Diversification Of Early Animals Research Articles

The Lower Cambrian deposits of the Le Rozel Formation (Normandy, NW France): Insight into a newly described ichnofossil assemblage

The rise of morphological complexity and taxonomic diversity of trace fossils during the Cambrian Period has been a prominent focus to track the evolution and diversification of early animals. Here, we present a newly described trace fossil assemblage from the lower Cambrian Le Rozel Formation in Normandy (Northwestern France) composed of the following ichnogenera: Archaeonassa, Bergaueria, Helminthoidichnites, Helminthopsis, Nereites, Psammichnites, and Treptichnus. Trace fossils are exquisitely preserved on top of strata (i.e., epirelief), and are distributed through the entire thickness of the siliciclastic succession of about 200 m. This thick sedimentary succession shows sedimentary environments with ripples generated by unidirectional current processes or by oscillatory processes, such as hummocky cross-stratification (HCS) storm facies, tidal facies with flaser stratification, and unidirectional current ripples. In addition, syneresis cracks and microbially induced sedimentary structures (MISS) suggest that the Le Rozel Formation was deposited in a shallow marine shelf environment. New U-Pb dating of detrital zircon grains suggests a late Ediacaran maximum deposition age of 549 ± 3 Ma. This dating is supplemented by the substantial preservation of the three-dimensional burrow systems (Treptichnus pedum) and the large sediment bulldozers (Nereites, Psammichnites). Collectively, our findings suggest that the trace fossils assemblage was deposited during the Terreneuvian Epoch, beneath the first archaeocyaths and the first trilobite-bearing facies from both other lower Cambrian Carteret and Saint-Jean-de-la-Rivière Formation. The ichnofauna described from the Le Rozel Formation complements the worldwide trace fossil record around this critical time of life history.

Redox heterogeneity of the Ediacaran ocean constrained by chromium isotopes

Ocean oxygenation during the Ediacaran Period (635–541 Ma) is thought to be a potential stimulus for the emergence and diversification of early animals. However, the atmospheric and oceanic redox states during this period remain controversial. The chromium (Cr) stable isotopic system is an emerging proxy for constraining the redox states of the ocean and atmosphere over the geological time. Here, we present systematic Cr isotopic (δ53Cr) data for two Ediacaran successions in the Yangtze Block of South China: the Wangji section deposited in shallow water and the Lantian section deposited in deep water. The positively fractionated δ53Cr values at the Wangji section (up to 1.15‰ for carbonates and up to 1.11‰ for shales) imply that atmospheric oxygen had risen to a relatively high level to allow the oxidative weathering of Cr(III) during the Ediacaran. Additionally, a peak of δ53Cr values is closely correlated to the δ13C positive excursion in the upper Doushantuo Formation, likely because of the enhanced biological productivity at that time. In contrast, positively fractionated δ53Cr values are suppressed (from −0.11‰ to 0.05‰) in the shales at the Lantian section, indicating that the contemporary deep water might have still been extensively anoxic, such that the terrestrial Cr(VI) with positive δ53Cr values would have been quantitatively reduced and deposited before being transferred to the deeper settings. A comparison of these two sections of different depositional environments and their δ53Cr profiles suggests that the redox conditions of the Ediacaran ocean were highly stratified and heterogeneous.

New Facies Model and Carbon Isotope Stratigraphy for an Ediacaran Carbonate Platform From South America (Tamengo Formation—Corumbá Group, SW Brazil)

The Ediacaran is a period characterized by the diversification of early animals and extensive neritic carbonate deposits. These deposits are still not well understood in terms of facies and carbon isotope composition (δ13C). In this study we focus on the Tamengo Formation, in southwestern Brazil, which constitutes one of the most continuous and well-preserved sedimentary record of the late Ediacaran in South America. We present new detailed lithofacies and stable isotopes data from two representative sections (Corcal and Laginha) and revise the paleoenvironmental and stratigraphic interpretation of the Tamengo Formation. The Corcal section consists of neritic deposits including shallow-water limestone beds, alternated with shale and subordinate marl beds. These facies yield specimens of the Ediacaran fossils Cloudina lucianoi and Corumbella werneri. On the other hand, the Laginha section shows more heterogeneous facies, such as impure carbonates, breccias, marls, and subordinate mudstone beds, as well as no evidence of Corumbella werneri. The stable carbon isotope record is also different between the two sections, despite belonging to the same unit. The Corcal section displays higher and more homogeneous δ13C values, consistent with those of Ediacaran successions worldwide. The Laginha section, instead, displays more variable δ13C values, which suggest the influence of local and post depositional processes. The difference between the two sections was attributed to the different distance from the shore. We propose that the difference is due to topographic variations of the continental platform, which, at the Laginha site, was steeper and controlled by extensional faults. Therefore, the Corcal section is a better reference for the Tamengo Formation, whereas the Laginha is more particular and influenced by local factors. Besides, the lithofacies associations of the Tamengo Formation are like those of the Doushantuo and Dengying formatios, in South China, with no significant biogenic carbonate buildups, and different from those of other important Ediacaran units, such as the Nama Group in Nmibia and the Buah Formation in Oman. Our work highlights the complexity and heterogeneity of Ediacaran carbonate platforms and of their carbon isotopic composition. In addition, we characterize the Corcal section as a possible reference for the Ediacaran in South America.

Late Ediacaran to Early Cambrian tectonic–sedimentary controls on Lower Cambrian black shales in the Tarim Basin, Northwest China

Lower Cambrian black shales contain unique metazoan fossils that record the origins and diversification of early animals, and are also important hydrocarbon source rocks with a global distribution. The spatial distribution and hydrocarbon potential of Lower Cambrian black shales in the Tarim Basin, northwest China, are still poorly constrained due to deep burial and the few previous studies of these rocks. In this study, we integrated outcrop, seismic, and drilling data in order to examine the tectonic–sedimentary evolution of the Tarim Basin from the late Ediacaran to Early Cambrian, and its effects on the distribution of Lower Cambrian source rocks. During rifting related to breakup of the Rodinia supercontinent, the Tarim Basin was filled with clastic rocks of Cryogenian to early Ediacaran age. Post-rift carbonates interbedded with siliceous and argillaceous sediments were then deposited during a widespread transgression in the late Ediacaran–Early Cambrian. The Tarim Basin can be subdivided into northern and southern parts by a central paleo-uplift region that had a high paleo-topography. Although tectonism at the end-Ediacaran is variably recorded in some outcrops and by seismic data, the Tarim Basin did not experience a change in its tectonic and sedimentary framework at this time, indicating it inherited the structure of the Early Cambrian and late Ediacaran basins. Therefore, the spatial distribution of the Lower Cambrian source rocks was controlled by the paleo-topography of the late Ediacaran basin, and these source rocks can be divided into three members. Black siliceous shales are one of the most productive source rocks in the Tarim Basin, are widely distributed in the northern Tarim Basin, and have excellent petroleum potential.

Evolution of a key enzyme of aerobic metabolism reveals Proterozoic functional subunit duplication events and an ancient origin of animals

The biological toolkits for aerobic respiration were critical for the rise and diversification of early animals. Aerobic life forms generate ATP through the oxidation of organic molecules in a process known as Krebs’ Cycle, where the enzyme isocitrate dehydrogenase (IDH) regulates the cycle's turnover rate. Evolutionary reconstructions and molecular dating of proteins related to oxidative metabolism, such as IDH, can therefore provide an estimate of when the diversification of major taxa occurred, and their coevolution with the oxidative state of oceans and atmosphere. To establish the evolutionary history and divergence time of NAD-dependent IDH, we examined transcriptomic data from 195 eukaryotes (mostly animals). We demonstrate that two duplication events occurred in the evolutionary history of NAD-IDH, one in the ancestor of eukaryotes approximately at 1967 Ma, and another at 1629 Ma, both in the Paleoproterozoic Era. Moreover, NAD-IDH regulatory subunits β and γ are exclusive to metazoans, arising in the Mesoproterozoic. Our results therefore support the concept of an ‘‘earlier-than-Tonian’’ diversification of eukaryotes and the pre-Cryogenian emergence of a metazoan IDH enzyme.

A Chronostratigraphic Framework for the Rise of the Ediacaran Macrobiota: New Constraints from Mistaken Point Ecological Reserve, Newfoundland

Abstract The Conception and St. John’s Groups of southeastern Newfoundland contain some of the oldest known fossils of the Ediacaran macrobiota. The Mistaken Point Ecological Reserve UNESCO World Heritage Site is an internationally recognized locality for such fossils and hosts early evidence for both total group metazoan body fossils and metazoan-style locomotion. The Mistaken Point Ecological Reserve sedimentary succession includes ∼1500 m of fossil-bearing strata containing numerous dateable volcanogenic horizons, and therefore offers a crucial window into the rise and diversification of early animals. Here we present six stratigraphically coherent radioisotopic ages derived from zircons from volcanic tuffites of the Conception and St. John’s Groups at Mistaken Point Ecological Reserve. The oldest architecturally complex macrofossils, from the upper Drook Formation, have an age of 574.17 ± 0.66 Ma (including tracer calibration and decay constant uncertainties). The youngest rangeomorph fossils from Mistaken Point Ecological Reserve, in the Fermeuse Formation, have a maximum age of 564.13 ± 0.65 Ma. Fossils of the famous “E” Surface are confirmed to be 565.00 ± 0.64 Ma, while exceptionally preserved specimens on the “Brasier” Surface in the Briscal Formation are dated at 567.63 ± 0.66 Ma. We use our new ages to construct an age-depth model for the sedimentary succession, constrain sedimentary accumulation rates, and convert stratigraphic fossil ranges into the time domain to facilitate integration with time-calibrated data from other successions. Combining this age model with compiled stratigraphic ranges for all named macrofossils within the Mistaken Point Ecological Reserve succession, spanning 76 discrete fossil-bearing horizons, enables recognition and interrogation of potential evolutionary signals. Peak taxonomic diversity is recognized within the Mistaken Point and Trepassey Formations, and uniterminal rangeomorphs with undisplayed branching architecture appear several million years before multiterminal, displayed forms. Together, our combined stratigraphic, paleontological, and geochronological approach offers a holistic, time-calibrated record of evolution during the mid–late Ediacaran Period and a framework within which to consider other geochemical, environmental, and evolutionary data sets.

Suboxic-dominated conditions in the Ediacaran shallow ocean: Geochemistry of pelitic siltstones in the Doushantuo Formation, South China

The Ediacaran witnessed a significant oxygenation of the Earth's surface in line with the emergence and diversification of early animals, but the redox conditions of the shallow waters during this period are still uncertain. Here, we present geochemical profiles from neritic pelitic siltstones of the Doushantuo Formation at the village of Xiaoyang on the northwestern margin of the Yangtze Platform in South China. Most sediments show moderate to low FeHR/FeT ratios (≤0.38), low FePy/FeHR ratios (<0.7), and low enrichment factors (EF ≤ 1) for most redox-sensitive elements, suggesting predominately oxygenated shallow waters. However, we also find moderately enriched Re (EF ~10–25) combined with depleted U and Mo (EF ≤ 1), indicating suboxic conditions. Furthermore, a few samples show FeHR/FeT > 0.38, suggesting episodic anoxia in the shallow ocean. Overall, the Xiaoyang section represents a suboxic shallow ocean with occasional episodes of anoxia during the Doushantuo stage, implying that the oxygenation level in the early-mid Ediacaran ocean was still limited, even under some shallow conditions. It also supports the high redox heterogeneity in the early-mid Ediacaran ocean, which could have restricted the evolution and distribution of early metazoans.

A case for low atmospheric oxygen levels during Earth's middle history.

The oxygenation of the atmosphere - one of the most fundamental transformations in Earth's history - dramatically altered the chemical composition of the oceans and provides a compelling example of how life can reshape planetary surface environments. Furthermore, it is commonly proposed that surface oxygen levels played a key role in controlling the timing and tempo of the origin and early diversification of animals. Although oxygen levels were likely more dynamic than previously imagined, we make a case here that emerging records provide evidence for low atmospheric oxygen levels for the majority of Earth's history. Specifically, we review records and present a conceptual framework that suggest that background oxygen levels were below 1% of the present atmospheric level during the billon years leading up to the diversification of early animals. Evidence for low background oxygen levels through much of the Proterozoic bolsters the case that environmental conditions were a critical factor in controlling the structure of ecosystems through Earth's history.

Heterogeneous and dynamic marine shelf oxygenation and coupled early animal evolution.

It is generally agreed that early diversification of animals and significant rise of atmospheric and oceanic oxygen (O2) levels occurred in the Ediacaran (635-541 million years ago, Ma) and early Cambrian (ca. 541-509 Ma). The strength and nature of their relationship, however, remain unclear and debated. A recent wave of paleoredox research - with a particular focus on the fossiliferous sections in South China - demonstrates high spatial heterogeneity of oceanic O2 (redox) conditions and dynamic marine shelf oxygenation in a dominantly anoxic ocean during the Ediacaran and early Cambrian. This pattern shows a general spatiotemporal coupling to early animal evolution. We attribute dynamic shelf oxygenation to a complex interplay among the evolving atmosphere, continents, oceans, and biosphere during a critical period in Earth history. Our review supports the idea of a complex coevolution between increasing O2 levels and early diversification of animals, although additional work is required to fully delineate the timing and patterns of this coevolution and the mechanistic underpinnings.

Rise to modern levels of ocean oxygenation coincided with the Cambrian radiation of animals.

The early diversification of animals (∼630 Ma), and their development into both motile and macroscopic forms (∼575–565 Ma), has been linked to stepwise increases in the oxygenation of Earth's surface environment. However, establishing such a linkage between oxygen and evolution for the later Cambrian ‘explosion' (540–520 Ma) of new, energy-sapping body plans and behaviours has proved more elusive. Here we present new molybdenum isotope data, which demonstrate that the areal extent of oxygenated bottom waters increased in step with the early Cambrian bioradiation of animals and eukaryotic phytoplankton. Modern-like oxygen levels characterized the ocean at ∼521 Ma for the first time in Earth history. This marks the first establishment of a key environmental factor in modern-like ecosystems, where animals benefit from, and also contribute to, the ‘homeostasis' of marine redox conditions.

Limited role of functional differentiation in early diversification of animals.

The origin of most animal phyla and classes during the Cambrian explosion has been hypothesized to represent an ‘early burst’ of evolutionary exploration of functional ecological possibilities. However, the ecological history of marine animals has yet to be fully quantified, preventing an assessment of the early-burst model for functional ecology. Here we use ecological assignments for 18,621 marine animal genera to assess the relative timing of functional differentiation versus taxonomic diversification from the Cambrian to the present day. We find that functional diversity increased more slowly than would be expected given the history of taxonomic diversity. Contrary to previous inferences of rapid ecological differentiation from the early appearances of all well-fossilized phyla and classes, explicit coding of functional characteristics demonstrates that Cambrian genera occupied comparatively few modes of life. Functional diversity increased in the Ordovician and, especially, during the recoveries from the end-Permian and end-Cretaceous mass extinctions. Permanent shifts in the relationship between functional and taxonomic diversity following the era-bounding extinctions indicates a critical role for these biotic crises in coupling taxonomic and functional diversity.

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."

Resolving Difficult Phylogenetic Questions: Why More Sequences Are Not Enough

In the quest to reconstruct the Tree of Life, researchers have increasingly turned to phylogenomics, the inference of phylogenetic relationships using genome-scale data (Box 1). Mesmerized by the sustained increase in sequencing throughput, many phylogeneticists entertained the hope that the incongruence frequently observed in studies using single or a few genes [1] would come to an end with the generation of large multigene datasets. Yet, as so often happens, reality has turned out to be far more complex, as three recent large-scale analyses, one published in PLoS Biology [2]–[4], make clear. The studies, which deal with the early diversification of animals, produced highly incongruent (Box 2) findings despite the use of considerable sequence data (see Figure 1). Clearly, merely adding more sequences is not enough to resolve the inconsistencies.