Sarah E. Gabbott

Preservation of Early Cambrian animals of the Chengjiang biota

The Chengjiang biota of Yunnan, China, documents the earliest extensive radiation of the Metazoa recorded in the fossil record. Gauging preservational bias is crucial in providing an assessment of the completeness of this biota and thereby elucidating whether it represents a comprehensive depiction of Early Cambrian life. We here present a model to explain the nature of the exceptional preservation of the Chengjiang biota and details of the decay process. This study indicates that Chengjiang fossils were preserved through two taphonomic pathways that may have captured tissues of distinct compositions, and this finding should provide a foundation for the interpretation of Chengjiang fossils. Many Chengjiang fossils are preserved by pyrite (later pseudomorphed by iron oxides); the clay-rich host sediment was deficient in organic carbon but replete in available Fe, and this composition ensured that a decaying carcass acted as a local substrate for Fe- and S-reducing bacteria. Pyrite morphology probably reflects contrasts in the decay rate, and hence the H 2 S production rate, of different tissues in a carcass. Reactive, rapidly decaying tissues would have quickly supplied H 2 S, producing many pyrite nuclei, resulting in framboidal habits. More recalcitrant tissues would have produced H 2 S more slowly, so that crystal growth operated on fewer nuclei, resulting in larger euhedral pyrite crystals. Reflective films, especially common on Chengjiang arthropods, represent the remains of degraded carbon.

Non-random decay of chordate characters causes bias in fossil interpretation.

Exceptional preservation of soft-bodied Cambrian chordates provides our only direct information on the origin of vertebrates. Fossil chordates from this interval offer crucial insights into how the distinctive body plan of vertebrates evolved, but reading this pre-biomineralization fossil record is fraught with difficulties, leading to controversial and contradictory interpretations. The cause of these difficulties is taphonomic: we lack data on when and how important characters change as they decompose, resulting in a lack of constraint on anatomical interpretation and a failure to distinguish phylogenetic absence of characters from loss through decay. Here we show, from experimental decay of amphioxus and ammocoetes, that loss of chordate characters during decay is non-random: the more phylogenetically informative are the most labile, whereas plesiomorphic characters are decay resistant. The taphonomic loss of synapomorphies and relatively higher preservation potential of chordate plesiomorphies will thus result in bias towards wrongly placing fossils on the chordate stem. Application of these data to Cathaymyrus (Cambrian period of China) and Metaspriggina (Cambrian period of Canada) highlights the difficulties: these fossils cannot be placed reliably in the chordate or vertebrate stem because they could represent the decayed remains of any non-biomineralized, total-group chordate. Preliminary data suggest that this decay filter also affects other groups of organisms and that ‘stem-ward slippage’ may be a widespread but currently unrecognized bias in our understanding of the early evolution of a number of phyla.

Mechanism for Burgess Shale-type preservation

Exceptionally preserved fossil biotas of the Burgess Shale and a handful of other similar Cambrian deposits provide rare but critical insights into the early diversification of animals. The extraordinary preservation of labile tissues in these geographically widespread but temporally restricted soft-bodied fossil assemblages has remained enigmatic since Walcott’s initial discovery in 1909. Here, we demonstrate the mechanism of Burgess Shale-type preservation using sedimentologic and geochemical data from the Chengjiang, Burgess Shale, and five other principal Burgess Shale-type deposits. Sulfur isotope evidence from sedimentary pyrites reveals that the exquisite fossilization of organic remains as carbonaceous compressions resulted from early inhibition of microbial activity in the sediments by means of oxidant deprivation. Low sulfate concentrations in the global ocean and low-oxygen bottom water conditions at the sites of deposition resulted in reduced oxidant availability. Subsequently, rapid entombment of fossils in fine-grained sediments and early sealing of sediments by pervasive carbonate cements at bed tops restricted oxidant flux into the sediments. A permeability barrier, provided by bed-capping cements that were emplaced at the seafloor, is a feature that is shared among Burgess Shale-type deposits, and resulted from the unusually high alkalinity of Cambrian oceans. Thus, Burgess Shale-type preservation of soft-bodied fossil assemblages worldwide was promoted by unique aspects of early Paleozoic seawater chemistry that strongly impacted sediment diagenesis, providing a fundamentally unique record of the immediate aftermath of the “Cambrian explosion.”

Decay of vertebrate characters in hagfish and lamprey (Cyclostomata) and the implications for the vertebrate fossil record

The timing and sequence of events underlying the origin and early evolution of vertebrates remains poorly understood. The palaeontological evidence should shed light on these issues, but difficulties in interpretation of the non-biomineralized fossil record make this problematic. Here we present an experimental analysis of decay of vertebrate characters based on the extant jawless vertebrates (Lampetra and Myxine). This provides a framework for the interpretation of the anatomy of soft-bodied fossil vertebrates and putative cyclostomes, and a context for reading the fossil record of non-biomineralized vertebrate characters. Decay results in transformation and non-random loss of characters. In both lamprey and hagfish, different types of cartilage decay at different rates, resulting in taphonomic bias towards loss of ‘soft’ cartilages containing vertebrate-specific Col2α1 extracellular matrix proteins; phylogenetically informative soft-tissue characters decay before more plesiomorphic characters. As such, synapomorphic decay bias, previously recognized in early chordates, is more pervasive, and needs to be taken into account when interpreting the anatomy of any non-biomineralized fossil vertebrate, such as Haikouichthys, Mayomyzon and Hardistiella.

Ubiquitous Burgess Shale–style “clay templates” in low-grade metamorphic mudrocks

Despite the Burgess Shale’s (British Columbia, Canada) paleobiological importance, there is little consensus regarding its taphonomy. Its organic fossils are preserved as compressions associated with phyllosilicate fi lms (“clay templates”). Debate focuses on whether these templates were fundamental in exceptional preservation or if they formed in metamorphism, meaning that it is important to establish the timing of their formation relative to decay. An early diagenetic origin has been proposed based on anatomy-specifi c variations in their composition, purportedly refl ecting contrasts in decay. However, we demonstrate that these films bear a remarkable similarity to those that occur on organic fossils in graptolitic mudrocks and form as a normal product of low-grade metamorphism. Such phyllosilicates may also occur within voids created by volume loss in maturation, a process that may have aided their formation. In bedding-plane assemblages from graptolitic mudrocks, different taxa are associated with distinct phyllosilicates. This likely refl ects stepwise maturation of their constituent kerogens in an evolving hydrothermal fl uid, with different hyllosilicates forming as each taxon progressively underwent maturation. These observations provide an analogue for the distribution and composition of phyllosilicates on Burgess Shale fossils, which we interpret as refl ecting variations in the maturation of their constituent tissues. Thus, their clay templates seem unremarkable, forming too late to account for exceptional preservation.

Sedimentation of the Phyllopod Bed within the Cambrian Burgess Shale Formation of British Columbia

We provide the most detailed sedimentological log to date through the Phyllopod Bed of the mid-Cambrian Burgess Shale Formation of British Columbia, based on millimetre-scale logging of a suite of thin sections. The sedimentary facies is dominated by alternations of homogeneous mudstone and a coarser-grained, laminated, variably sandy and shelly mudstone that is locally micronodular. Most boundaries between these two lithologies are gradational, and discrete fining-upwards turbidite units were rarely recognized. Such a pattern is interpreted to indicate rapid sedimentation of up to decimetre-thick units at this location from pulsatory, quasi-continuous density currents consistent with earlier proposals of exceptional preservation through rapid burial; the density currents responsible were probably largely akin to mud-rich slurries, helping explain the transport and entombment of the fossils. The homogeneous mudstone units are characterized by numerous distinctive lenses of pyrite framboids or subeuhedral crystals, previously interpreted as small ripples. Their 3D shape, however, suggests an origin as subspherical early diagenetic aggregates; their present morphology is consistent with the high levels of compaction inferred from the preservation of fossils.

Chemical, experimental, and morphological evidence for diagenetically altered melanin in exceptionally preserved fossils

Significance Melanin is a widespread pigment that provides black to reddish brown hues to organisms. Recent evidence has shown that melanin is retained in exceptionally preserved fossils, including feathered dinosaurs, allowing the reconstruction of ancient color patterns. However, little is known about the chemical preservation of melanin or its distribution in the fossil record. Here, we show that melanin is preserved in a number of soft-bodied fossils, but its burial under high pressure and temperature for millions of years alters its original chemistry. The widespread occurrence of melanin substantiates the applicability of reconstructing aspects of original color patterns and allows us to dismiss the alternative suggestion that these structures are microbial in origin. In living organisms, color patterns, behavior, and ecology are closely linked. Thus, detection of fossil pigments may permit inferences about important aspects of ancient animal ecology and evolution. Melanin-bearing melanosomes were suggested to preserve as organic residues in exceptionally preserved fossils, retaining distinct morphology that is associated with aspects of original color patterns. Nevertheless, these oblong and spherical structures have also been identified as fossilized bacteria. To date, chemical studies have not directly considered the effects of diagenesis on melanin preservation, and how this may influence its identification. Here we use time-of-flight secondary ion mass spectrometry to identify and chemically characterize melanin in a diverse sample of previously unstudied extant and fossil taxa, including fossils with notably different diagenetic histories and geologic ages. We document signatures consistent with melanin preservation in fossils ranging from feathers, to mammals, to amphibians. Using principal component analyses, we characterize putative mixtures of eumelanin and phaeomelanin in both fossil and extant samples. Surprisingly, both extant and fossil amphibians generally exhibit melanosomes with a mixed eumelanin/phaeomelanin composition rather than pure eumelanin, as assumed previously. We argue that experimental maturation of modern melanin samples replicates diagenetic chemical alteration of melanin observed in fossils. This refutes the hypothesis that such fossil microbodies could be bacteria, and demonstrates that melanin is widely responsible for the organic soft tissue outlines in vertebrates found at exceptional fossil localities, thus allowing for the reconstruction of certain aspects of original pigment patterns.

Orthoconic cephalopods and associated fauna from the late Ordovician Soom Shale Lagerstatte, South Africa

. Orthoconic cephalopods from the Soom Shale Member (Ashgill) are exceptionally preserved and are colonized by lingulate brachiopods and cornulitids. Other fossils commonly associated with orthocones include myodocopid ostracodes and chitinozoans. Size distribution analysis of the brachiopods on one orthocone indicates that it was colonized in vivo. Four orthocone radulae are preserved extending the record of these structures 50 My back to the late Ordovician. Orthocone radula configuration is more similar to that of ammonoids and coleoids than to that of nautiloids.

Chitinozoans and the age of the Soom Shale, an Ordovician black shale Lagerstätte, South Africa

Isolated chitinozoans from the Soom Shale Member of the Cedarberg Formation, SW South Africa are described and provide a date of the latest Hirnantian–earliest Rhuddanian. The recovered chitinozoans are typical of the latest Ordovician Spinachitina oulebsiri Biozone, although an earliest Silurian age is possible. They indicate a very short time span (less than 1 Ma) across the Ordovician–Silurian boundary. This is currently the highest biostratigraphical resolution attainable for the Soom Shale Lagerstätte. Correlation of the Soom Shale chitinozoans with identical assemblages in post-glacial, transgressive deposits of Northern Africa is possible; both faunas occur in shales that overlie glacial diamictites of the Hirnantian glaciation. A new species, Spinachitina verniersi n. sp. is described.

The earliest myodocopes: ostracodes from the late Ordovician Soom Shale Lagerstätte of South Africa

The late Ordovician Soom Shale Lagerstatte of South Africa has yielded Myodoprimigenia fistuca n. gen. and n. sp., the earliest and only known Ordovician occurrence of myodocopes, one of the major groups of ostracodes. M. fistuca is a likely sister group of the Upper Silurian ‘cypridinid’ myodocopes and allied forms. It had a thin, lightly mineralized and flexible shell with microstructures resulting from in vivo calcification processes. It probably fed on cephalopod carrion, thus extending evidence for a carnivorous scavenging lifestyle in ostracodes back by 200 Ma. The species was probably nektobenthic and thus consistent with the notion that the origin of the late Silurian pelagic myodocopes - and therefore of pelagic ostracodes - is to be charted in a benthic to pelagic ecological shift in the group.