Microbial biofilms and the preservation of the Ediacara biota

Laflamme, M., Schiffbauer, J.D., Narbonne, G.M., & Briggs, D.E.G. 2011: Microbial biofilms and the preservation of the Ediacara biota. Lethaia, Vol. 44, pp. 203–213. The terminal Neoproterozoic Ediacaran Period is typified by the Ediacara biota (ca. 579–542 Ma), which includes the first morphologically complex macroscopic organisms. Both the taphonomic setting that promoted the preservation of the soft‐bodied Ediacara biota in coarse‐grained sediments, and the influence of associated microbial coatings on this process, have generated debate. Specimens of Ediacaran discs (Aspidella) from the Fermeuse Formation of Newfoundland, Canada, were analysed using environmental scanning electron microscopy (ESEM) and focused ion beam electron microscopy (FIB‐EM) to determine the relationship between the fossil specimens and the surrounding sediment. The presence of chemically distinct (Al–Mg–Fe–K‐ and to a lesser extent S‐rich), finer‐grained sediment (with organized iron sulphides) surrounding the upper and lower margins of the Ediacaran fossils is consistent with elemental analyses of well preserved bacterial biofilms from other localities. ESEM analyses reveal a contrast in the composition of the sediment bound within the discs, which contains a higher concentration of Al, Ca and K, and the purer Si‐rich sediment that forms the surrounding matrix. This suggests that the coarse grained sediment was incorporated into the organism during life. Ediacaran discs were likely surrounded by a bacterial biofilm or thin microbial mat composed primarily of extracellular polymeric substances (or exopolysaccharide) during life, which added structural stability to these frond holdfasts, and facilitated their fossilization. Microbially mediated preservation in Fermeuse‐style Ediacaran taphonomy provides an explanation for the dominance of Aspidella holdfasts in these settings, and suggests that preservation of Ediacaran fossils in the round may be much more prevalent than previously recognized. We suggest that the overwhelming dominance of circular to bulbous forms such as Aspidella in Ediacaran biotas around the world is a direct result of the interplay between microbial ecology and microbially mediated taphonomy. □Aspidella, Ediacaran preservation, environmental scanning electron microscopy, focused ion beam electron microscopy, palaeoecology, taphonomic bias.     

Global marine redox changes drove the rise and fall of the Ediacara biota

The role of O₂ in the evolution of early animals, as represented by some members of the Ediacara biota, has been heavily debated because current geochemical evidence paints a conflicting picture regarding global marine O₂ levels during key intervals of the rise and fall of the Ediacara biota. Fossil evidence indicates that the diversification the Ediacara biota occurred during or shortly after the Ediacaran Shuram negative C‐isotope Excursion (SE), which is often interpreted to reflect ocean oxygenation. However, there is conflicting evidence regarding ocean oxygen levels during the SE and the middle Ediacaran Period. To help resolve this debate, we examined U isotope variations (δ²³⁸U) in three carbonate sections from South China, Siberia, and USA that record the SE. The δ²³⁸U data from all three sections are in excellent agreement and reveal the largest positive shift in δ²³⁸U ever reported in the geologic record (from ~ ?0.74‰ to ~ ?0.26‰). Quantitative modeling of these data suggests that the global ocean switched from a largely anoxic state (26%–100% of the seafloor overlain by anoxic waters) to near‐modern levels of ocean oxygenation during the SE. This episode of ocean oxygenation is broadly coincident with the rise of the Ediacara biota. Following this initial radiation, the Ediacara biota persisted until the terminal Ediacaran period, when recently published U isotope data indicate a return to more widespread ocean anoxia. Taken together, it appears that global marine redox changes drove the rise and fall of the Ediacara biota.     

The youngest Ediacaran fossils from southern Africa

Discovery of fossils of the Ediacara biota near the top of the Spitzkopf Member at farm Swanpunt extends the known range of these remains in Namibia more than 600 m to near the sub-Cambrian unconformity. The fossiliferous beds occur approximately 100 m above a volcanic ash dated at 543 +/- 1 Ma, and thus may be the youngest Proterozoic Ediacara-type fossils reported anywhere in the world. Fossils are preserved within and on the tops of dm-thick beds of storm-deposited sandstone at two stratigraphic levels; the environment is interpreted as open marine, generally calm but with episodic disruptions by storm waves, and probably within the euphotic zone. The presence of Pteridinium carolinaense (St. Jean), which is also known from the classic sections in Ediacara and the White Sea among others, reinforces evidence from geochronology and chemostratigraphy that the Swartpunt section is terminal Neoproterozoic in age. The new genus and species Swartpuntia germsi is a large, multifoliate frond that exhibits at least three quilted petaloids. Macroscopically, Swartpuntia resembles Pteridinium and Ediacara-type fronds such as Charniodiscus traditionally interpreted as Cnidaria, whereas microscopically it exhibits segmentation that is remarkably similar to that of the putative worm Dickinsonia. Combination of diagnostic characters of these supposedly disparate groups in a single species suggests that many species of quilted Ediacaran organisms were more similar to each other than they were to any modern groups, and provides support for the concept of the "Vendobionta" as a late Neoproterozoic group of mainly multifoliate organisms with a distinctive quilted segmentation.     

Dark extinction: the problem of unknown historical extinctions

The extinction of species before they are discovered and named (dark extinction, DE) is widely inferred as a significant part of species loss in the ‘pre-taxonomic’ period (approx. 1500–1800 CE) and, to some extent, in the ‘taxonomic period’ (approx. 1800–present) as well. The discovery of oceanic islands and other pristine habitats by European navigators and the consequent introduction of destructive mammals, such as rats and goats, started a process of anthropogenic extinction. Much ecosystem change happened before systematic scientific recording, so has led to DE. Statistical methods are available to robustly estimate DE in the ‘taxonomic period’. For the ‘pre-taxonomic period’, simple extrapolation can be used. The application of these techniques to world birds, for example, suggests that approximately 56 DEs occurred in the ‘taxonomic period’ (1800–present) and approximately 180 in the ‘pre-taxonomic period’ (1500–1800). Targeting collection activities in extinction hotspots, to make sure organisms are represented in collections before their extinction, is one way of reducing the number of extinct species without a physical record (providing that collection efforts do not themselves contribute to species extinction).     

Effaced preservation in the Ediacara biota and its implications for the early macrofossil record

Abstract: Ediacaran structures known as ‘pizza discs’ or Ivesheadia have long been considered enigmatic. They are amongst the oldest known members of the Ediacara biota, apparently restricted to the Avalonian successions of Newfoundland and the UK, c. 579–560 Ma. Here, we suggest that these impressions are taphomorphs, resulting from the post‐mortem decay of the frondose Ediacaran biota. Ediacaran fossils range from well‐preserved, high‐fidelity variants to almost completely effaced specimens. The effaced specimens are inferred to have undergone modification of their original morphology by post‐mortem microbial decay on the sea floor, combined with sediment trapping and binding. In this style of preservation, morphological details within the organism became variously subdued as a function of the extent of organic decay prior to casting by overlying sediments. Decay and effacement were progressive in nature, producing a continuum of grades of preservation on Ediacaran bedding planes. Fossils preserved by such ‘effaced preservation’ are those that have suffered these processes to the extent that only their gross form can be determined. We suggest that the lack of detailed morphology in effaced specimens renders such fossils unsuitable for use as type material, as it is possible that several taxa may, upon degradation and burial, generate similar morphological taphomorphs. We here reinterpret the genus Ivesheadia as a taphomorph resulting from extensive post‐mortem decay of frondose organisms. Blackbrookia, Pseudovendia and Shepshedia from beds of comparable age in England are likewise regarded as taphomorphs broadly related to Charnia or Charniodiscus spp. To reflect the suggestion that such impressions are likely to be taphomorphs, and not taxonomically discrete, we propose the term ivesheadiomorphs to incorporate all such effaced taphonomic expressions of Ediacaran macrofossil taxa in Avalonian assemblages. Our recognition of effaced preservation has significant implications for Ediacaran taxonomy, and consequently for measures of Ediacaran diversity and disparity. It is implied that Avalonian assemblages preserve both organisms that were alive and organisms that were already dead at the time of burial. As such, the fossil assemblages cannot be taken to represent census populations of living organisms, as in prior interpretations.     

Metacommunity analyses show an increase in ecological specialisation throughout the Ediacaran period

The first animals appear during the late Ediacaran (572 to 541 Ma); an initial diversity increase was followed reduction in diversity, often interpreted as catastrophic mass extinction. We investigate Ediacaran ecosystem structure changes over this time period using the “Elements of Metacommunity Structure” framework to assess whether this diversity reduction in the Nama was likely caused by an external mass extinction, or internal metacommunity restructuring. The oldest metacommunity was characterised by taxa with wide environmental tolerances, and limited specialisation or intertaxa associations. Structuring increased in the second oldest metacommunity, with groups of taxa sharing synchronous responses to environmental gradients, aggregating into distinct communities. This pattern strengthened in the youngest metacommunity, with communities showing strong environmental segregation and depth structure. Thus, metacommunity structure increased in complexity, with increased specialisation and resulting in competitive exclusion, not a catastrophic environmental disaster, leading to diversity loss in the terminal Ediacaran. These results reveal that the complex eco-evolutionary dynamics associated with Cambrian diversification were established in the Ediacaran.

This study shows that the eco-evolutionary dynamics of metazoan diversification known from the Cambrian Period started earlier in the Ediacaran Period with the Avalon assemblage and increased in complexity towards the Phanerozoic as new anatomical innovations appeared, culminating in the “Cambrian Explosion."     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

Testing the protozoan hypothesis for Ediacaran fossils: a developmental analysis of Palaeopascichnus

Abstract: The hypothesis that the Ediacara biota were giant protozoans is tested by considering the external morphology, internal organization, suggested fossil representatives and molecular phylogeny of the xenophyophores. From this analysis, we find no case to support a direct relationship. Rather, the xenophyophores are here regarded as a group of recently evolved Foraminifera and are hence unlikely to have a record from the Ediacaran Period. Further from the growth dynamics of Foraminifera, they are also unlikely to be related to the Palaeopascichnus organism. We also find significant distinctions in the growth dynamics of Palaeopascichnus and organisms usually referred to the Ediacara biota, such as Charnia and Dickinsonia. Developmental analysis of the Palaeopascichnus– central to the xenophyophore hypothesis – reveals unusual, protozoan features, including evidence for chaotic repair structures, for mergence of coeval forms, as well as complex bifurcations. These observations suggest that Palaeopascichnus is a body fossil of an unidentified protozoan but is unrepresentative of Ediacaran body construction, in general.     

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.     

Facies selectivity of benthic invertebrates in a Permian/Triassic boundary microbialite succession: Implications for the “microbialite refuge” hypothesis

Thrombolite and stromatolite habitats are becoming increasingly recognized as important refuges for invertebrates during Phanerozoic Oceanic Anoxic Events (OAEs); it is posited that oxygenic photosynthesis by cyanobacteria in these microbialites provided a refuge from anoxic conditions (i.e., the “microbialite refuge” hypothesis). Here, we test this hypothesis by investigating the distribution of ~34, 500 benthic invertebrate fossils found in ~100 samples from a microbialite succession that developed following the latest Permian mass extinction event on the Great Bank of Guizhou (South China), representing microbial (stromatolites and thrombolites) and non‐microbial facies. The stromatolites were the least taxonomically diverse facies, and the thrombolites also recorded significantly lower diversities when compared to the non‐microbial facies. Based on the distribution and ornamentation of the bioclasts within the thrombolites and stromatolites, the bioclasts are inferred to have been transported and concentrated in the non‐microbial fabrics, that is, cavities around the microbial framework. Therefore, many of the identified metazoans from the post‐extinction microbialites are not observed to have been living within a microbial mat. Furthermore, the lifestyle of many of the taxa identified from the microbialites was not suited for, or even amenable to, life within a benthic microbial mat. The high diversity of oxygen‐dependent metazoans in the non‐microbial facies on the Great Bank of Guizhou, and inferences from geochemical records, suggests that the microbialites and benthic communities developed in oxygenated environments, which disproves that the microbes were the source of the oxygenation. Instead, we posit that microbialite successions represent a taphonomic window for exceptional preservation of the biota, similar to a Konzentrat‐Lagerstätte, which has allowed for diverse fossil assemblages to be preserved during intervals of poor preservation.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

Small but Powerful: Top Predator Local Extinction Affects Ecosystem Structure and Function in an Intermittent Stream

Top predator loss is a major global problem, with a current trend in biodiversity loss towards high trophic levels that modifies most ecosystems worldwide. Most research in this area is focused on large-bodied predators, despite the high extinction risk of small-bodied freshwater fish that often act as apex consumers. Consequently, it remains unknown if intermittent streams are affected by the consequences of top-predators’ extirpations. The aim of our research was to determine how this global problem affects intermittent streams and, in particular, if the loss of a small-bodied top predator (1) leads to a ‘mesopredator release’, affects primary consumers and changes whole community structures, and (2) triggers a cascade effect modifying the ecosystem function. To address these questions, we studied the top-down effects of a small endangered fish species, Barbus meridionalis (the Mediterranean barbel), conducting an enclosure/exclosure mesocosm experiment in an intermittent stream where B. meridionalis became locally extinct following a wildfire. We found that top predator absence led to ‘mesopredator release’, and also to ‘prey release’ despite intraguild predation, which contrasts with traditional food web theory. In addition, B. meridionalis extirpation changed whole macroinvertebrate community composition and increased total macroinvertebrate density. Regarding ecosystem function, periphyton primary production decreased in apex consumer absence. In this study, the apex consumer was functionally irreplaceable; its local extinction led to the loss of an important functional role that resulted in major changes to the ecosystem’s structure and function. This study evidences that intermittent streams can be affected by the consequences of apex consumers’ extinctions, and that the loss of small-bodied top predators can lead to large ecosystem changes. We recommend the reintroduction of small-bodied apex consumers to systems where they have been extirpated, to restore ecosystem structure and function.     

CHARNIA AT 50: DEVELOPMENTAL MODELS FOR EDIACARAN FRONDS

Abstract:  Correct interpretation of the Ediacara biota is critical to our understanding of the dramatic events at the base of the Cambrian. We review here the history of thought and examine new laser images of the holotype of Charnia masoni Ford, 1958 , of the Ediacara biota, in terms of growth and development. Growth and development are argued to provide critical tools for understanding this and other enigmatic fossil groups. We show that Charnia cannot be related to the modern cnidarian group, the sea pens, with which it has for so long been compared, because they have opposite growth polarities. This is shown by our work on material collected by HMS challenger. Recent evolutionary studies also show that sea pens are a highly derived group of actively burrowing cnidarians that are likely to have evolved later than the Palaeozoic. The traditional paradigm of translating Phanerozoic animal phyla back into the Ediacaran is therefore questioned.     

Ediacaran fronds

Leaf-shaped Ediacaran fronds such as Charniodiscus, Charnia, Rangea and Swartpuntia are among the most widespread and readily recognized elements of the soft-bodied Ediacara Biota (late Neoproterozoic). Recent papers have proposed a variety of possible lifestyles for Ediacaran fronds, but functional morphological analysis supports the traditional view that Ediacaran fronds gathered nutrients from the water column through filter feeding or direct nutrient absorption, resulting in the onset of Phanerozoic-style ecological competition complete with denizens occupying distinct tiers in the water column. Within the standard Ediacaran frond morphoshape, four distinct branching architectures have been isolated: Arborea-type branching, characterized by pea-pod like primary branches which house several secondary branches within the protective sheath; Charnia-type branching, resulting in a zigzagging central axis due to the overlapping of alternating sigmoidal primary branches which overly an inferred internal stalk; Rangea-type branching, consisting of petaloids composed of several overlapping primary branches which are self-similar (fractal) over three orders of branching and attached to a straight central stalk; and Swartpuntia-type branching, consisting of petaloids with unornamented tube-like primary branches. Traditional Ediacaran taxonomy has emphasized unity of fronds as a high-level taxon, but the frond morphology most likely represents convergent evolution resulting from competition for nutrients in the water column. It is recommended that any new, higher-order classification schemes highlight the differences in branching architecture as a way of recognizing clades within the Ediacara biota.     

In situ filamentous communities from the Ediacaran (approx. 563 Ma) of Brazil

Precambrian filamentous microfossils are common and diverse. Nevertheless, their taxonomic assignment can be difficult owing to their overall simple shapes typically lacking in diagnostic features. Here, we report in situ communities of well-preserved, large filamentous impressions from the Ediacaran Itajaí Basin (ca 563 Ma) of Brazil. The filaments are uniserial (unbranched) and can reach up to 200 µm in width and up to 44 mm in length. They occur as both densely packed or sparsely populated surfaces, and typically show a consistent orientation. Although simple in shape, their preferred orientation suggests they were tethered to the seafloor, and their overall flexibility (e.g. bent, folded and twisted) supports a biological (rather than sedimentary) affinity. Biometric comparisons with modern filamentous groups further support their biological affinity, suggesting links with either large sulfide-oxidizing bacteria (SOB) or eukaryotes. Other morphological and palaeoecological characteristics further corroborates their similarities with modern large filamentous SOB. Their widespread occurrence and association with complex Ediacaran macrobiota (e.g. frondose organisms, Palaeopascichnus) suggest that they probably played an important role in the ecological dynamics of these early benthic communities by providing firm substrates for metazoans to inhabit. It is further hypothesized that the dynamic redox condition in the latest Ediacaran, with the non-continuous rise in oxygen concentration and periods of hypoxia, may have created ideal conditions for SOB to thrive.     

Ancient life and moving fluids

Over 3.7 billion years of Earth history, life has evolved complex adaptations to help navigate and interact with the fluid environment. Consequently, fluid dynamics has become a powerful tool for studying ancient fossils, providing insights into the palaeobiology and palaeoecology of extinct organisms from across the tree of life. In recent years, this approach has been extended to the Ediacara biota, an enigmatic assemblage of Neoproterozoic soft-bodied organisms that represent the first major radiation of macroscopic eukaryotes. Reconstructing the ways in which Ediacaran organisms interacted with the fluids provides new insights into how these organisms fed, moved, and interacted within communities. Here, we provide an in-depth review of fluid physics aimed at palaeobiologists, in which we dispel misconceptions related to the Reynolds number and associated flow conditions, and specify the governing equations of fluid dynamics. We then review recent advances in Ediacaran palaeobiology resulting from the application of computational fluid dynamics (CFD). We provide a worked example and account of best practice in CFD analyses of fossils, including the first large eddy simulation (LES) experiment performed on extinct organisms. Lastly, we identify key questions, barriers, and emerging techniques in fluid dynamics, which will not only allow us to understand the earliest animal ecosystems better, but will also help to develop new palaeobiological tools for studying ancient life.     

Crocodyliform Feeding Traces on Juvenile Ornithischian Dinosaurs from the Upper Cretaceous (Campanian) Kaiparowits Formation,Utah

Crocodyliforms serve as important taphonomic agents, accumulating and modifying vertebrate remains. Previous discussions of Mesozoic crocodyliform feeding in terrestrial and riverine ecosystems have often focused on larger taxa and their interactions with equally large dinosaurian prey. However, recent evidence suggests that the impact of smaller crocodyliforms on their environments should not be discounted. Here we present direct evidence of feeding by a small crocodyliform on juvenile specimens of a ‘hypsilophodontid’ dinosaur from the Upper Cretaceous (Campanian) Kaiparowits Formation of southern Utah. Diagnostic crocodyliform bite marks present on a left scapula and a right femur, as well as a partial probable crocodyliform tooth crown (ovoid in cross-section) preserved within a puncture on the right femur, comprise the bulk of the feeding evidence. Computed tomography scans of the femoral puncture reveal impact damage to the surrounding bone and that the distal tip of the embedded tooth was missing prior to the biting event. This is only the second reported incidence of a fossil crocodyliform tooth being found embedded directly into prey bone. These bite marks provide insight into the trophic interactions of the ecosystem preserved in the Kaiparowits Formation. The high diversity of crocodyliforms within this formation may have led to accentuated niche partitioning, which seems to have included juvenile dinosaurian prey.     

Environmental context for the terminal Ediacaran biomineralization of animals

In terminal Ediacaran strata of South China, the onset of calcareous biomineralization is preserved in the paleontological transition from Conotubus to Cloudina in repetitious limestone facies of the Dengying Formation. Both fossils have similar size, funnel‐in‐funnel construction, and epibenthic lifestyle, but Cloudina is biomineralized, whereas Conotubus is not. To provide environmental context for this evolutionary milestone, we conducted a high‐resolution elemental and stable isotope study of the richly fossiliferous Gaojiashan Member. Coincident with the first appearance of Cloudina is a significant positive carbonate carbon isotope excursion (up to +6‰) and an increase in the abundance and ³⁴S composition of pyrite. In contrast, δ³⁴S values of carbonate‐associated sulfate remain steady throughout the succession, resulting in anomalously large (>70‰) sulfur isotope fractionations in the lower half of the member. The fractionation trend likely relates to changes in microbial communities, with sulfur disproportionation involved in the lower interval, whereas microbial sulfate reduction was the principal metabolic pathway in the upper. We speculate that the coupled paleontological and biogeochemical anomalies may have coincided with an increase in terrestrial weathering fluxes of sulfate, alkalinity, and nutrients to the depositional basin, which stimulated primary productivity, the spread of an oxygen minimum zone, and the development of euxinic conditions in subtidal and basinal environments. Enhanced production and burial of organic matter is thus directly connected to the carbon isotope anomaly, and likely promoted pyritization as the main taphonomic pathway for Conotubus and other soft‐bodied Ediacara biotas. Our studies suggest that the Ediacaran confluence of ecological pressures from predation and environmental pressures from an increase in seawater alkalinity set the stage for an unprecedented geobiological response: the evolutionary novelty of animal biomineralization.     

Disturbance Alters the Phylogenetic Composition and Structure of Plant Communities in an Old Field System

The changes in phylogenetic composition and structure of communities during succession following disturbance can give us insights into the forces that are shaping communities over time. In abandoned agricultural fields, community composition changes rapidly when a field is plowed, and is thought to reflect a relaxation of competition due to the elimination of dominant species which take time to re-establish. Competition can drive phylogenetic overdispersion, due to phylogenetic conservation of ‘niche’ traits that allow species to partition resources. Therefore, undisturbed old field communities should exhibit higher phylogenetic dispersion than recently disturbed systems, which should be relatively ‘clustered’ with respect to phylogenetic relationships. Several measures of phylogenetic structure between plant communities were measured in recently plowed areas and nearby ‘undisturbed’ sites. There was no difference in the absolute values of these measures between disturbed and ‘undisturbed’ sites. However, there was a difference in the ‘expected’ phylogenetic structure between habitats, leading to significantly lower than expected phylogenetic diversity in disturbed plots, and no difference from random expectation in ‘undisturbed’ plots. This suggests that plant species characteristic of each habitat are fairly evenly distributed on the shared species pool phylogeny, but that once the initial sorting of species into the two habitat types has occurred, the processes operating on them affect each habitat differently. These results were consistent with an analysis of correlation between phylogenetic distance and co-occurrence indices of species pairs in the two habitat types. This study supports the notion that disturbed plots are more clustered than expected, rather than ‘undisturbed’ plots being more overdispersed, suggesting that disturbed plant communities are being more strongly influenced by environmental filtering of conserved niche traits.