A fossil Lagerstätte from the 700–750 Ma old Svanbergfjellet Formation of northeastern Spitsbergen offers a substantially enhanced view of late Proterozoic paleobiology. Fossils occur primarily as Organic‐walled compressions in shales and permineralizations in chert: secondary modes of preservation include bedding‐plane imprints and mineral replacements in apatite and goethite (?). The depositional setting of all fossiliferous horizons is broadly peritidal with highest taxonomic diversity occurring in shallow subtidal settings; the details of included fossil assemblages contribute to improved paleoecological resolution. The often distinct constituents of shale‐and chert‐hosted fossil assemblages appear to be a product of both paleoenvironment and fundamentally dissimilar taphonomic pathways, such that only forms with inferred wide ecological tolerance appear in both. Consideration of taphonomic processes also provides a variety of useful taxonomic insights, on the one hand permitting some resolution of so‐called wastebasket taxa. such as Chuaria, and on the other acknowledging the taxonomic disparity that can occur in simple forms like Siphonophycus and Oscillatoriopsis. True multicellular (including coenocytic) eukaryotes are a conspicuous Component of the Svanbergfjellet assemblage: of eight distinct taxa, one can be identified as a coenobial/colonial chlorococcalean and three as filamentous siphonocladaleans (Chlorophyta). Other forms are problematic, but several show significant cell, or possibly tissue, differentiation. A review of Proterozoic multicellular organisms reveals that a coenocytic grade of organization was common among early metaphytes and supports the view (that a cellularity is a derived condition in many ‘multicellular’ lineages. Nineteen acritarch taxa are preserved in the Svanbergfjellet sediments. Ten of these show a readily identifiable ornamentation and contribute significantly to Neoproterozoic biostratigraphy; a world‐wide and exclusively Late Riphean distribution of the acanthomorph Trachyhystrichosphaera aimika identifies it as a particularly valuable index fossil. The Svanbergfjellel fossil assemblage preserves a total of 63 distinct forms, of which 56 are treated taxonomically. As much as possible, principles of ‘natural’ taxonomy are applied, such that taphonomic and ontogenetic variants are declined separate taxonomy status. Major taxonomic revisions are offered for the acritarchs Trachyhystrichosphaera and Chuaria as well as for the prokaryotic‐grade filaments: Cephalonyx, Cyanonema, Oscillatoriopsis, Palaeolyngbya, Rugossopsis, Siphonophycus, Tortunema, and Veteronostocale. Newly erected taxa include 7 new genera: Palaeastrum. Proterocladus, Pseudotawuia, Valkyria, Cerebrosphaera, Osculosphera and Pseudodendron; 14 new species in 12 genera: Palaestrum ***dyptocranum, Proterocladus major, Proterocladus minor, Proterocladus ***hermannae. Pseudodendron birenifera, Valkyria borealis, Cerebrosphaera buickii, Osculosphaera hyaline, Pseudodendron anteridium fullerne, Germinosphaera jan...

Multicellular filaments from the ca. 1200-Ma Hunting Formation (Somerset Island, arctic Canada) are identified as bangiacean red algae on the basis of diagnostic cell-division patterns. As the oldest taxonomically resolved eukaryote on record Bangiomorpha pubescens n. gen. n. sp. provides a key datum point for constraining protistan phylogeny. Combined with an increasingly resolved record of other Proterozoic eukaryotes, these fossils mark the onset of a major protistan radiation near the Mesoproterozoic/Neoproterozoic boundary.Differential spore/gamete formation shows Bangiomorpha pubescens to have been sexually reproducing, the oldest reported occurrence in the fossil record. Sex was critical for the subsequent success of eukaryotes, not so much for the advantages of genetic recombination, but because it allowed for complex multicellularity. The selective advantages of complex multicellularity are considered sufficient for it to have arisen immediately following the appearance of sexual reproduction. As such, the most reliable proxy for the first appearance of sex will be the first stratigraphic occurrence of complex multicellularity.Bangiomorpha pubescens is the first occurrence of complex multicellularity in the fossil record. A differentiated basal holdfast structure allowed for positive substrate attachment and thus the selective advantages of vertical orientation; i.e., an early example of ecological tiering. More generally, eukaryotic multicellularity is the innovation that established organismal morphology as a significant factor in the evolutionary process. As complex eukaryotes modified, and created entirely novel, environments, their inherent capacity for reciprocal morphological adaptation, gave rise to the “biological environment” of directional evolution and “progress.” The evolution of sex, as a proximal cause of complex multicellularity, may thus account for the Mesoproterozoic/Neoproterozoic radiation of eukaryotes.

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Molecular clocks have the potential to shed light on the timing of early metazoan divergences, but differing algorithms and calibration points yield conspicuously discordant results. We argue here that competing molecular clock hypotheses should be testable in the fossil record, on the principle that fundamentally new grades of animal organization will have ecosystem-wide impacts. Using a set of seven nuclear-encoded protein sequences, we demonstrate the paraphyly of Porifera and calculate sponge͞eumetazoan and cnidarian͞bilaterian divergence times by using both distance [minimum evolution (ME)] and maximum likelihood (ML) molecular clocks; ME brackets the appearance of Eumetazoa between 634 and 604 Ma, whereas ML suggests it was between 867 and 748 Ma. Significantly, the ME, but not the ML, estimate is coincident with a major regime change in the Proterozoic acritarch record, including: (i) disappearance of low-diversity, evolutionarily static, pre-Ediacaran acanthomorphs; (ii) radiation of the high-diversity, short-lived Doushantuo-Pertatataka microbiota; and (iii) an orderof-magnitude increase in evolutionary turnover rate. We interpret this turnover as a consequence of the novel ecological challenges accompanying the evolution of the eumetazoan nervous system and gut. Thus, the more readily preserved microfossil record provides positive evidence for the absence of pre-Ediacaran eumetazoans and strongly supports the veracity, and therefore more general application, of the ME molecular clock.Porifera ͉ acritarchs ͉ Ediacaran ͉ coevolution T he sudden appearance of diverse metazoan fossils Ϸ530 million years ago (Ma) is the focus of ongoing and heated debate: Is this recording a true ''Cambrian explosion'' of early metazoan evolution or merely the onset of extensive burrowing and biomineralization (1)? Certainly the invisibility of microscopic, nonburrowing, and͞or nonbiomineralizing metazoans in the fossil record allows for the possibility of deep Proterozoic origins, so independent lines of evidence must be sought. Molecular clocks offer a potentially powerful approach for testing such evolutionary hypotheses (2), but recent analyses have yielded conspicuously discordant results. Pisani et al. (3), for example, estimate that the protostome-deuterostome ancestor evolved 900-1,100 Ma, whereas Douzery et al. (4) place this node at 642-761 Ma and Peterson et al. (5) at Ϸ570 Ma. With estimated divergence times differing by Ͼ500 million years (myr), there is clearly a need to assess both the methods used and predictions made by individual molecular-clocks analyses.Pisani et al. (3) have argued that their deep estimate for metazoan origins is robust because it agrees with a previous analysis calibrated by using different taxa, i.e., between chick and mouse (6) vs. centipedes and millipedes and spiders and horseshoe crabs (3). However, Peterson et al. (5) demonstrated that a significant rate reduction is associated with the vertebrate sequences, such that a vertebrate-calibrated clock produces a spurious 2-fold ove...