Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes

The extinct echinoderm clade Stylophora consists of some of the strangest known deuterostomes. Stylophorans are known from complete, fully articulated skeletal remains from the middle Cambrian to the Pennsylvanian, but remain difficult to interpret. Their bizarre morphology, with a single appendage extending from a main body, has spawned vigorous debate over the phylogenetic significance of stylophorans, which were long considered modified but bona fide echinoderms with a feeding appendage. More recent interpretation of this appendage as a posterior “tail-like” structure has literally turned the animal back to front, leading to consideration of stylophorans as ancestral chordates, or as hemichordate-like, early echinoderms. Until now, the data feeding the debate have been restricted to evaluations of skeletal anatomy. Here, we apply novel elemental mapping technologies to describe, for the first time, soft tissue traces in stylophorans in conjunction with skeletal molds. The single stylophoran appendage contains a longitudinal canal with perpendicular, elongate extensions projecting beyond hinged biserial plates. This pattern of soft tissues compares most favorably with the hydrocoel, including a water vascular canal and tube feet found in all typical echinoderms. Presence of both calcite stereom and now, an apparent water vascular system, supports echinoderm and not hemichordate-like affinities.     

A new eocrinoid from the Guanshan Biota (Cambrian Series 2, Stage 4), with implication of the development of different attachment modes in early Cambrian

Eocrinoids are scarce in the Guanshan Biota (Cambrian Series 2, Stage 4), Yunnan Province, southwest China. Here, we introduce a new gogiid: Kunmingeocrinus cupuliformis n. gen. n. sp. which is characterized by a short stalk and a well-developed attachment disc. Preservation may indicate a weakly biomineralized body for the new taxon. Morphological similarities between the new taxon and other eocrinoids with attachment discs from Cambrian Lagerstätten of Guizhou Province (Series 2, Stage 4 and Miaolingian, Wuliuan) might suggest a similar mode of life. The eocrinoids from the Guanshan Biota possibly utilized different attachment modes.     

Tomographic techniques for the study of exceptionally preserved fossils

Three-dimensional fossils, especially those preserving soft-part anatomy, are a rich source of palaeontological information; they can, however, be difficult to work with. Imaging of serial planes through an object (tomography) allows study of both the inside and outside of three-dimensional fossils. Tomography may be performed using physical grinding or sawing coupled with photography, through optical techniques of serial focusing, or using a variety of scanning technologies such as neutron tomography, magnetic resonance imaging and most usefully X-ray computed tomography. This latter technique is applicable at a variety of scales, and when combined with a synchrotron X-ray source can produce very high-quality data that may be augmented by phase-contrast information to enhance contrast. Tomographic data can be visualized in several ways, the most effective of which is the production of isosurface-based 'virtual fossils' that can be manipulated and dissected interactively.     

Three-dimensional anatomy of the Tully monster casts doubt on its presumed vertebrate affinities

Tullimonstrum gregarium, also known as the Tully monster, is a well-known phylogenetic enigma, fossils of which have been found only in the Mazon Creek Lagerstätte. The affinities of Tullimonstrum have been debated since its discovery in 1966, because its peculiar morphology with stalked eyes and a proboscis cannot easily be compared with any known animal morphotypes. Recently, the possibility that Tullimonstrum was a vertebrate has attracted much attention, and it has been postulated that Tullimonstrum might fill a gap in the fossil record of early vertebrates, providing important insights into vertebrate evolutionary history. With the hope of resolving this debate, we collected 3D surface data from 153 specimens of Tullimonstrum using a high-resolution laser 3D scanner and conducted x-ray micro-computed tomographic (μCT) analysis of stylets in the proboscis. Our investigation of the resulting comprehensive 3D morphological dataset revealed that structures previously regarded as myomeres, tri-lobed brain, tectal cartilages and fin rays are not comparable with those of vertebrates. These results raise further doubts about its vertebrate affinities, and suggest that Tullimonstrum may have been either a non-vertebrate chordate or a protostome.     

Rusophycus carleyi (James, 1885), Trace Fossils from the Lower Ordovician of Southern Morocco,and the Trilobites that Made Them

Thin-bedded, pyrite-rich, fine sandstones and mudstones of the Floian-Dapingian Upper Fezouata Formation contain abundant trace fossils Rusophycus carleyi in close association with a species of the asaphid trilobite Asaphellus. The sizes and shapes of this trilobite and the traces match closely. Five specimens have even been found where an articulated specimen of Asaphellus appears to be directly located over a specimen of Rusophycus carleyi within a thin bed of sandstone, suggesting that the trilobite animal may have been trapped on top of a trace that it had just made. Such intimate associations between a putative tracemaker and a trace are rare in the fossil record and particularly rare for Trilobita. The number of coxal impressions that form part of R. carleyi, eleven, matches the number expected for an asaphid trilobite (one for each of eight thoracic segments and one for each of three post-oral cephalic appendages). Impressions of the hypostome, thoracic tip impressions, cephalic margin, and pygidial margin in a few of the traces also match those of this asaphid trilobite. R. carleyi has been found in Ordovician strata of other parts of the world in association with asaphid trilobites.     

Nanometer‐scale characterization of exceptionally preserved bacterial fossils in Paleocene phosphorites from Ouled Abdoun (Morocco)

Micrometer‐sized spherical and rod‐shaped forms have been reported in many phosphorites and often interpreted as microbes fossilized by apatite, based on their morphologic resemblance with modern bacteria inferred by scanning electron microscopy (SEM) observations. This interpretation supports models involving bacteria in the formation of phosphorites. Here, we studied a phosphatic coprolite of Paleocene age originating from the Ouled Abdoun phosphate basin (Morocco) down to the nanometer‐scale using focused ion beam milling, transmission electron microscopy (TEM), and scanning transmission x‐ray microscopy (STXM) coupled with x‐ray absorption near‐edge structure spectroscopy (XANES). The coprolite, exclusively composed of francolite (a carbonate‐fluroapatite), is formed by the accumulation of spherical objects, delimited by a thin envelope, and whose apparent diameters are between 0.5 and 3 μm. The envelope of the spheres is composed of a continuous crown dense to electrons, which measures 20–40 nm in thickness. It is surrounded by two thinner layers that are more porous and transparent to electrons and enriched in organic carbon. The observed spherical objects are very similar with bacteria encrusting in hydroxyapatite as observed in laboratory experiments. We suggest that they are Gram‐negative bacteria fossilized by francolite, the precipitation of which started within the periplasm of the cells. We discuss the role of bacteria in the fossilization mechanism and propose that they could have played an active role in the formation of francolite. This study shows that ancient phosphorites can contain fossil biological subcellular structures as fine as a bacterial periplasm. Moreover, we demonstrate that while morphological information provided by SEM analyses is valuable, the use of additional nanoscale analyses is a powerful approach to help inferring the biogenicity of biomorphs found in phosphorites. A more systematic use of this approach could considerably improve our knowledge and understanding of the microfossils present in the geological record.     

Fossil liberation: a model to explain high biodiversity in the Triassic Cassian Formation

With 1429 animal species, the Triassic Cassian Formation in the Dolomites, Southern Alps (Italy), yields the highest species richness reported from any spatially constrained pre-Quaternary formation known to science. The high preserved diversity is partly attributable to a high primary diversity governed by the tropical setting, increasing alpha diversity, and the breadth of habitats spurring beta diversity. More important is the excellent preservation of fossils and the ease with which they can be extracted from the poorly lithified sediments. We propose the term ‘liberation Lagerstätten’ to capture this preservational window. In contrast to conservation Lagerstätten, liberation Lagerstätten like the Cassian Formation originate from normal marine conditions but low-grade diagenesis. Molluscs contribute substantially to species richness, comprising 67% of all invertebrate species in the Cassian Formation. The gastropod dominance (39% of all species) is nearly as great as in Recent tropical settings, contradicting the concept of a substantial Cenozoic rise.     

Facies of a Lower Ordovician carbonate shelf (Mungok Formation: Taebaeksan Basin,Korea)

Summary The lithologic associations within the Lower Ordovician Mungok Formation in Korea define four depositional facies that formed across a continental margin fringing the Sino-Korean block: these facies represent lagoonal/restricted marine, shoal, inner shelf, and outer shelf environments. The stacking pattern of these facies reveals two systems tracts composed of five depositional sequences. The lower highstand systems tract consists of the lagoonal/restricted marine and shoal facies, whereas the upper lowstand systems tract comprises, in ascending order, inner shelf, outer shelf, and inner shelf facies. Three trilobite biofacies are recognized in the Mungok Formation: i.e.,Yosimuraspis, Kainella, andShumardia biofacies in ascending order. TheYosimuraspis Biofacies is dominated byYosimuraspis but also containsJujuyaspis andElkanaspis. The predominance of the endemic eponymous taxon suggests a lagoonal/restricted marine environment. The nearly monotaxicKainella Biofacies, which comprises pandemic genera such asKainella and occasionallyLeiostegium, may represent a less restricted environment than theYosimuraspis Biofacies. TheShumardia Biofacies occurs in the marlstone/shale lithofacies through relatively thick stratigraphic interval and is dominated by cosmopolitan trilobite taxa with some endemic species. The lithofacies and cosmopolitan trilobite assemblage of theShumardia Biofacies indicate that it occupied an outer shelf environment. The vertical succession of lithofacies and trilobite biofacies in the Mungok Formation records in general a shift from a restricted, shallow water environment to deeper-water environment.     

An exceptional record of Cambrian trilobite moulting behaviour preserved in the Emu Bay Shale,South Australia

Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock 1964 being extremely abundant, and the larger Redlichia takooensis Lu 1950 , being common. Many specimens within the EBS represent complete moulted exoskeletons, which is unusual for Cambrian fossil deposits. The abundance of complete moults provides an excellent record that has allowed the recognition of various recurrent moult configurations for both species, enabling the inference of movement sequences required to produce such arrangements. Moult configurations of E. bilobata are characterized by slight displacement of the joined rostral plate and librigenae, often accompanied by detachment of the cranidium, suggesting ecdysis was achieved by anterior withdrawal via opening of the cephalic sutures. Moulting in R. takooensis often followed the same method, but configurations show greater displacement of cephalic sclerites, suggesting more vigorous movement by the animal during moulting. Both species also show rare examples of Salter's configuration, with the entire cephalon anteriorly inverted, and several other unusual configurations. These results indicate that moulting in trilobites was a more variable process than originally thought. In contrast, other Cambrian Konservat‐Lagerstätten with an abundance of trilobites (e.g. Wheeler Shale, USA, and Mount Stephen Trilobite Beds, Canada) show larger numbers of ‘axial shields’ and isolated sclerites, often interpreted as disarticulated exuviae. This points to a higher level of disturbance from factors, such as animal activity, depositional processes or water movement, compared to that of the EBS, where quiescent conditions and intermittent seafloor anoxia contributed to an unparalleled trilobite moulting record.     

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.     

Treasure with blood: On the discovery of Traumatocrinus (Echinodermata,Triassic) crowns in China

A team of three paleontologists led by the important Triassic worker Te-You Hsu were killed in Guizhou during a field mapping project in 1944. A.T. Mu studied Hsu's crinoid collection and named a new species Traumatocrinus hsui after him in 1949. The first edition of crinoid Treatise did not acknowledge Mu's work and placed Traumatocrinus as junior synonym of Encrinus. However, it is now regarded as a valid taxon in the new edition published in 2011. During this study, original specimens reported by Mu were re-examined. Among them, 21 specimens survived to this day, and the rest of specimens mentioned in Mu were lost during the Second World War. The collection is illustrated with modern standards for the first time. Despite the hard working conditions endured by the Chinese workers during and shortly after the Second World War, this collection, including eleven slabs of crowns, one cup, eight columnals, and one holdfast curated in Nanjing, is the basis for family Traumatocrinidae.     

Trace fossils analysis of fluvial to open marine transitional sediments: Example from the Upper Devonian (Geirud Formation), Central Alborz,Iran

This study integrates ichnological and sedimentological data to interpret depositional environments of the mixed siliciclastic-carbonate fluvial to marine sediments of the Geirud Formation (Upper Devonian) in the central Alborz, northern Iran. Lithofacies analysis shows that these sediments are deposited in fluvial, tidal, shoreface, and shelf environments. Fluvial and tidal deposits are characterized by the presence of bi- to multi-directional cross beddings, reverse directional current ripples, low angle cross beddings, and herringbone cross beddings, with a few scattered Skolithos and Palaeophycus. Shoreface sediments, accumulated in a storm-influenced setting, are characterized by a preferentially interface and low diversity Cruziana ichnoassemblage (Rhizocorallium, Thalassinoides, Palaeophycus, Chondrites, and Ophiomorpha). In contrast to the fluvial-tidal assemblage, the storm-influenced shelf sediments display a highly diversified, mixture of dwelling and feeding forms (Arenicolites, Protovirgularia, Diplocraterion, Palaeophycus, Thalassinoides, Chondrites, and Helminthopsis), reflecting the presence of adequate food resources both in substrate and water column under normal salinity conditions. A fluvial-shelf replacement of the weakly to scarcely bioturbated sediments by the Rhizocorallium-Thalassinoides suite (Cruziana) – Chondrites-Helminthopsis (distal Cruziana) suite from the lower to upper parts of the succession clearly indicates an overall deepening upward in the Geirud Formation. In contrast to the lower part, generally of restricted environment, the upper part of the succession mainly shows open marine conditions. Ichnofabric development is controlled primarily by depositional conditions, e.g., bottom water oxygenation, sediment type, food abundance, and hydrodynamic level, which all exert control on substrate colonization style.     

Variable preservation potential and richness in the fossil record of vertebrates

Variation in preservation and sampling probability clouds our estimates of past biodiversity. The most extreme examples are Lagerstätten faunas and floras. Although such deposits provide a wealth of information and represent true richness better than other deposits, they can create misleading diversity peaks because of their species richness. Here, we investigate how Lagerstätten formations add to time series of vertebrate richness in the UK, Germany and China. The first two nations are associated with well-studied fossil records and the last is a country where palaeontology has a much shorter history; all three nations include noted Lagerstätten in their fossil records. Lagerstätten provide a larger proportion of China's sampled richness than in Germany or the UK, despite comprising a smaller proportion of its fossiliferous deposits. The proportions of taxa that are unique to Lagerstätten vary through time and between countries. Further, in all regions, we find little overlap between the taxa occurring in Lagerstätten and in ‘ordinary’ formations within the same time bin, indicating that Lagerstätten preserve unusual faunas. As expected, fragile taxa make up a greater proportion of richness in Lagerstätten than the remainder of the fossil record. Surprisingly, we find that Lagerstätten account for a minority of peaks in the palaeodiversity curves of all vertebrates (18% in the UK; 36% in Germany and China), and Lagerstätten count is generally not a good overall predictor of the palaeodiversity signal. Vastly different sampling probabilities through taxa, locations and time require serious consideration when analysing palaeodiversity curves.     

Geochemistry and palaeogeography of upper Ordovician glaciogenic sedimentary rocks in the Table Mountain Group, South Africa

Evidence for a late Ordovician glaciation is best known from northern Africa and locally in South Africa, and western South America. The South African glacial deposits (Pakhuis Formation of the Table Mountain Group) are present in the western part of the Cape Fold Belt. Two thin diamictites are succeeded by a coarsening upward shale-siltstone unit known as the Cedarberg Formation. These glaciogenic rocks are associated with one of the world's greatest accumulations of mature sandstones. The diamictites are under- and overlain by shales and sandstones that have undergone extensive alteration, presumably as a result of chemical weathering. Some diamictite samples have low values for a Chemical Index of Alteration (CIA), in keeping with their inferred glacial origin. Immediately above the glaciogenic diamictites, black shales, comprising the basal Soom Member of the Cedarberg Formation preserve an unusual soft bodied fauna and are enriched in Mo, U and other trace elements. The unusual chemical composition of these shales is attributed to starved basin conditions and a reducing environment during a rapid sea level rise that accompanied demise of the late Ordovician glaciers. The late Ordovician glacial deposits of western South Africa are widely regarded as the distal deposits of a large continental ice sheet centred on North Africa but they may have been partly derived from piedmont glaciers that descended from highlands formed as a result of late Ordovician orogeny.     

Palaeoecological and sedimentological characteristics of the Lower Tortonian scleractinian reef corals of Gavdos Island,southern Greece

Early Late Miocene coral assemblages from five outcrops of Gavdos Island, Southern Greece, are investigated with respect to their palaeoecological implications. Small patch reefs with Porites assemblage are a common feature of the low-diversity coral occurrences. The determined hermatypic colonies indicate a nearshore palaeoecosystem prevailing in a tropical to subtropical coastal sea at depths ranging from 5 to 50 m with an average temperature of 22–26 °C. Microfacially, the studied Scleractinian patch reefs are represented by Coral Framestones-Floatstones. The reefal facies has been affected by syndepositional processes (boring activities-micritization), as well as by post-depositional diagenesis in the meteoric realm (dissolution, cementation and intense pedogenesis). The palaeoecological and sedimentological analysis indicates a restricted to open-marine inner platform setting of moderate to high energy, possibly of ramp-type (inner-mid ramp). Coral reef growth took place mainly during stages of accommodation (i.e., transgressive episode, cycle 3.1 of Vail curve) and of relatively low siliciclastic input.     

Aligned trace fossils from the Utica Shale: implications for mode of life and feeding in the trilobite Triarthrus beckii

The olenid trilobite Triarthrus commonly occurs in nearly monospecific assemblages within otherwise relatively barren black shales. As such, it has been proposed that these trilobites preferred dysoxic or even anoxic habitats and suggested feeding habits range from predation and particle feeding to chemoautotrophism. A unique bedding‐plane assemblage of aligned traces that grade from Rusophycus to Cruziana, with associated Triarthrus beckii carcasses, are described from three localities in the Indian Castle Formation (upper Utica Shale) just below a K‐bentonite bed. Although few body fossils are preserved, it is clear that the 15‐cm‐thick, laterally extensive Thruway K‐bentonite created a unique taphonomic window that preserved the activities of numerous olenid trilobites. Rusophycus and Cruziana, consistently observed in densities above 100 trackways/m², were excavated by the trilobites into the upper surface of a micro‐graded bed, likely a distal turbidite deposit. Sedimentological and trace metal data (Mo, Mn, V and U) support dysoxic but not persistently anoxic conditions through this interval. Measurements from over 500 individual trackways have mean orientations of 259.1° (WSW) at the Myers Rd. locality and 224.59° (SW) at the Dolgeville Dam locality. These orientations are closely similar to independent sedimentological indicators of current transport direction at these sites, which indicates that the trilobites preferentially faced into the prevailing current. The trackways provide compelling evidence that Triarthrus beckii individuals were engaged in feeding, probably assisted by current transport of particles. These results do not support previous suggestions that Triarthrus may have relied upon a chemoautotrophic life habit, but are consistent with suggestions of predation or scavenging from the sediment.     

Characteristics of the Lofer cyclicity in the type locality of the Dachstein Limestone (Dachstein Plateau,Austria)

In the present paper, the results of our studies in the type locality of the Dachstein Limestone are summarised in order to contribute to the correct interpretation of the Lofer cycles. In the sections studied on the Dachstein Plateau, the boundaries of the Lofer cycles are usually erosional disconformities showing karstification features. Penetration by karstic solution was not more than a few decimetres, since during the recurrent sea-level drops the platform only slightly emerged above sea level. The reddish or greenish argillaceous carbonate interlayers (facies A) cannot be interpreted as in situ palaeosol horizons. They are tidal flat deposits consisting predominantly of subtidal carbonate mud redeposited by storms that was mixed with reworked airborne fine carbonate particles and argillite and/or reworked lateritic soil, which were accumulated on the subaerially exposed platform. Rip-ups from consolidated sediment, blackened intraclasts and skeletons of tidal flat biota may have also contributed to the sediment of facies A. Erosional boundaries of most of the investigated cycles, and definite features of karstic solution beneath the disconformities, suggest periodical drops of sea level followed by a renewed transgression. This appears to confirm the allocyclic model for the explanation of the origin of the Lofer cycles.     

‘Cambrian’ demosponges in the Ordovician of Morocco: Insights into the early evolutionary history of sponges

Cambrian spicular sponge faunas are dominated by a distinctive assemblage of demosponges and hexactinellids that are known from Burgess shale-type faunas worldwide. Most of these are previously unknown outside the Lower-Middle Cambrian (and perhaps Tremadoc) and have no obvious close relatives in later sequences. This paper describes examples of Choia sp., Pirania auraeum sp. nov. and Hamptonia christi sp. nov. from the Arenig of Morocco, associated with isolated hexactinellid spicules. A summary of the stratigraphic ranges of the major Cambrian sponge lineages is provided. These indicate an environmental contrast in the Lower Palaeozoic evolution of hexactinellids and non-lithistid demosponges, with demosponges probably undergoing cryptic diversification in nearshore environments during the Upper Ordovician.     

Characteristics of peritidal facies and evidences for subaerial exposures in Dachstein-type cyclic platform carbonates in the Transdanubian Range,Hungary

In the Late Triassic, an extremely large carbonate platform system (Dachstein-type platforms) developed on the margin of the Neotethys. On the wide inner platform cyclic peritidal, lagoonal successions were deposited. In the Transdanubian Range (Hungary), the lower part of the 1.5–2-km-thick cyclic succession (Upper Tuvalian–mid-Norian) is pervasively dolomitised, the upper part (Upper Norian–Rhaetian) is non-dolomitised; there is a transitional interval between them made up of partially dolomitised cycles. The peritidal–lagoonal cycles are commonly bounded by well-developed disconformity surfaces reflecting subaerial erosion that punctuated the marine carbonate accumulation. Truncation of the cycles was preceded by pervasive cementation of the previously deposited cycle. In the early stage of the platform evolution, tidal flat dolomitisation under semi-arid conditions led to the consolidation of the previously deposited sediments. The truncation surfaces were commonly covered by dolocretes. During the more humid Late Norian–Rhaetian period, the early cementation was followed by karstification, accumulation of wind-blown dust and pedogenesis. Erosion during regularly recurring subaerial exposure that commonly reached the previously deposited subtidal beds suggests eustatic control of the cyclicity and supports the application of an allocyclic model, even if the Milankovitch signal is imperfect.