Exceptional Fossil Preservation Research Articles

Statistical estimation of the early to middle Ediacaran ocean redox architecture in the Yangtze block of South China

Early to middle Ediacaran organic-rich black shales host several famous fossil biotas and provide key evidence for understanding the coevolution of multicellular organisms and palaeoceanic environment. The water column redox states play critical roles in organic-rich shale deposition. Amongst multiple geochemical redox indexes, iron-speciation chemistry (FeHR/FeT, Fepy/FeHR) in shales constitutes a reliable proxy to reconstruct the first-order redox framework of the ocean basin. However, iron-speciation is a local marine redox proxy, and only compiled data collected from multiple different sedimentary facies record statistically significant changes of oxidation states within a depositional basin. Here we compiled the spatiotemporal distribution of iron-speciation data in early to middle Ediacaran black shales of the Yangtze block in South China. New high-resolution analysis on two outcrops and one drill-core reflects overall ferruginous condition and dominant euxinic condition locally interrupted by oxic states, respectively, generally indicating pervasive anoxic depositional environment for the Member II shales of the Ediacaran Doushantuo Formation in the Lower Yangtze block. Compilation of fourteen Ediacaran sections from shallow-, slope- and deep-water facies demonstrates frequent spatiotemporal oscillation of the redox conditions throughout the Yangtze block. Widespread ferruginous states, accompanied with euxinic zones mainly focusing in lower slope facies, are detected during the early Ediacaran period, in contrast with more widespread euxinic settings in the middle Ediacaran period. Statistical data including new high-resolution results from the Lower Yangtze block show generally low enrichment for redox sensitive elements (RSEs, e.g., Mo, V and U) in the Member II shales probably due to their contemporary low seawater concentrations. An increase of RSEs concentrations occurs in the Member IV shales, which probably reflects more oxidized Earth surface environment during middle Ediacaran period. The increasing Earth oxygenation documents the occurrence of widespread euxinic states during deposition of the Member IV shales. Compilation of spatial distribution of TOC contents in black shales demonstrates their apparent relevance to spatial distribution of the euxinic rather than ferruginous states, which is likely due to the favorable role of sulfurization process on burial and preservation of organic matters. The euxinic states may also facilitate the exceptional preservation of the soft-bodied Ediacaran fossils.

Quantifying the effects of exceptional fossil preservation on the global availability of phylogenetic data in deep time.

Fossil deposits with exceptional preservation ("lagerstätten") provide important details not typically preserved in the fossil record, such that they hold an outsized influence on our understanding of biodiversity and evolution. In particular, the potential bias imparted by this so-called "lagerstätten effect" remains a critical, but underexplored aspect of reconstructing evolutionary relationships. Here, we quantify the amount of phylogenetic information available in the global fossil records of 1,327 species of non-avian theropod dinosaurs, Mesozoic birds, and fossil squamates (e.g., lizards, snakes, mosasaurs), and then compare the influence of lagerstätten deposits on phylogenetic information content and taxon selection in phylogenetic analyses to other fossil-bearing deposits. We find that groups that preserve a high amount of phylogenetic information in their global fossil record (e.g., non-avian theropods) are less vulnerable to a "lagerstätten effect" that leads to disproportionate representation of fossil taxa from one geologic unit in an evolutionary tree. Additionally, for each taxonomic group, we find comparable amounts of phylogenetic information in lagerstätten deposits, even though corresponding morphological character datasets vary greatly. Finally, we unexpectedly find that ancient sand dune deposits of the Late Cretaceous Gobi Desert of Mongolia and China exert an anomalously large influence on the phylogenetic information available in the squamate fossil record, suggesting a "lagerstätten effect" can be present in units not traditionally considered lagerstätten. These results offer a phylogenetics-based lens through which to examine the effects of exceptional fossil preservation on biological patterns through time and space, and invites further quantification of evolutionary information in the rock record.

Trace element evidence for diverse origins of superheavy pyrite in Neoproterozoic sedimentary strata

Sedimentary pyrite has long been used as an archive of marine environments in Earth history. To capture reliable paleoenvironmental signals, however, we need to first evaluate pyrite in sedimentary strata as it can be altered and masked by later diagenetic and/or hydrothermal processes. Here, we trained two supervised machine learning algorithms on a large LA-ICP-MS pyrite trace element database to distinguish pyrite of different origins. The analysis validates that two models built on the co-behavior of 12 trace elements (Co, Ni, Cu, Zn, As, Mo, Ag, Sb, Te, Au, Tl, and Pb) can be used to accurately predict pyrite origins. Further statistical analysis suggests four trace element clusters behaving differently among sedimentary (syngenetic and early diagenetic), synsedimentary hydrothermal (syngenetic hydrothermal), and post-sedimentary hydrothermal (epigenetic hydrothermal) pyrite, which is probably driven by chemical and physical properties of source fluids, interactions between elements, competition among coprecipitating minerals, and pyrite growth rate. Armed with this initial success and aided by new LA-ICP-MS trace element data from 9 samples, we then demonstrated the efficacy of this approach in identifying the origins of pyrite from two Neoproterozoic sedimentary successions in South China. The first set of samples contain isotopically superheavy pyrite (i.e., whose bulk-sample δ34S values greater than those of contemporaneous seawater sulfate and whose origins remain controversial) from the Cryogenian Tiesi’ao and Datangpo formations. The second set of samples contain pyritic rims (associated with fossiliferous chert nodules and thought to be critical in exceptional fossil preservation) from the Ediacaran Doushantuo Formation. For the superheavy pyrite, the models consistently show high confidence levels (mostly > 80 % probability) in identifying its genesis type, and three out of four samples were given sedimentary origins. For the pyritic nodule rims, the models suggest that early diagenetic pyrite was subsequently altered by hydrothermal fluids and therefore shows mixed signals. The study highlights the importance of pyrite trace elements in deciphering and distinguishing the origins of pyrite in sedimentary strata.

Taphonomy of non-biomineralized trilobite tissues preserved as calcite casts from the Ordovician Walcott-Rust Quarry, USA

Trilobites with appendages from the Rust Formation of New York State were discovered in the 1870s and represent one of the earliest known cases of exceptional preservation of non-biomineralized tissues. The Rust Formation trilobites feature three-dimensionally preserved walking legs and delicate respiratory lamellae, but the mechanism behind their fossilization remains unknown. Here we show that after burial, carcass decay produced framboidal pyrite, while fibrous calcite precipitated on the visceral side of the body, followed by the widespread formation of sparry calcite crystal replicating non-biomineralized morphological features. Trilobites and co-occurring calcite veins show no chemical or petrographic differences, rejecting the hypothesis that exceptional preservation was caused by a local microenvironment within enrolled trilobites. These results suggest that fine-grained sediment provided support for the appendages and facilitated their fossilization through calcite replacement. Our findings carry broader implications for understanding the exceptional three-dimensional preservation of animal Paleozoic body fossils through calcite casts.

Taphonomy of early life (2.1 Ga) in the francevillian basin (Gabon): Role of organic mineral interactions

The taphonomy of early soft-bodied organisms in Palaeoproterozoic sediments is not yet clearly understood, even though some locations where these fossils are found present all the conditions for exceptional fossil preservation. Indeed, the degree of fossil preservation has received attention, but better knowledge of the environmental conditions and associated taphonomic processes is also essential. In the Gabonese Francevillian Basin, the discovery of macrofossils (2.1 Ga) of multicellular organisms in black shales is an outstanding example of this degree of preservation. Indeed, the biological diversity, as a wide variety of fossil morphologies are observed, is associated with two major taphonomic processes – moulding (lenticular-shaped forms) or early pyritization, while the fossil host rocks were not deeply buried and were affected only by weak to moderate diagenesis. However, usually, the mechanisms of this preservation remain difficult to assess, as the original taphonomic processes are impacted by diagenesis and still misunderstood. In this way, by closely observing fossil mineralization in four morphotypes of macrofossils and associated host rocks from mineralogical and textural points of view, this work aims to provide some keys to a taphonomic comprehension of soft-bodied organism preservation. After the deposition of dead organisms on the clayey sediment, an illitization process, which depends on the availability of dissolved K driven by bacterial activity, started from the first stages of preservation by moulding the lenticular-shaped forms and proceeded in the pores of the other macrofossils after their pyritization. At the fossil level, the intensity of illitization is controlled by the mode of preservation and the evolution of the associated permeability. In the nonpyritized lenticular-shaped specimens, illitization was not achieved, preserving I-S mixed-layer minerals, while in the pyritized forms, the illitization degree was more extensive. In comparison, I-S mixed-layer minerals are almost absent in pyritized abiotic concretions. A second process, which occurred later, consists of general chloritization from fluid circulation. Our detailed results show that each specimen behaved like a microsystem with its own physico-chemical and mineralogical evolution during preservation/diagenesis. This finding allows us to propose a conceptual model of the taphonomic history, describing the fossilization stages for each type of specimen.

Examining the ontogeny of the Pennsylvanian cladid crinoid Erisocrinus typus Meek and Worthen, 1865

AbstractCrinoids were major constituents of late Carboniferous (Pennsylvanian) marine ecosystems, but their rapid disarticulation rates after death result in few well-preserved specimens, limiting the study of their growth. This is amplified for cladids, who had among the highest disarticulation rates of all Paleozoic crinoids due to the relatively loose suturing of the calyx plates. However, Erisocrinus typus Meek and Worthen, 1865 has been found in unusually large numbers, most preserved as cups but some as nearly complete crowns, in the Barnsdall Formation in Oklahoma. The Barnsdall Formation, a Koncentrat Lagerstätte, is composed predominantly of fine- to medium-grained sandstone, overlain by mudstone and shale; severe compaction of the fossils in the mudstone and shale layer in this formation allowed for exceptional preservation of the plates. Herein, we summarize a growth study based on 10 crowns of E. typus, showcasing a well-defined growth series of this species from the Barnsdall Formation, including fossils from juvenile stages of development, which are rarely preserved. We used high-resolution photographs imported into ImageJ and recorded measurements of the cup and arms for all nondistorted or disarticulated plates. Results show that the plates of the cup grew anisometrically with both positive and negative allometry. The primibrachial plates of E. typus grew with positive allometry. The brachial plates started as uniserial (i.e., cuneiform) as juveniles but shifted to be biserial. Erisocrinus typus broadly shares similar growth trajectories with other cladids. These growth patterns provide insight into feeding strategies and can aid in understanding crinoid evolutionary paleoecological trends.

Fungal-induced fossil biomineralization

Exceptional preservation of fossils has often been attributed to the actions of bacteria that aid in the preservation of soft tissues that normally decay rapidly. However, it is well known that fungi play a major role in organic matter decomposition, biogeochemical cycling of elements, and metal-mineral transformations in modern ecosystems. Although the fungal fossil record can be traced back over a billion years, there are only a few recorded examples of fungal roles in fossilization. In this research, we have carried out a detailed geobiological investigation on early Pleistocene hyena coprolites (fossilized dung) in an attempt to ascertain possible fungal involvement in their formation. Using an advanced microscopic and mineralogical approach, we found that numerous hydroxyapatite nanofibers (25-34nm on average), interwoven to form spheroidal structures, constituted the matrix of the coprolites in addition to food remains. These structures were found to be extremely similar in texture and mineral composition to biominerals produced during laboratory culture of a common saprophytic and geoactive fungus, Aspergillus niger, in the presence of a solid source of calcium (Ca) and phosphorus (P). This observation, and our other data obtained, strongly suggests that fungal metabolism can provide a mechanism that can result in fossil biomineralization, and we hypothesize, therefore, that this may have contributed to the formation of well-preserved fossils (Lagerstätten) in the geological record. The characteristic polycrystalline nanofibers may also have served as a potential biosignature for fungal life in early Earth and extraterrestrial environments.

Exceptional fossil preservation and evolution of the ray-finned fish brain.

Brain anatomy provides key evidence for the relationships between ray-finned fishes1, but two majorlimitations obscure our understanding of neuroanatomical evolution in this major vertebrate group. First, the deepest branching living lineages are separated from the group's common ancestor by hundreds of millions of years, with indications that aspects of their brain morphology-like other aspects of their anatomy2,3-are specialized relative to primitive conditions. Second, there are no direct constraints on brain morphology in the earliest ray-finned fishes beyond the coarse picture provided by cranial endocasts: natural or virtual infillings of void spaces within the skull4-8. Here we report brain and cranial nerve soft-tissue preservation in Coccocephalus wildi, an approximately 319-million-year-old ray-finned fish. This example of a well-preserved vertebrate brain provides a window into neural anatomy deep within ray-finned fish phylogeny. Coccocephalus indicates a more complicated pattern of brain evolution than suggested by living species alone, highlighting cladistian apomorphies1 and providing temporal constraints on the origin of traits uniting all extant ray-finned fishes1,9. Our findings, along with a growing set of studies in other animal groups10-12, point to the importance of ancient soft tissue preservation in understanding the deep evolutionary assembly of major anatomical systems outside of the narrow subset of skeletal tissues13-15.

High productivity promoted exceptional fossil preservation of the early Middle Triassic Luoping biota of Yunnan Province, China

The early Middle Triassic (Anisian) Luoping biota is representative of marine ecosystems following their full recovery after the Permian-Triassic mass extinction; however, its exceptional preservation remains poorly understood. In this paper, we report multiple geochemical proxies (TOC, TN, P/Al, Cuxs, Nixs, Baxs, δ13Ccarb, and δ13Corg) from Member II of the Middle Triassic Guanling Formation at the Xiangdongpo section in Luoping County, Yunnan Province, China, to assess the relationship between primary productivity and exceptional preservation of the Luoping biota. Variations in TOC, TN, P/Al, Cuxs, and Nixs values show that exceptional preservation coincides with two intervals of high productivity. Relatively low C/N ratios and distributed Δ13Ccarb-org values indicate that primary productivity was dominated by eukaryotic algae and prokaryotic microbes. Based on geochemical analyses and regional correlations, we conclude that, following sea level rise, nutrients were supplied to the Luoping area by upwelling from the open ocean and facilitated the blooming of marine communities in an open platform setting. In the intra-platform depression, increased nutrients supply also fueled high primary productivity in surface waters and oxygen consumption in the water column, causing bottom water anoxia. The lack of oxygen in the bottom water reduced the rate of biomass degradation and bioturbation, promoting growth of microbial mats, leading to the exceptional preservation of macrofauna. This study highlights the important role of elevated primary productivity in exceptional fossil preservation by triggering anoxia of bottom waters. Our findings confirm the widespread development of anoxic conditions during the middle Anisian (Pelsonian), from the eastern Tethys to the Panthalassa, and also reveal the interactions between organisms and the environment after Permian-Triassic mass extinction.

Mineralogy, geochemistry, and K-Ar dating of feldspars and clays from an exceptional Cretaceous fossil locality (Tlayúa, Puebla, Mexico): Insights into the depositional and diagenetic ages and processes

The exceptional preservation of fossils in a small number of deposits is difficult to explain. The precise conditions for preserving soft-bodied animals or delicate invertebrate and vertebrate structures during their death, burial, and diagenesis must be examined case by case. We studied the Middle Member of the Tlayúa Formation, Puebla, Mexico, a limestone lithostratigraphic unit that contains exceptionally preserved fauna, mainly fishes. Its diagenetic history has been investigated using clay mineralogy and geochemistry, volcanic and authigenic alkali feldspar characterization, and K-Ar dating of clays and feldspars. Our results indicate that the death and initial decay of the fossilized organisms was controlled by the repeated deposition of thin layers, probably volcaniclastic, interrupting the precipitation of limestones in a shallow marine environment associated with algal mats. These layers exhibit high concentrations of heavy metals (V, Cr, Ni, Cu, As, Mo) indicative of anoxia. The Tlayúa Formation experienced a low degree of thermal diagenesis, as determined by the incipient smectite illitization and the presence of liptinite, but chemical diagenesis was observed as clay formation and authigenesis of feldspars. Two generations of feldspars were identified and dated. The first is primary volcanic sanidine, whose K-Ar ages in mineral concentrates and K-Ar in situ on single crystals, gave the deposition age of the upper part of the Middle Member of the Tlayúa Formation at ∼104 to ∼100 Ma (Late Albian-Lower Cenomanian) and a range of inherited volcanic crystals up to 118 Ma. The second feldspar, an authigenic adularia formed between 98 and 82 Ma, indicates the main stage of burial (Cenomanian to Campanian). Later incipient illitization was dated at 70–60 Ma (Maastrichtian-Danian), coincidental with the Mexican Orogen development and later exhumation.

Dataset accompanying "Exceptional fossil preservation and evolution of the ray-finned fish brain"

Dataset accompanying "Exceptional fossil preservation and evolution of the ray-finned fish brain"

Iron from continental weathering dictated soft‐part preservation during the Early Ordovician

AbstractThe Fezouata Shale in Morocco is the most diverse Lower Ordovician unit yielding soft‐tissue preservation. Iron played a crucial role in the preservation of soft parts in this formation through the damage of bacterial membranes under oxic conditions and the pyritization of soft parts under the activity of bacterial sulphate reduction. However, the origin of Fe in this formation remains largely speculative. Herein, trace and rare earth elements were investigated in drilled‐core sediments from the Fezouata Shale. It is shown that a correlation exists between Fe and Al suggesting that most Fe has a detrital source. Elemental concentrations in the Fezouata Shale are most comparable to rivers and are the least similar to loess and sediments deposited near active island arcs. In this sense, continental weathering and its related Fe in river fluxes dictated occurrences of exceptional fossil preservation in the Fezouata Shale.

Ultrastructure and in-situ chemical characterization of intracellular granules of embryo-like fossils from the early Ediacaran Weng’an biota

Embryo-like fossils from the early Ediacaran Weng’an biota provide a window of exceptional fossil preservation onto the period of life history in which molecular clocks estimate the fundamental animal lineages to have diverged. However, their diversity and biological affinities have proven controversial, because they are morphologically simple and, consequently, their interpretation lacks phylogenetic constraint. The subcellular structures preserved in these embryo-like fossils might help to understand their cytology, biology, and diversity, but the potential of these structures has not been fully realized, because detailed microscale physical and chemical investigations are lacking. Here, to remedy this deficiency, we performed a comprehensive study to characterize their micro- and ultra-structures as well as in-situ chemical components. Our results reveal three types of subcellular structure that differ in size, shape, and mineral components: (1) relatively small and spheroidal granules in embryo-like fossils with equal cell division pattern; (2) relatively large, spheroidal, or polygonal granules in embryo-like fossils with unequal and asynchronous cell division pattern; and (3) irregular multi-layered rim-bounded granules in embryo-like fossils with unequal and asynchronous cell division pattern. We propose that the three types may be rationalized to a single taphonomic pathway of preferential mineralization of the cell cytoplasm, preserving an external mould of subcellular granules. We followed the previous interpretation that the spheroidal and polygonal granules should be fossilized lipid droplets or yolk platelets. The distinction between these subcellular structures are largely the result of postmortem degradation processes such as autolysis. The widely preserved lipid droplets or yolk platelets within these Ediacaran embryo-like fossils are compatible with the interpretion of large yolky embryos with maternal nourishment and direct development.

Taphonomy of marine vertebrates of the Pisco Formation (Miocene, Peru): Insights into the origin of an outstanding Fossil-Lagerstätte.

The Miocene Pisco Formation, broadly exposed in the Ica Desert of southern Peru, is among the most outstanding Cenozoic marine Fossil-Lagerstätten worldwide. It is renowned for its exceptional preservation and abundance of vertebrate fossils, including a rich assemblage of whales and dolphins (Cetacea). Here, we integrate taphonomic data on 890 marine vertebrate fossils, gathered through 16 different localities. Our observations range from the taxonomic distribution, articulation, completeness, disposition and orientation of skeletons, to the presence of bite marks, associations with shark teeth and macro-invertebrates, bone and soft tissue preservation, and the formation of attendant carbonate concretions and sedimentary structures. We propose that the exceptional preservation characterising many Pisco vertebrates, as well as their exceptionally high abundance, cannot be ascribed to a single cause like high sedimentation rates (as proposed in the past), but rather to the interplay of several favourable factors including: (i) low levels of dissolved oxygen at the seafloor (with the intervention of seasonal anoxic events); (ii) the early onset of mineralisation processes like apatite dissolution/recrystallisation and carbonate mineral precipitation; (iii) rapid burial of carcasses in a soupy substrate and/or a novel mechanism involving scour-induced self-burial; and (iv) original biological richness. Collectively, our observations provide a comprehensive overview of the taphonomic processes that shaped one of South America’s most important fossil deposits, and suggest a model for the formation of other marine vertebrate Fossil-Lagerstätten.

Evolutionary drivers, morphological evolution and diversity dynamics of a surviving mammal clade: cainotherioids at the Eocene-Oligocene transition.

The Eocene-Oligocene transition (EOT) represents a period of global environmental changes particularly marked in Europe and coincides with a dramatic biotic turnover. Here, using an exceptional fossil preservation, we document and analyse the diversity dynamics of a mammal clade, Cainotherioidea (Artiodactyla), that survived the EOT and radiated rapidly immediately after. We infer their diversification history from Quercy Konzentrat-Lagerstätte (south-west France) at the species level using Bayesian birth-death models. We show that cainotherioid diversity fluctuated through time, with extinction events at the EOT and in the late Oligocene, and a major speciation burst in the early Oligocene. The latter is in line with our finding that cainotherioids had a high morphological adaptability following environmental changes throughout the EOT, which probably played a key role in the survival and evolutionary success of this clade in the aftermath. Speciation is positively associated with temperature and continental fragmentation in a time-continuous way, while extinction seems to synchronize with environmental change in a punctuated way. Within-clade interactions negatively affected the cainotherioid diversification, while inter-clade competition might explain their final decline during the late Oligocene. Our results provide a detailed dynamic picture of the evolutionary history of a mammal clade in a context of global change.

Distraction sinking and fossilized coleoid predatory behaviour from the German Early Jurassic

Exceptional fossil preservation is required to conserve soft-bodied fossils and even more so to conserve their behaviour. Here, we describe a fossil of a co-occurrence of representatives of two different octobrachian coleoid species. The fossils are from the Toarcian Posidonienschiefer of Ohmden near Holzmaden, Germany. The two animals died in the act of predation, i.e. one had caught the other and had begun to nibble on it, when they possibly sank into hypoxic waters and suffocated (distraction sinking). This supports the idea that primitive vampyromorphs pursued diverse feeding strategies and were not yet adapted to being opportunistic feeders in oxygen minimum zones like their modern relative Vampyroteuthis.

The link between exceptional fossil preservation and palaeo‐redox conditions in the Middle Triassic Luoping Biota from South China

The Luoping Biota represents of the recovery of marine ecosystems after the Permian–Triassic Mass Extinction. To better understand the palaeoenvironmental background of the Luoping Biota, sedimentary analysis and multi‐geochemical proxies (Mo, U, V, V/Cr, MoEF, UEF, and VEF) were assessed in Member II of the Middle Triassic Guanling Formation of the Xiangdongpo section in Luoping County, Yunnan Province. According to the facies characteristics and redox‐sensitive elemental proxies, in combination with macrofossil and bioturbation evidence, five intervals (I, II, III, IV, and V) were recognized. The exceptional preservation of the fossils is related to two anoxic intervals (II and IV) sandwiched by three suboxic intervals (I, III, and V). Based on this study and regional correlation, it is concluded that anoxia in the water‐column increased the preservation potential of macrofauna fossils and inhibited the survival of trace‐making organisms. In addition, microbial mats could produce local anoxic and restricted micro‐environments, preventing the disarticulation of the carcasses by water turbulence. Therefore, we demonstrate that anoxic conditions played a key role in the exceptional fossil preservation of the Luoping Biota.

Taxon- and senescence-specific fluorescence of colored leaves from the Pliocene Willershausen Lagerst\xe4tte, Germany

UV-light-induced fluorescence is widely used in the study of coal macerals and palynological samples, but to date has not been described in great detail for plant macrofossils. Here, we report the characteristics of bright UV-light-induced fluorescence of various fossil angiosperm leaf taxa from the Upper Pliocene of Willershausen, Lower Saxony, Germany. The fluorescence is exceptional, since different fluorescence colors ranging from green to yellow to red can be observed and fluorescence properties are found to be related to genera. Using confocal laser scanning microscopy, fluorescence was studied in detail and emission spectra were obtained that allowed to differentiate broad groups of fluorophores. Fluorescence emissions attributed to chlorophyll degradation products demonstrate that fluorescence can be used as an indicator for exceptional chemical preservation of leaf fossils. Comparison with present-day senescing plants suggests that the fluorescence differences in the fossil leaves are mainly caused by taxon-specific degeneration of organic compounds during senescence. The occurrence of various leaf taxa with different fluorescence properties, preserved under identical conditions of fossilization, indicate that diagenesis was not crucial for the differences in leaf fluorescence.

Oxidative conditions can lead to exceptional preservation through phosphatization

Abstract Exceptional preservation through phosphatization is primarily controlled by a reduction in pH, favoring the precipitation of apatite over that of calcite. Laboratory experiments have suggested that phosphatization results from anoxic decay. Here we report results of the fine-scale mineralogical characterization of Cretaceous phosphatized fossils of teleost fishes and crustaceans from the Jebel oum Tkout Lagerstätte (Morocco). Data collected using complementary laboratory and synchrotron-based X-ray techniques reveal that oxidative conditions were established at a certain step of decay. Supporting these conclusions are the presence, covering and embedded in the phosphatized tissues, of Fe(III)-rich mineral phases, the precipitation of which was likely biologically induced during decay. The present study highlights that the establishment of oxidative conditions during decay can be compatible with exceptional preservation of fossils through phosphatization.

Developmental biology of Helicoforamina reveals holozoan affinity, cryptic diversity, and adaptation to heterogeneous environments in the early Ediacaran Weng'an biota (Doushantuo Formation, South China).

The exceptional fossil preservation of the early Ediacaran Weng'an biota provides a unique window on the interval of Earth history in which animal lineages emerged. It preserves a diversity of similarly ornamented encysted developmental stages previously interpreted as different developmental stages of one taxon. Although Helicoforamina wenganica is distinguished from other forms by a helical groove or canal, it has been interpreted as a developmental stage of cooccurring metazoan, nonmetazoan holozoan, or green algal taxa. Using x-ray microtomography, we show that Helicoforamina developed through one-, four-, and eight-cell stages, to hundreds and thousands of cells. Putative hatchlings are artifacts of incompletely preserved cyst walls. Our results preclude inclusion of Helicoforamina into life cycles assembled from other components of the Weng'an biota but support a holozoan affinity. The similarly ornamented encysted forms shared among the diverse Weng'an biota represent parallel adaptations to the temporally and spatially heterogeneous Ediacaran shallow marine environments.