The paleoredox context of early eukaryotic evolution: insights from the Tonian Mackenzie Mountains Supergroup, Canada.

Tonian (ca. 1000-720 Ma) marine environments are hypothesised to have experienced major redox changes coinciding with the evolution and diversification of multicellular eukaryotes. In particular, the earliest Tonian stratigraphic record features the colonisation of benthic habitats by multicellular macroscopic algae, which would have been powerful ecosystem engineers that contributed to the oxygenation of the oceans and the reorganisation of biogeochemical cycles. However, the paleoredox context of this expansion of macroalgal habitats in Tonian nearshore marine environments remains uncertain due to limited well-preserved fossils and stratigraphy. As such, the interdependent relationship between early complex life and ocean redox state is unclear. An assemblage of macrofossils including the chlorophyte macroalga Archaeochaeta guncho was recently discovered in the lower Mackenzie Mountains Supergroup in Yukon (Canada), which archives marine sedimentation from ca. 950-775 Ma, permitting investigation into environmental evolution coincident with eukaryotic ecosystem evolution and expansion. Here we present multi-proxy geochemical data from the lower Mackenzie Mountains Supergroup to constrain the paleoredox environment within which these large benthic macroalgae thrived. Two transects show evidence for basin-wide anoxic (ferruginous) oceanic conditions (i.e., high FeHR/FeT, low Fepy/FeHR), with muted redox-sensitive trace metal enrichments and possible seasonal variability. However, the weathering of sulfide minerals in the studied samples may obscure geochemical signatures of euxinic conditions. These results suggest that macroalgae colonized shallow environments in an ocean that remained dominantly anoxic with limited evidence for oxygenation until ca. 850 Ma. Collectively, these geochemical results provide novel insights into the environmental conditions surrounding the evolution and expansion of benthic macroalgae and the eventual dominance of oxygenated oceanic conditions required for the later emergence of animals.

Annelids from the Cambrian (Wuliuan Stage, Miaolingian) Spence Shale Lagerstätte of northern Utah, USA.

The Spence Shale Member of the Langston Formation in northern Utah and southern Idaho preserves generally non-biomineralized fossil assemblages referred to as the Spence Shale Lagerstätte. The biota of this Lagerstätte is dominated by panarthropods, both biomineralized and soft-bodied examples, but also preserves diverse infaunal organisms, including species of scalidophorans, echinoderms, lobopodians, stalked filter feeders, and various problematic taxa. To date, however, only a single annelid fossil, originally assigned to Canadia sp., has been described from the Spence Shale. This lone specimen and another recently collected specimen were analyzed in this study using scanning electron microscopy and energy dispersive X-ray spectrometry. The previous occurrence is reassigned to Burgessochaeta cf. B. setigera Walcott, 1911. The new fossil, however, is identified as a novel polychaete taxon, Shaihuludia shurikeni gen. et sp. nov., characterized by the presence of fused, bladed chaetae and a wide body. The occurrence of Burgessochaeta is the first outside the Burgess Shale and its vicinity, whereas Shaihuludia shurikeni gen. et sp. nov. adds to the diversity of annelids in the middle Cambrian and highlights the diversity of the Spence Shale Lagerstätte.

A modern definition of Fossil-Lagerstätten.

Fossil-Lagerstätten are amongst the most important windows onto the paleobiology of ancient ecosystems. Inconsistencies surrounding what constitutes a Lagerstätte limits our ability to compare sites and thus their scientific potential. Here, we provide a modern and utilitarian classification scheme for Konservat-Lagerstätten, allowing for more consistent and improved scientific discourse.

Resolving taphonomic and preparation biases in silicified faunas through paired acid residues and X-ray microscopy.

Paired petrography and acid maceration has shown that preferential silicification of shelly faunas can bias recovery based on taxon and body size. Here, silicified fossils from the Upper Ordovician Edinburg Formation, Strasburg Junction, Virginia, USA, were analyzed using X-ray tomographic microscopy (µCT) in conjunction with recovered residues from acid maceration of the same materials to further examine sources of potential bias. Results reveal that very small (<~1 mm) fossils are poorly resolved in µCT when scanning at lower resolutions (~30 µm), underestimating abundance of taxa including ostracods and bryozoans. Acid maceration, meanwhile, fails to recover poorly silicified fossils prone to disarticulation and/or fragmentation during digestion. Tests for patterns of breakage, however, indicate no significant size or taxonomic bias during extraction. Comparisons of individual fossils from 3-D fossil renders and maceration residues reveal patterns of fragmentation that are taxon-specific and allow the differentiation of biostratinomic and preparational breakage. Multivariate ordinations and cluster analyses of µCT and residue data in general produce concordant results but indicate that the variation in taxonomic composition of our samples is compromised by the resolvability of small size classes in µCT imaging, limiting the utility of this method for addressing paleoecological questions in these specific samples. We suggest that comparability of results will depend strongly on the sample size, taphonomic history, textural, and compositional characteristics of the samples in question, as well as µCT scan parameters. Additionally, applying these methods to different deposits will test the general applicability of the conclusions drawn on the relative strengths and weaknesses of the methods.

Herpetogaster collinsi from the Cambrian of China elucidates the dispersal and palaeogeographic distribution of early deuterostomes and the origin of the ambulacrarian larva.

The Cambrian Radiation represents one of the largest diversification events in Earth history. While the resulting taxonomic diversity is exceptional, relatively few of these novel species can be traced outside the boundaries of a single palaeocontinent. Many of those species with cosmopolitan distributions were likely active swimmers, presenting opportunity and means to conquer new areas, but this would not have been the case for sessile organisms. Herpetogaster is a lower to middle Cambrian (Series 2-Miaolingian, Stage 3-Wuliuan) genus of sessile, stalked, filter-feeding deuterostomes with two species, H. collinsi and H. haiyanensis, known respectively from Laurentia and Gondwana. Here, we expand the distribution of H. collinsi to Gondwana with newly discovered specimens from the Balang Formation of Hunan, China. This discovery raises questions on the origin of the genus and how sessile organisms were able to disperse over such a broad distance in the lower Cambrian. As Herpetogaster has been recovered at the base of the Ambulacrarian tree in recent phylogenies, a planktonic larval stage is suggested, which implies, that the last common ancestor of the Ambulacraria might have already had planktonic larvae or that such larvae developed multiple times within the Ambulacraria.

The role of iron in the formation of Ediacaran 'death masks'.

The Ediacara biota are an enigmatic group of Neoproterozoic soft-bodied fossils that mark the first major radiation of complex eukaryotic and macroscopic life. These fossils are thought to have been preserved via pyritic "death masks" mediated by seafloor microbial mats, though little about the chemical constraints of this preservational pathway is known, in particular surrounding the role of bioavailable iron in death mask formation and preservational fidelity. In this study, we perform decay experiments on both diploblastic and triploblastic animals under a range of simulated sedimentary iron concentrations, in order to characterize the role of iron in the preservation of Ediacaran organisms. After 28 days of decay, we demonstrate the first convincing "death masks" produced under experimental laboratory conditions composed of iron sulfide and probable oxide veneers. Moreover, our results demonstrate that the abundance of iron in experiments is not the sole control on death mask formation, but also tissue histology and the availability of nucleation sites. This illustrates that Ediacaran preservation via microbial death masks need not be a "perfect storm" of paleoenvironmental porewater and sediment chemistry, but instead can occur under a range of conditions.

Paleontology: Paleogastronomy in the Ediacaran.

Preserved lipid biomarkers provide a unique window onto the biology of fossil organisms. A new study uses lipids to assess the gut contents of three fossils from the 'dawn of animals' in the Ediacaran, with implications for their feeding behaviors.

Preservation of early Tonian macroalgal fossils from the Dolores Creek Formation, Yukon.

The rise of eukaryotic macroalgae in the late Mesoproterozoic to early Neoproterozoic was a critical development in Earth's history that triggered dramatic changes in biogeochemical cycles and benthic habitats, ultimately resulting in ecosystems habitable to animals. However, evidence of the diversification and expansion of macroalgae is limited by a biased fossil record. Non-mineralizing organisms are rarely preserved, occurring only in exceptional environments that favor fossilization. Investigating the taphonomy of well-preserved macroalgae will aid in identifying these target environments, allowing ecological trends to be disentangled from taphonomic overprints. Here we describe the taphonomy of macroalgal fossils from the Tonian Dolores Creek Formation (ca. 950 Ma) of northwestern Canada (Yukon Territory) that preserves cm-scale macroalgae. Analytical microscopy, including scanning electron microscopy and tomographic x-ray microscopy, was used to investigate fossil preservation, which was the result of a combination of pyritization and aluminosilicification, similar to accessory mineralization observed in Paleozoic Burgess Shale-type fossils. These new Neoproterozoic fossils help to bridge a gap in the fossil record of early algae, offer a link between the fossil and molecular record, and provide new insights into evolution during the Tonian Period, when many eukaryotic lineages are predicted to have diversified.

Global controls on phosphatization of fossils during the toarcian oceanic anoxic event.

Konservat-Lagerstätten-deposits with exceptionally preserved fossils-vary in abundance across geographic and stratigraphic space due to paleoenvironmental heterogeneity. While oceanic anoxic events (OAEs) may have promoted preservation of marine lagerstätten, the environmental controls on their taphonomy remain unclear. Here, we provide new data on the mineralization of fossils in three Lower Jurassic Lagerstätten-Strawberry Bank (UK), Ya Ha Tinda (Canada), and Posidonia Shale (Germany) -and test the hypothesis that they were preserved under similar conditions. Biostratigraphy indicates that all three Lagerstätten were deposited during the Toarcian OAE (TOAE), and scanning electron microscopy (SEM) and energy dispersive X-ray spectroscopy (EDS) show that each deposit contains a variety of taxa preserved as phosphatized skeletons and tissues. Thus, despite their geographic and paleoenvironmental differences, all of these Lagerstätten were deposited in settings conducive to phosphatization, indicating that the TOAE fostered exceptional preservation in marine settings around the world. Phosphatization may have been fueled by phosphate delivery from climatically-driven sea level change and continental weathering, with anoxic basins acting as phosphorus traps.

Discovery of bilaterian-type through-guts in cloudinomorphs from the terminal Ediacaran Period.

The fossil record of the terminal Ediacaran Period is typified by the iconic index fossil Cloudina and its relatives. These tube-dwellers are presumed to be primitive metazoans, but resolving their phylogenetic identity has remained a point of contention. The root of the problem is a lack of diagnostic features; that is, phylogenetic interpretations have largely centered on the only available source of information-their external tubes. Here, using tomographic analyses of fossils from the Wood Canyon Formation (Nevada, USA), we report evidence of recognizable soft tissues within their external tubes. Although alternative interpretations are plausible, these internal cylindrical structures may be most appropriately interpreted as digestive tracts, which would be, to date, the earliest-known occurrence of such features in the fossil record. If this interpretation is correct, their nature as one-way through-guts not only provides evidence for establishing these fossils as definitive bilaterians but also has implications for the long-debated phylogenetic position of the broader cloudinomorphs.

Ultrastructure of Ediacaran cloudinids suggests diverse taphonomic histories and affinities with non-biomineralized annelids.

Cloudinids have long been considered the earliest biomineralizing metazoans, but their affinities have remained contentious and undetermined. Based on well-preserved ultrastructures of two taxa, we here propose new interpretations regarding both their extent of original biomineralization and their phylogenetic affinity. One of these taxa is a new cloudinid from Mongolia, Zuunia chimidtsereni gen. et sp. nov., which exhibits key characteristics of submicrometric kerogenous lamellae, plastic tube-wall deformation, and tube-wall delamination. Multiple carbonaceous lamellae are also discovered in Cloudina from Namibia and Paraguay, which we interpret to have originated from chitinous or collagenous fabrics. We deduce that these cloudinids were predominantly originally organic (chitinous or collagenous), and postmortem decay and taphonomic mineralization resulted in the formation of aragonite and/or calcite. Further, based on our ultrastructural characterization and other morphological similarities, we suggest that the cloudinids should most parsimoniously be assigned to annelids with originally organic tubes.

Ediacaran biozones identified with network analysis provide evidence for pulsed extinctions of early complex life.

Rocks of Ediacaran age (~635-541 Ma) contain the oldest fossils of large, complex organisms and their behaviors. These fossils document developmental and ecological innovations, and suggest that extinctions helped to shape the trajectory of early animal evolution. Conventional methods divide Ediacaran macrofossil localities into taxonomically distinct clusters, which may represent evolutionary, environmental, or preservational variation. Here, we investigate these possibilities with network analysis of body and trace fossil occurrences. By partitioning multipartite networks of taxa, paleoenvironments, and geologic formations into community units, we distinguish between biostratigraphic zones and paleoenvironmentally restricted biotopes, and provide empirically robust and statistically significant evidence for a global, cosmopolitan assemblage unique to terminal Ediacaran strata. The assemblage is taxonomically depauperate but includes fossils of recognizable eumetazoans, which lived between two episodes of biotic turnover. These turnover events were the first major extinctions of complex life and paved the way for the Cambrian radiation of animals.

Melanosome diversity and convergence in the evolution of iridescent avian feathers-Implications for paleocolor reconstruction.

Some of the most varied colors in the natural world are created by iridescent nanostructures in bird feathers, formed by layers of melanin-containing melanosomes. The morphology of melanosomes in iridescent feathers is known to vary, but the extent of this diversity, and when it evolved, is unknown. We use scanning electron microscopy to quantify the diversity of melanosome morphology in iridescent feathers from 97 extant bird species, covering 11 orders. In addition, we assess melanosome morphology in two Eocene birds, which are the stem lineages of groups that respectively exhibit hollow and flat melanosomes today. We find that iridescent feathers contain the most varied melanosome morphologies of all types of bird coloration sampled to date. Using our extended dataset, we predict iridescence in an early Eocene trogon (cf. Primotrogon) but not in the early Eocene swift Scaniacypselus, and neither exhibit the derived melanosome morphologies seen in their modern relatives. Our findings confirm that iridescence is a labile trait that has evolved convergently in several lineages extending down to paravian theropods. The dataset provides a framework to detect iridescence with more confidence in fossil taxa based on melanosome morphology.

Decoupling biogeochemical records, extinction, and environmental change during the Cambrian SPICE event.

Several positive carbon isotope excursions in Lower Paleozoic rocks, including the prominent Upper Cambrian Steptoean Positive Carbon Isotope Excursion (SPICE), are thought to reflect intermittent perturbations in the hydrosphere-biosphere system. Models explaining these secular changes are abundant, but the synchronicity and regional variation of the isotope signals are not well understood. Examination of cores across a paleodepth gradient in the Upper Cambrian central Missouri intrashelf basin (United States) reveals a time-transgressive, facies-dependent nature of the SPICE. Although the SPICE event may be a global signal, the manner in which it is recorded in rocks should and does vary as a function of facies and carbonate platform geometry. We call for a paradigm shift to better constrain facies, stratigraphic, and biostratigraphic architecture and to apply these observations to the variability in magnitude, stratigraphic extent, and timing of the SPICE signal, as well as other biogeochemical perturbations, to elucidate the complex processes driving the ocean-carbonate system.

Proteins from the past.

Some fragments of ancient protein are less prone to degradation because they bind strongly to the surfaces of minerals.

A unifying model for Neoproterozoic-Palaeozoic exceptional fossil preservation through pyritization and carbonaceous compression.

Soft-tissue fossils capture exquisite biological detail and provide our clearest views onto the rise of animals across the Ediacaran-Cambrian transition. The processes contributing to fossilization of soft tissues, however, have long been a subject of debate. The Ediacaran Gaojiashan biota displays soft-tissue preservational styles ranging from pervasive pyritization to carbonaceous compression, and thus provides an excellent opportunity to dissect the relationships between these taphonomic pathways. Here geochemical analyses of the Gaojiashan fossil Conotubus hemiannulatus show that pyrite precipitation was fuelled by the degradation of labile tissues through bacterial sulfate reduction (BSR). Pyritization initiated with nucleation on recalcitrant tube walls, proceeded centripetally, decelerated with exhaustion of labile tissues and possibly continued beneath the BSR zone. We propose that pyritization and kerogenization are regulated principally by placement and duration of the decaying organism in different microbial zones of the sediment column, which hinge on post-burial sedimentation rate and/or microbial zone thickness.

Validation of the MR simulation approach for evaluating the effects of immersion on visual analysis of volume data.

In our research agenda to study the effects of immersion (level of fidelity) on various tasks in virtual reality (VR) systems, we have found that the most generalizable findings come not from direct comparisons of different technologies, but from controlled simulations of those technologies. We call this the mixed reality (MR) simulation approach. However, the validity of MR simulation, especially when different simulator platforms are used, can be questioned. In this paper, we report the results of an experiment examining the effects of field of regard (FOR) and head tracking on the analysis of volume visualized micro-CT datasets, and compare them with those from a previous study. The original study used a CAVE-like display as the MR simulator platform, while the present study used a high-end head-mounted display (HMD). Out of the 24 combinations of system characteristics and tasks tested on the two platforms, we found that the results produced by the two different MR simulators were similar in 20 cases. However, only one of the significant effects found in the original experiment for quantitative tasks was reproduced in the present study. Our observations provide evidence both for and against the validity of MR simulation, and give insight into the differences caused by different MR simulator platforms. The present experiment also examined new conditions not present in the original study, and produced new significant results, which confirm and extend previous existing knowledge on the effects of FOR and head tracking. We provide design guidelines for choosing display systems that can improve the effectiveness of volume visualization applications.

Effects of immersion on visual analysis of volume data.

Volume visualization has been widely used for decades for analyzing datasets ranging from 3D medical images to seismic data to paleontological data. Many have proposed using immersive virtual reality (VR) systems to view volume visualizations, and there is anecdotal evidence of the benefits of VR for this purpose. However, there has been very little empirical research exploring the effects of higher levels of immersion for volume visualization, and it is not known how various components of immersion influence the effectiveness of visualization in VR. We conducted a controlled experiment in which we studied the independent and combined effects of three components of immersion (head tracking, field of regard, and stereoscopic rendering) on the effectiveness of visualization tasks with two x-ray microscopic computed tomography datasets. We report significant benefits of analyzing volume data in an environment involving those components of immersion. We find that the benefits do not necessarily require all three components simultaneously, and that the components have variable influence on different task categories. The results of our study improve our understanding of the effects of immersion on perceived and actual task performance, and provide guidance on the choice of display systems to designers seeking to maximize the effectiveness of volume visualization applications.

Comment on "Fossilized nuclei and germination structures identify Ediacaran 'animal embryos' as encysting protists".

On the basis of putative nuclei and endospores, Huldtgren et al. (Reports, 23 December 2011, p. 1696) propose that embryo-like Doushantuo microfossils are nonmetazoan holozoans akin to mesomycetozoeans. However, both size and preservation preclude interpretation of internal structures as nuclei. Moreover, the authors may have conflated two different populations; some specimens display a pseudoparenchymatous organization incompatible with a mesomycetozoean comparison.

Ultrastructural and geochemical characterization of Archean-Paleoproterozoic graphite particles: implications for recognizing traces of life in highly metamorphosed rocks.

Abundant graphite particles occur in amphibolite-grade quartzite of the Archean-Paleoproterozoic Wutai Metamorphic Complex in the Wutaishan area of North China. Petrographic thin section observations suggest that the graphite particles occur within and between quartzite clasts and are heterogeneous in origin. Using HF maceration techniques, the Wutai graphite particles were extracted for further investigation. Laser Raman spectroscopic analysis of a population of extracted graphite discs indicated that they experienced a maximum metamorphic temperature of 513 +/- 50 degrees C, which is consistent with the metamorphic grade of the host rock and supports their indigenicity. Scanning and transmission electron microscopy revealed that the particles bear morphological features (such as hexagonal sheets of graphite crystals) related to metamorphism and crystal growth, but a small fraction of them (graphite discs) are characterized by a circular morphology, distinct marginal concentric folds, surficial wrinkles, and complex nanostructures. Ion microprobe analysis of individual graphite discs showed that their carbon isotope compositions range from -7.4 per thousand to -35.9 per thousand V-PDB (Vienna Pee Dee Belemnite), with an average of -20.3 per thousand, which is comparable to bulk analysis of extracted carbonaceous material. The range of their size, ultrastructures, and isotopic signatures suggests that the morphology and geochemistry of the Wutai graphite discs were overprinted by metamorphism and their ultimate carbon source probably had diverse origins that included abiotic processes. We considered both biotic and abiotic origins of the carbon source and graphite disc morphologies and cannot falsify the possibility that some circular graphite discs characterized by marginal folds and surficial wrinkles represent deflated, compressed, and subsequently graphitized organic-walled vesicles. Together with reports by other authors of acanthomorphic acritarchs from greenschist-amphibolite-grade metamorphic rocks, this study suggests that it is worthwhile to examine carbonaceous materials preserved in highly metamorphosed rocks for possible evidence of ancient life.