A Middle and Late Cambrian age for the Booley Bay Formation,County Wexford,Ireland: New acritarch data and its implications

A well preserved Ediacara-type fauna was recorded for the first time a few years ago from the bases of seven sandstone beds in the type section of the Lower Palaeozoic Booley Bay Formation in County Wexford, Ireland. Four microfossil samples from mudstones interbedded with the sandstones yielded an acritarch microflora considered indicative of a late Late Cambrian age for the Ediacara fauna, thus suggesting extension of its range from the Neoproterozoic to the Late Cambrian, rather than to the Middle Cambrian as had been previously suggested. The current study examines the microfossils in more detail, with 18 samples, all of which were productive. One additional Ediacara bed approximately 100 m above those recorded earlier was also observed. The exceptionally well preserved acritarch assemblages indicate a clear Middle Cambrian age for the Ediacara beds, thus predating the previously suggested upper limit for Ediacara faunas. The uppermost part of the formation, above known occurrences of Ediacara beds, is younger; that is, early Late Cambrian, probably coeval with the Olenus trilobite zone. In addition, the age now proposed for the Booley Bay Formation shows that it is part of the Lower Palaeozoic Ribband Group of southeast Ireland, and not part of the Cahore Group as has recently been suggested by the Irish Geological Survey. Thirty-seven acritarch species (or species groups) are distinguished and illustrated. One new species, Stellechinatum ? mariatheresae Vanguestaine, is established.     

A new type of Precambrian megascopic fossils: the Jinxian biota from northeastern China

Precambrian fossils are crucial for our understanding of the evolution of early organisms. Megascopic body fossils are more important because they potentially represent macroorganisms. However, the Precambrian fossil record is sparse and dominated by microfossils and microbial structures. Here we show a new type of megascopic fossils recovered from the Xingmincun Formation (probably Neoproterozoic age), northeastern China. The specimens are flat, flexible (easily corrugated) and discoidal in outline. Concentric or spiral ridges are preserved on both sides. Petrographical thin section examination indicates that the specimen consist of a thin layer of microcrystalline quartz grains (about 20–30 μm thick) wrapped by an outer sheath, composed primarily of chlorites. Field Emission Scanning Electron Microscopy (FE-SEM) coupled with an x-ray energy dispersive spectrometer system (EDX) analysis shows microstructures and relative element abundance of the fossils, but contributes little in solving their biological affinities. The fossils have previously been linked to discoidal impressions of the Ediacara biota. Close examination on new materials indicates that they are radically different from either the Ediacara impressions or any other Precambrian megascopic remains. Concentric or spiral ridges may result from rhythmic growth and the presence of twin specimens may suggest that the organisms undergo asexual reproduction or inhibition of growth in one direction. Referring them to any known fossil or living group has proved to be difficult. We conclude that they represent a distinct group of Precambrian megascopic organisms regardless of their affinities remaining problematic.     

Underground Vendobionta From Namibia

The late Precambrian fossils from Namibia have generally been regarded as soft-bodied organisms whose three-dimensional preservation resulted from smothering in fluidized sand. The sedimentological context of Pteridinium and Namalia within a sandstone bed, however, allows us to distinguish between two taphocoenoses: (1) winnowed, laterally collapsed, current-transported specimens accumulated as a lag deposit of turbidite-like flows, and (2) specimens 'floating' in the top part of an event bed with their vanes extending upwards to the upper bedding surface. The second taphocoenosis is interpreted as an in situ preserved 'infaunal' community. The immobile underground life habit and the bizarre modes of growth of Pteridinium and Namalia do not fit any extinct or modern group of multicellular organisms. Similar statements can be made for Ernietta and Rangea , thus reviving the Vendobionta hypothesis.     

The rise of bilaterians: a few closing comments

Freeman (2009, 2010) argue that Ediacara “faunas” should be analysed using “promorphologies” of extant animals, that rangeomorphs and erniettomorphs are colonial animals, and that Vernanimalcula is a bilaterian animal. Recent studies of Ediacara fossils have provided multiple lines of evidence that these fossils represent a sample of phylogenetically diverse marine organisms that included crown-group animals as well as stem-group animals and non-animals. Thus, it is inappropriate to analyse Ediacara fossils using “promorphologies” of extant animals. The interpretation of rangeomorphs and erniettomorphs as colonial animals is inconsistent with the functional morphologies of these Ediacara fossils. Although Vernanimalcula is a fossil, the purported germ layers of Vernanimalcula are of diagenetic origin and there is no morphological evidence in support of its bilaterian affinity.     

Restudy of the worm-like carbonaceous compression fossils Protoarenicola,Pararenicola, and Sinosabellidites from early Neoproterozoic successions in North China

The carbonaceous compression fossils Protoarenicola baiguashanensis Wang, 1982, Pararenicola huaiyuanensis Wang, 1982, and Sinosabellidites huainanensis Zheng, 1980, from the early Neoproterozoic Liulaobei and Jiuliqiao formations in northern Anhui, North China, were previously interpreted as worm-like metazoans, largely on the basis of transverse annulations and purported proboscis structures. If correct, these would be some of the earliest known bilaterian animals and would provide a key paleontological calibration to molecular clock analyses. In this study, we examine a large population of these carbonaceous fossils, clarify their taxonomy, and provide new insights into their morphological, paleoecological, and phylogenetic interpretations.Although all three species are characterized by annulated tubes, P. baiguashanensis bears a bulbous terminal structure at one end of its tube. P. huaiyuanensis is characterized by a constricted opening at one end and a closed termination at the other. The two ends of S. huainanensis tubes are both closed and round. The bulbous terminal structure in P. baiguashanensis was previously interpreted as an animal proboscis, but new observations suggest that it was more likely a holdfast structure analogous to discoidal holdfast structures of the Mesoproterozoic Tawuia-like fossil Radhakrishnania Kumar, 2001, and the frondose Ediacara fossil Charniodiscus Ford, 1958. Furthermore, it is possible that at least P. baiguashanensis and P. huaiyuanensis may represent reproductive or taphonomic fragments of the same organism. This reinterpretation weakens the previous interpretation that P. baiguashanensis and P. huaiyuanensis were worm-like bilaterian animals. Instead, they can be alternatively interpreted as erect epibenthic organisms, possibly coenocytic algae reaching a tiering height of 30 mm. The predominance of discoidal holdfasts, as opposed to rhizoidal holdfasts, in pre-Ediacaran epibenthic organisms was probably related to more stable substrates in the presence of microbial mats and in the absence of bioturbating animals.     

The first named Ediacaran body fossil, Aspidella Terranovica

Aspidella terranovica Billings, 1872 was first described from the late Neoproterozoic Fermeuse Formation (St. John's Group) on the Avalon Peninsula of eastern Newfoundland, approximately 1km stratigraphically above the famous Ediacaran biota at Mistaken Point, and several kilometres below the base of the Cambrian. Aspidella has been reinterpreted perhaps more than any other Precambrian taxon, and has variously been regarded as a fossil mollusc or ‘medusoid’, a gas escape structure, a concretion, or a mechanical suction mark. Our studies indicate that Aspidella includes a wide variety of preservational morphs varying from negative hyporeliefs with a raised rim and ridges radiating from a slit (Aspidella‐type preservation), to flat discs with a central boss and sharp outer ring (Spriggia preservation), to positive hyporeliefs with concentric ornamentation (Ediacaria preservation). Specimens occur in a continuum of sizes, with preservational styles dependent on the size of the specimen and the grain size of the host lithology; the elongation of specimens is tectonic. Aspidella is confirmed as a body fossil from observations of complex radial and concentric ornamentation, mutually deformed borders in clusters of specimens, and occurrence on the same bedding planes as certain distinctive Ediacaran taxa. Aspidella is indistinguishable from, and has priority over, several of the most common genera of late Neoproterozoic discoidal body fossils worldwide. Similar fossils from Australia are interpreted as holdfasts of frond‐like organisms. The density of specimens in the Aspidella beds suggests levels of benthic biomass in the Neoproterozoic that could rival those of modern marine communities. The serial growth forms, PalaeopascichnusIntrites, Neonereites renariusYelovichnus, associated with Aspidella, are interpreted as body fossils of unknown affinities rather than trace fossils. A new, trilobed, Ediacaran body fossil, Triforillonia costellae gen. et sp. nov., is described from the Aspidella beds of the Fermeuse Formation.     

Cnidarian taphonomy and affinities of the Ediacara biota

Plaster impressions and sand casts of extant medusae, a chondrophoran, and a pennatulid share basic structural characteristics with fossils in the Upper Proterozoic Ediacara assemblage. Impressions of extant medusae and Proterozoic circular impressions show general similarities in arrangement and position of radial and concentric structures and a central raised boss. However, annular rings and radial grooves are more numerous in the Proterozoic fossils and strongly folded or deformed fossils are rare as compared with impressions of modem medusae. Recent pennatulids yield impressions that are more deformed and irregular than the Proterozoic genus Charniodiscus. The greater frequency of deformation of most simulated fossil medusoids relative to Precambrian circular impressions implies that Proterozoic medu-soids were substantially stiffer than many modern taxa of comparable sizes. Many fossils with abundant circular rings have no constructional counterparts among the extant forms studied here and their medusoid affinities should remain in doubt. The structural simplicity of impressions of Ediacara organisms and extant cnidarians suggests that their mutual similarities may be due to convergence. However, there is no compelling morphological reason to reject the claim that some Proterozoic fossils may share affinities with living cnidarians. □ Taphonomy. Ediacara biota, cnidarians, phylogenetic relationships.     

Ediacaran microbial colonies

Enigmatic discoidal fossils are common in Neoproterozoic sedimentary sequences and in the stratigraphic record pre-date the first appearance of diverse Ediacaran fossil assemblages. Termed 'medusoids', these Neoproterozoic discoidal fossils have generally been interpreted as coelenterate-grade organisms implying a radially symmetrical body plan for ancestral eumetazoans. Analysis of exceptionally preserved discoidal fossils from the White Sea area, however, indicates that most of these discoidal forms represent colonial microbes. Localized pyritization, for example, reveals the presence of a conspicuous filamentous substructure in Ediacaria , whereas concentric rings, radial sectors and central structures in Cyclomedusa and Paliella compare directly with Recent microbial colonies growing in a nutritionally heterogeneous environment. At least some Ediacaran discoids can be compared with extant concentric ring-shaped microbial colonies that grow in hypersaline microbial mats. Insofar as most of the remaining record of Ediacaran discoids can be attributed to the holdfast structures of non-radiate modular organisms, there is no support from the fossil record for identifying a radiate ancestry for the Metazoa.     

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.     

Conulariid–like fossil from the Vendian of Russia: a metazoan clade across the Proterozoic/Palaeozoic boundary

A conulariid–like fossil is described for the first time from Upper Vendian (late Neoproterozoic) deposits. Vendoconularia triradiata gen. et sp. nov. is found with other fossils in poorly cemented, siliciclastic deposits of the lower part of the Ust'–Pinega Formation, exposed in the middle reaches of the Onega River, north of the Russian Platform. This fossil assemblage, stratigraphically lowermost in the region, includes Swartpuntia Narbonne, endemic Ventogyrus Ivantsov and Grazhdankin, Calyptrina–like tubular forms, and other new, as yet undescribed, species. Vendoconularia is described here using morphological terms developed for conulariids owing to their striking similarity. However, the six faces of the conical test and three–fold symmetry may reflect some phylogenetic connection with the larger clade of Trilobozoa, a cnidarian class that radiated during the Late Vendian–Early Cambrian and then became extinct.     

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.     

Dickinsonia from the Ediacaran Dengying Formation in the Yangtze Gorges area,South China

Dickinsonia is an iconic fossil of the Ediacara biota (~575–539 Ma). It was previously known from siliciclastic successions of the White Sea assemblage in Australia, Baltica, and possibly India. Here we describe Dickinsonia sp. from the terminal Ediacaran Shibantan Member limestone (ca. 551–543 Ma) of the Dengying Formation in the Yangtze Gorges area of South China. The stratigraphic distribution of Ediacara-type fossils in the Shibantan Member indicates that this biota uniquely preserves both the White Sea and Nama assemblages in stratigraphic succession. The new data presented here suggests that Dickinsonia had wider paleogeographic and paleoenvironmental distributions, implying its strong dispersal capability and environmental tolerance.     

A reinterpretation of Rangea schneiderhoehni and the discovery of a related new fossil from the Nama Group, South West Africa

Two new specimens of Rangea schneiderhoehni Gürich 1930 and a new fossil are here described from the Nama Group of South West Africa. A careful study of the two Rangea specimens and a reexamination of the holotype shows that R. schneiderhoehni forma plana and R. schneiderhoehni forma turgida are the same species as already suggested by Gürich, Richter, and Pflug, that the puzzling outer flange of forma plana is part of an underlying second leaf-like body, and that the Ediacara 'Rangea' does not belong to the genus Rangea . Here, therefore, it is introduced as the genus Glaessnerina . The new fossil, Nasepia altae , is related to Pteridinium simplex Gürich 1930 and Rangea schneiderhoehni Gürich 1930, and also to the problematic fossil Namalia villiersiensis Germs 1968.     

On the eve of animal radiation: phylogeny, ecology and evolution of the Ediacara biota

Ediacara fossils document an important evolutionary episode just before the Cambrian explosion and hold critical information about the early evolution of macroscopic and complex multicellular life. They also represent an enduring controversy in paleontology. How are the Ediacara fossils related to living animals? How did they live? Do they share any evolutionary patterns with other life forms? Recent developments indicate that Ediacara fossils epitomize a phylogenetically diverse biosphere, probably including animals, protists, algae, fungi and others. Their simple ecology is dominated by epibenthic osmotrophs, deposit feeders and grazers, but few if any predators. Their evolution started with an early morphospace expansion followed by taxonomic diversification within confined morphospace, and concluded by extinction of many taxa at the Ediacaran-Cambrian boundary.     

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.     

陕西镇巴下寒武统水井沱组小壳化石新属种

本文描述了陕西镇巴下寒武统水井沱组中的小壳化石３新属５新种。样品采自镇巴小洋坝曾家坡,共生化石有三叶虫、古介形虫、海绵骨针和大量软舌螺等。化石组合面貌表明其地质时代属筇竹寺期。另外,在峡东水井沱组中发现了微网虫（Ｍｉｃｒｏｄｉｃｔｙｏｎｓｐ．）化石骨片（图版Ｉ,图１８,１９）,可与云南筇竹寺组及陕南水井沱组对比,说明Ｍｉｃｒｏｄｉｃｔｙｏｎ广泛分布于扬子地台区。     

The proterozoic and earliest cambrian trace fossil record; patterns, problems and perspectives

The increase in trace fossil diversity across the Neoproterozoic-Cambrianboundary often is presented in terms of tabulations of ichnogenera.However, a clearer picture of the increase in diversity andcomplexity can be reached by combining trace fossils into broadgroups defined both on morphology and interpretation. This alsofocuses attention on looking for similarites between Neoproterozoicand Cambrian trace fossils. Siliciclastic sediments of the Neoproterozoicpreserve elongate tubular organisms and structures of probablealgal origin, many of which are very similar to trace fossils.Such enigmatic structures include Palaeopascichnus and Yelovichnus,previously thought to be trace fossils in the form of tightmeanders. A preliminary two or tripartite terminal Neoproterozoic tracefossil zonation can be be recognized. Possibly the earliesttrace fossils are short unbranched forms, probably younger thanabout 560 Ma. Typical Neoproterozoic trace fossils are unbranchedand essentially horizontal forms found associated with diverseassemblages of Ediacaran organisms. In sections younger thanabout 550 Ma a modest increase in trace fossil diversity occurs,including the appearance of rare three-dimensional burrow systems(treptichnids), and traces with a three-lobed lower surfaces.     

湖北三峡地区埃迪卡拉系灯影组“蝌蚪状”遗迹化石*

遗迹化石是埃迪卡拉纪存在两侧对称动物最有力的证据。但多数埃迪卡拉纪遗迹化石为简单、水平的表面爬迹或潜穴。在湖北三峡地区灯影组石板滩段含典型埃迪卡拉软躯体化石的地层中新发现一类形态特别的化石,呈蝌蚪状,一端膨大,一端细管状。通过对化石形态、同位素分析以及沉积学特征的研究,说明该化石为遗迹化石,而不是实体化石。蝌蚪状化石为一种复合迹,垂向活动形成球状的膨大端,平行藻席层活动形成近于水平的潜穴,反映了造迹生物垂向切穿藻席层并沿藻席层进行觅食的行为。该发现说明了在埃迪卡拉纪晚期已有两侧对称动物开始形成较为复杂的潜穴。     

Giant Bathysiphon (Foraminiferida) from Cretaceous turbidites, northern California

Straight-sided, tubular fossils occur together with a diverse assemblage of deep-water trace fossils in linegrained Franciscan turbidites at Point Saint George. Pacific Coast of California. Based on comparison with living abyssal organisms and on skeletal microstructure, these tubes are interpreted as recrystallized siliceous tests of the large abyssohenthic foraminiferid, Bathysiphon. Tubes found preserved in-place show that life orientation of the Franciscan species was vertical with the broader, mature end protruding several millimetres above the muddy seafloor. Positioned in this way the organisms probably were suspension feeders or carnivores. Bathysiphon and trace fossils taken together reflect a diverse deep-water benthic community that was disrupted (or locally eliminated) repeatedly by turbidity currents reaching the outer margins of a submarine fan. ▭ Bathysiphon, deep-water benthos, outer fan/basin floor, trace fossils, Franciscan Complex, flysch, Cretaceous.     

Proterozoic photosynthesis – a critical review

Chlorophyll‐based photosynthesis has fuelled the biosphere since at least the early Archean, but it was the ecological takeover of oxygenic cyanobacteria in the early Palaeoproterozoic, and of photosynthetic eukaryotes in the late Neoproterozoic, that gave rise to a recognizably modern ocean–atmosphere system. The fossil record offers a unique view of photosynthesis in deep time, but is deeply compromised by differential preservation and non‐diagnostic morphologies. The pervasively polyphyletic expression of modern cyanobacterial phenotypes means that few Proterozoic fossils are likely to be members of extant clades; rather than billion‐year stasis, their similarity to modern counterparts is better interpreted as a combination of serial convergence and extinction, facilitated by high levels of horizontal gene transfer. There are few grounds for identifying cyanobacterial akinetes or crown‐group Nostocales in the Proterozoic record. Such recognition undermines the results of various ancestral state reconstruction analyses, as well as molecular clock estimates calibrated against demonstrably problematic Proterozoic fossils. Eukaryotic organisms are likely to have acquired their (stem‐group nostocalean) photoendosymbionts/plastids by at least the Palaeoproterozoic, but remained ecologically marginalized by incumbent cyanobacteria until the late Neoproterozoic appearance of suspension‐feeding animals.