly the fossils of soft-bodied organisms. The first relatively intense bioturbation in this region is found in this formation and is dominated by horizontal Thalassinoides burrows, which could represent undermat mining behavior. The evidence for the presence of microbial mats in the Lower Cambrian Meishucun Formation, and for metazoan lifestyles associated with such mat-bound seafloors, reveals that normal marine environments dominated by typical Proterozoic-style soft substrates still existed during the Cambrian substrate revolution.     

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.     

Quantitative analysis of horizontal bioturbation from Brioverian (Ediacaran - Fortunian) deposits of Brittany (Armorican Massif,NW of France)

Ediacaran-Cambrian bioturbation on bedding planes provides physical and chemical data about the environmental conditions and biological activities of early metazoans. We propose a quantitative method to estimate horizontal disruption of the substrate using bedding-plane trace fossils from Brioverian deposits. This methodology provides the first quantitative analysis of the trace fossil assemblage from the Armorican Massif (Brittany, NW of France). The Ediacaran-Cambrian transition within the Brioverian series is characterized by the abundance of simple and horizontal trace fossils such as Helminthopsis, Helminthoidichnites, and Palaeophycus as well as rare Gordia and Spirodesmos. Recent U-Pb dating has been carried out on detrital zircon grains from the upper Brioverian series suggesting a late Ediacaran to Fortunian age of the fossiliferous deposits (ca. 550 to 530 Ma). With Arc-Gis software, we use a semi-quantitative approach to estimate the relative 2D bioturbation rate recorded on bedding planes. The dataset is used to discuss the feeding strategies and how the seafloor ecospace was colonized by the early bilaterian metazoans. This approach combines new values such as: the length of a trace fossil, the surface of a trace, the cumulative length, and the cumulative surfaces from all of the ichnofossils recorded on the same slate surface, to finally suggest that the bioturbation rate of the microbial grazers impacted the use of the seafloor ecospaces and feeding strategies through the biomats.     

Early Cambrian Teichichnus-dominated ichnofabrics and palaeoenvironmental analysis of the Caerfai Group, Southwest Wales, UK

The early Cambrian Caerfai Group of Pembrokeshire in Southwest Wales yields an intensely bioturbated Teichichnus-dominated ichnofabric, and provides evidence for the early evolution of infaunal organisms. Three units of the Caerfai Group comprise the focus of this study in sections exposed at Caerfai Bay. In the St. Non's Sandstone few primary sedimentary structures are observed, bedding being predominantly massive with subordinate parallel and current ripple cross-lamination. The topmost St. Non's Sandstone contains common soft-sediment deformation phenomena, and collectively deposition is believed to have taken place in the proximal subaqueous reaches of a steep-fronted delta system. Teichichnus is the dominant trace, with subordinate Planolites, Palaeophycus and possible Rhizocorallium.The overlying Caerfai Bay Shales were deposited from mass flows with evidence of downslope creep suggesting deposition on a steep delta front. Sandstone debrites are common, as are tuffaceous beds with evidence of reworking by bottom currents. Sandstone and tuff event beds contain common Skolithos and Arenicolites with less frequent Planolites. Interbedded mudstones contain infrequent Teichichnus.The Caerfai Bay Shales coarsen and thicken-upwards into the Caerbwdy Sandstone. Debrite sandstone beds dominate, and bed thickening upsection indicates progradation of the delta system. The dominant trace fossil within the unit is Teichichnus which is constrained to the top few centimetres of individual sandstone beds. Less common are Planolites and Palaeophycus.The size and abundance of Teichichnus in the Caerfai Group are similar to, or exceed those in contemporaneous sections in Avalonia and Baltica. Local environmental factors were probably the primary control on trace fossil distribution. Substrate and salinity are proposed as the main influences on ichnodiversity and size in the Caerfai Group. Teichichnus is most abundant in the St. Non's Sandstone, where the proximal reaches of the delta experienced reduced salinity and possibly higher oxygen levels. Burrow diameter progressively decreases upsection, and bioturbation becomes less pervasive in subsequent units of the Caerfai Group. This indicates that by the Cambrian Stage 2, significant advances into reduced salinity environments had occurred, along with the dispersal of the Teichichnus producer throughout the Avalonia-Baltica seaway. Correlation within Avalonia indicates that a transgression in the Cambrian Stage 2 initiated the deposition of the Caerfai Group marking the onset of the Welsh Basin.     

Coexisting living stromatolites and infaunal metazoans

Microbialites, bioaccretionary structures formed during the growth and metabolism of microorganisms (principally cyanobacteria) were the dominant lifeform in shallow late-Archean and Proterozoic oceans. During the Cambrian radiation of metazoan life, which began ~540 Mya, microbialite abundance and diversity further declined following a peak in the Mesoproterozoic. Notwithstanding contention, grazing and bioturbation effects of metazoans have been hypothesized as the dominant driver of modern microbialite scarcity. However, this metazoan–microbialite exclusion has not been fully explored in the few extant microbialites. Here we provide further evidence showing that living marine layered microbialites (stromatolites) coexist with a persistent assemblage of benthic macro-invertebrates, as has previously been demonstrated in some thrombolitic (clotted) microbialites. Surprisingly, these metazoans have active habits, such as burrowing, which should be expected to disrupt the layered matrix. As other studies have shown, through a network of burrows, metazoans can exploit local diurnal oxygen refugia within microbialites as well as escape predation. Our results, therefore, add novel evidence in support of the hypotheses that geologically, metazoans are not always incompatible with stromatolites, while ecologically, microbialites may act as micro-refugia for modern metazoans and historically have performed a similar inferred role in past ecosystems.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

Does bioturbation by a benthic fish modify the effects of sediment contamination on saltmarsh benthic microalgae and meiofauna?

A microcosm experiment was conducted using a replicated factorial design to determine if a benthic fish modifies the effects of sediment-bound contaminants (diesel fuel and two levels of a Cu, Cr, Pb, Hg and Cd mixture) on saltmarsh benthic primary producers and consumers. The naked goby, Gobiosoma bosc, a burrowing fish that preys on small macrofauna, was added to experimental microcosms. Goby burrowing and foraging significantly increased turbidity and disrupted the sediment surface. Results were typified by complex and varied responses with many statistically significant effects and interactions among treatments. Although G. bosc modified the responses to both diesel and metal pollution in invertebrates (but not microalgae), bioturbation did not increase or decrease the toxic effects of metals or diesel, and diesel-metal interactions did not vary in response to G. bosc. Specifically, G. bosc inhibited a trend toward diesel-induced increases in nematode abundance, and diesel toxicity inhibited increases in ostracod abundance stimulated by G. bosc. Diatoms, nematodes and the copepod Pseudostenhelia wellsi decreased in treatments with G. bosc. However, G. bosc lead to increases in cyanobacteria and ostracods and a trend toward increases in the copepod Pseudobradya sp. Our findings suggest that microcosm experiments are potentially poor mimics of natural systems without bioturbation. Conclusions about the direct and indirect effects of contaminants may differ with and without bioturbation. Finally, our work suggests that the direct effects of toxicants may inhibit or mask bioturbation effects that stimulate population growth of some meiofauna.     

Effect of fish predation on intertidal benthic fauna is modified by crab bioturbation

The burrowing crab Chasmagnathus granulatus is an important bioturbator in SW Atlantic estuaries where they generate dense and extended intertidal beds. Its bioturbation leads to profound changes in the structure, quality and dynamics of sediments with concomitant impacts on the entire benthic community. In this study, we evaluate whether the presence of this crab affects the predator-prey interaction between juvenile fishes and their benthic prey. Gut content and benthic prey selection by juvenile fishes inside and outside crab beds were evaluated, and predation effect was experimentally contrasted between areas using fish exclosures. The results show that in crab beds the percentage of fish with empty guts was lower and the number of polychaetes consumed by fish higher than outside crab beds. The silverside Odontesthes argentinensis and the catfish Pimelodella laticeps fed on larger polychaetes outside than inside crab bed areas, while the white mouth croaker Micropogonias furnieri preyed upon larger polychaetes inside crab beds. In addition, field experiments shows that fish predation decreases polychaete abundances only in crab beds. These results suggest that crab bioturbation facilitate fish predation on benthic prey.     

Ediacaran–Cambrian bioturbation did not extensively oxygenate sediments in shallow marine ecosystems

The radiation of bioturbation during the Ediacaran–Cambrian transition has long been hypothesized to have oxygenated sediments, triggering an expansion of the habitable benthic zone and promoting increased infaunal tiering in early Paleozoic benthic communities. However, the effects of bioturbation on sediment oxygen are underexplored with respect to the importance of biomixing and bioirrigation, two bioturbation processes which can have opposite effects on sediment redox chemistry. We categorized trace fossils from the Ediacaran and Terreneuvian as biomixing or bioirrigation fossils and integrated sedimentological proxies for bioturbation intensity with biogeochemical modeling to simulate oxygen penetration depths through the Ediacaran–Cambrian transition. Ultimately, we find that despite dramatic increases in ichnodiversity in the Terreneuvian, biomixing remains the dominant bioturbation behavior, and in contrast to traditional assumptions, Ediacaran–Cambrian bioturbation was unlikely to have resulted in extensive oxygenation of shallow marine sediments globally.     

The origin of the animals and a ‘Savannah’ hypothesis for early bilaterian evolution

The earliest evolution of the animals remains a taxing biological problem, as all extant clades are highly derived and the fossil record is not usually considered to be helpful. The rise of the bilaterian animals recorded in the fossil record, commonly known as the ‘Cambrian explosion’, is one of the most significant moments in evolutionary history, and was an event that transformed first marine and then terrestrial environments. We review the phylogeny of early animals and other opisthokonts, and the affinities of the earliest large complex fossils, the so‐called ‘Ediacaran’ taxa. We conclude, based on a variety of lines of evidence, that their affinities most likely lie in various stem groups to large metazoan groupings; a new grouping, the Apoikozoa, is erected to encompass Metazoa and Choanoflagellata. The earliest reasonable fossil evidence for total‐group bilaterians comes from undisputed complex trace fossils that are younger than about 560 Ma, and these diversify greatly as the Ediacaran–Cambrian boundary is crossed a few million years later. It is generally considered that as the bilaterians diversified after this time, their burrowing behaviour destroyed the cyanobacterial mat‐dominated substrates that the enigmatic Ediacaran taxa were associated with, the so‐called ‘Cambrian substrate revolution’, leading to the loss of almost all Ediacara‐aspect diversity in the Cambrian. Why, though, did the energetically expensive and functionally complex burrowing mode of life so typical of later bilaterians arise? Here we propose a much more positive relationship between late‐Ediacaran ecologies and the rise of the bilaterians, with the largely static Ediacaran taxa acting as points of concentration of organic matter both above and below the sediment surface. The breaking of the uniformity of organic carbon availability would have signalled a decisive shift away from the essentially static and monotonous earlier Ediacaran world into the dynamic and burrowing world of the Cambrian. The Ediacaran biota thus played an enabling role in bilaterian evolution similar to that proposed for the Savannah environment for human evolution and bipedality. Rather than being obliterated by the rise of the bilaterians, the subtle remnants of Ediacara‐style taxa within the Cambrian suggest that they remained significant components of Phanerozoic communities, even though at some point their enabling role for bilaterian evolution was presumably taken over by bilaterians or other metazoans. Bilaterian evolution was thus an essentially benthic event that only later impacted the planktonic environment and the style of organic export to the sea floor.     

Trilobites from the lower and lowermost middle Cambrian of the Kiya River reference section (Kuznetsk Alatau)

The Kiya River reference section is characterized by trilohites confined to units X–XV of the Lower Cambrian Usa Formation and to the base of the lowermost Middle Cambrian Berikul Formation. Trilobites are represented by 130 species assigned to 77 genera. The given paper describes 38 species belonging to 27 genera. 16 species, 2 genera and one subfamily are new. All of them occur in the Usa Formation and in the Lower Cambrian pebbles of the Berikul Formation. Biostratigraphic analysis shows that the trilobite fauna contains characteristic trilobite assemblage elements of the Lower Cambrian regional stages in the Altay-Sayan Fold Belt, i.e. the Kiya and Kameshki regional stages (Atdabanian Stage), the Sanashtykgol regional stage (Botomian Stage), the Obruchev regional stage (Toyonian Stage). Table I shows trilobite species composition and vertical distribution in the reference section.     

Palaeoecological significance of calcimicrobial communities during ramp evolution: An example from the Lower Cambrian of Germany

Summary The palaeoecology of calcimicrobial communities from the only Gondwana-related Lower Cambrian in Central Europe (eastern Germany, carbonates and siliciclastics) has been studied. Six morphological groups of calcimicrobes are described. Some of them show a significant ability for sediment stabilization and construction of biohermal reef mounds. Other types of calcimicrobes were more common in biostromal thickets. Some of them were capable of populating different environments, growing in different modes and on different substrates. AnEpiphyton-archaeocyathan reef mound is described, illustrating the importance of calcimicrobes for mound formation. The fossil communities together with a complex of sedimentary features allow a reconstruction of the depositional history of the environment. Based on comparison with similar Gondwanan Lower Cambrian successions (Sardinia/Italy, Spain) and facies development a model is proposed describing the sedimentary history. Three depositional stages are distinguished: (1) deep subtidal ramp, (2) shallow subtidal ramp, (3) shallow subtidal to intertidal mixed siliciclastic-carbonate ramp with migrating oolitic shoals. In spite of similarities with the deposits in southern Europe, some distinct differences exist with respect to the succession of facies, the completeness of the sections, the fossil spectrum, and the nature of the siliciclastic sediments. For the German Lower Cambrian, a facies development from a low energy deep environment to a high energy shallow environment (partly restricted and with some evaporites) can be reconstructed. As compared with Sardinia and Spain, the depositional environment of the eastern German Lower Cambrian successions was predominantly characterized by low-energy conditions.     

Late Cambrian hard substrate communities from Montana/Wyoming: the oldest known hardground encrusters

Hardground surfaces from the Late Cambrian Snowy Range Formation in Montana/Wyoming are the oldest known non-reefal hard substrates exhibiting encrusting fossils. These surfaces range in age from Early Franconian to early Trempealeauan. Hardgrounds were developed on slightly hummocky to planar, truncated surfaces of glauconite-rich, carbonate, flat pebble conglomerates, which were deposited during episodes of storm scouring in shallow subtidal environments of the Montana/Wyoming shelf. Snowy Range hardgrounds are encrusted by a low diversity assemblage of fossils dominated by simple discoidal holdfasts of pelmatozoans, probably crinoids, and including small conical spongiomorph algae? and probable stromatolites. Macroborings (e.g. Trypanites) are notably absent from all hardground surfaces, although sharp-walled, vertical, cylindrical holes (borings?) occur in micrite clasts imbedded in certain flat pebble conglomerates. No evidence of faunal succession or microecologic partitioning of irregular surfaces was observed on these Cambrian hardgrounds.     

Grain-size controls on the occurrence of bioturbation

Grain size and grain-size related stresses impart a significant influence on the ichnological character of marginal-marine deposits. This is evident on the New Brunswick coastline of the Bay of Fundy, Canada, where three coarse-grained marginal-marine deposits are studied to assess grain-size controls on the occurrence and type of bioturbation. Firm mud and sand substrates exhibit the greatest diversity and density of bioturbation (i.e., bioturbation intensity). The types of organisms colonizing sands and firm-mud substrates are variable; however, the resultant trace assemblages are similar. Thixotropic muds exhibit significantly reduced trace diversity and density relative to firm mud, reflecting the additional stress placed on the organisms by the relatively soupy consistency of the sediment. A significant change in the trace assemblage occurs when sediment caliber passes the gradational sand—fine gravel boundary.Four main conclusions can be drawn from this study. First, for mixed sand and gravel, fine gravel, and coarse-gravel deposits, the degree of bioturbation (diversity ^? density) decreases more rapidly onshore (across the intertidal zone) than is noted in sand or mud deposits. Second, there is a decrease in the degree of bioturbation with increasing grain size for substrates composed of sand-sized and larger clasts. Third, burrows in gravels tend to be lined and/or robust, likely to maintain a stable environment within the burrow. Fourth, in coarse-gravel substrates or substrates with a significant component of coarse gravel, burrows are developed between the clasts and tend to be more permanent structures (than those developed in sand or mud), which are generally continuously occupied.The degree of burrowing noted in these modern gravel deposits contrasts with the relative paucity of biogenic structures reported in conglomerates preserved in the rock record. Based on the intensity of burrowing observed in the gravels, we hypothesize that ancient marginal-marine conglomerates are likely bioturbated, but that these burrows are likely distorted during burial and compaction.     

Facies control on lower Cambrian wrinkle structure development and paleoenvironmental distribution, southern Great Basin, United States

Assessing the role that physical processes play in restricting microbial mat distribution has been difficult due to the primary control of bioturbation in the modern ocean. To isolate and determine the physical controls on microbial mat distribution and preservation, a time in Earth’s history must be examined when bioturbation was not the primary control. This restricts the window of observation primarily to the Precambrian and Cambrian, which precede the development of typical Phanerozoic and modern levels of bioturbation. Lower Cambrian strata of the southern Great Basin, United States, record the widespread development of seafloor microbial mats in shallow shelf and nearshore settings. These microbial mats are recorded by wrinkle structures, which consist of millimeter-scale ridges and sinuous troughs that represent the former presence of a surface microbial mat. Wrinkle structures within these strata occur exclusively within heterolithic deposits of the offshore transition, i.e., between fair-weather wave base and storm wave base, and within heterolithic tidal-flat deposits. Wrinkle structures are not preserved in siltstone-dominated offshore deposits or amalgamated shoreface sandstones. The preservation of wrinkle structures within these environments is due to: (1) the development of microbial mats atop clean quartz-rich sands for growth and casting of the structures; and (2) the draping of the microbial mat by finer-grained sediment to inhibit erosion. The exclusion from offshore deposits may be due to a lack of sufficient sunlight, whereas the restriction from the shoreface is likely due to the amalgamation of proximal tempestites, resulting in the erosion of any incipient microbial mat development.     

Benthic foraminiferal assemblages of the Cretaceous platform carbonate succession in the Yavca area (Bolkar Mountains, S Turkey): biostratigraphy and paleoenvironments

The microbiostratigraphic analysis of the three outcrop sections from the Cretaceous inner platform carbonate succession in the Yavca area (Bolkar Mountains) allows to recognize the four local benthic foraminiferal zones. These are: (1) Voloshinoides murgensis and Praechrysalidina infracretacea Cenozone in the Lower Aptian; (2) Pseudorhapydionina dubia and Biconcava bentori Cenozone in the Middle-Upper Cenomanian; (3) Ostracoda and Miliolidae Interval Zone in the probable Turonian, represented by dolomitized limestones without any significant markers; (4) Moncharmontia compressa and Dicyclina schlumbergeri Cenozone in the Coniacian-Santonian. The benthic foraminiferal assemblages correspond to those in other areas of the Mediterranean realm, with the exception of a lack of alveolinids and orbitolinids due to unfavorable environmental conditions (inner platform, restricted shelf). After the regionally well-known emergence during the late Aptian, Albian and early Cenomanian, very shallow subtidal to intertidal conditions were re-established during the middle-late Cenomanian time. The Coniacian-Santonian benthic foraminiferal assemblage shows an increase in diversity and abundance as a result of open marine influence, confirmed by the presence of larger foraminifera (Dicyclina), Rotaliidae and radiolitid fragments. Thaumatoporella and Aeolisaccus-bearing wackestone intercalations still indicate the existence of sporadic restricted environment conditions. The Cretaceous shallow-water platform carbonate succession of the Yavca area is conformably overlain by gray pelagic limestones with calcispheres and planktonic foraminifera. The Campanian flooding of the Bolkar Da? carbonate platform resulted in drowning of the pre-existing biota and facies.     

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.     

Facies distribution patterns of some marine benthic faunas in early paleozoic platform environments

Worldwide Late Cambrian—Silurian lithofacies patterns indicate that the platforms of that time were sites of accumulation of two essentially different rocks suites: the platform carbonate rocks and the platform terrigenous rocks. Most of the platform rocks accumulated as sediments in shallow marine environments similar to those of the present but far more widely spread.Present-day marine benthic faunas are distributed in depth zones which are primarily controlled by temperature. Faunas tend to occur in substrate-related discrete clusters (communities) within each life zone; similar substrates in different depth zones commonly have different faunal associations. Individual phyletic stocks may encounter environmental optimum or near-optimum conditions in certain areas, that commonly are revealed by an abundance of species and individuals within species in each stock. Environmental optimum conditions depend upon availability of food that may be utilized, modes of feeding of the animals present, water motion, and substrate, among other factors. Organisms in past seas were distributed in patterns similar to those of the present.Carbonate platforms were particularly widespread during the latest Cambrian—Early Ordovician. Intertidal environments spread widely across those platforms during that time and characteristic faunal associations developed in them. Saukiid and related tribolites dominated latest Cambrian carbonate platform intertidal faunas. The Early Ordovician carbonate platform intertidal was dominated by archeogastropod-nautiloid cephalopod faunas. These animals were joined by tabulate corals and certain brachiopods during the latter part of the Ordovician and Silurian as prominent faunal elements in the carbonate platform intertidal—shallow subtidal. Cruziana and related trace fossils, bivalves, and certain tribolites (notably homalonotids and dalmanitids) dominated most terrigenous platform intertidal—shallow subtidal faunas of the Ordovician and Silurian.Articulate brachiopods (primarily orthoids, strophomenoids, and rhynchonelloids) appear to have been relatively prominent during the Early Ordovician in shallow subtidal environments on both carbonate and terrigenous platforms and to have spread down the bathymetric gradient into increasingly deeper subtidal areas of both platforms during the latter part of the Ordovician. Tribolites dominated faunas in relatively moderate to deep subtidal environments on both platforms during the early part of the Ordovician. They were gradually replaced by brachiopods in first the shallower, and later the deeper subtidal as dominant members of the faunas. Brachiopods (primarily pentameroids and spiriferoids) dominated nearly all Silurian warm-water subtidal environments from the shallow subtidal to the edges of the platforms.Platform uplifts in the Middle Ordovician and glacio-eustatic sea-level fluctuations in the Late Ordovician caused environmental changes across the platforms that were accompanied by marked replacements among marine benthic faunas in all environments. The distribution of Ordovician carbonate platforms and glacial deposits suggests that an Ordovician polar region may have been close to present-day equatorial Africa and that Ordovician warm temperate-tropical regions lay close to the present-day North Pole.