Saccorhytus coronarius: a Basal Deuterostome From the Ediacaran/Cambrian Boundary

Researchers have identified traces of what they believe is the earliest known prehistoric ancestor of humans -- a microscopic, bag-like sea creature, which lived about 540 million years ago.

Named Saccorhytus, after the sack-like features created by its elliptical body and large mouth, the species is new to science and was identified from microfossils found in China. It is thought to be the most primitive example of a so-called "deuterostome" -- a broad biological category that encompasses a number of sub-groups, including the vertebrates.

If the conclusions of the study, published in the journal Nature, are correct, then Saccorhytus was the common ancestor of a huge range of species, and the earliest step yet discovered on the evolutionary path that eventually led to humans, hundreds of millions of years later.

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Evidence Ediacaran Biota Members Survived Into the Cambrian

Transitional Ediacaran–Cambrian small skeletal fossil assemblages from South China and Kazakhstan: Implications for chronostratigraphy and metazoan evolution

Authors:

Yang et al

Abstract:

The Ediacaran–Cambrian transition records distinct evolutionary changes of metazoans. The Ediacaran fossils (i.e., Ediacara-type biota and tubular fossils) are contrasting with the diverse small skeletal fossils (SSFs) from the early Cambrian. The apparent dissimilarities hindered studies deciphering their evolutionary relationships. This also led to a popular assumption that there exists a mass extinction of the Ediacara biota and cloudinids at the Precambrian–Cambrian (Pc–C) boundary. Here we report for the first time a transitional fauna which consists of typical elements of Ediacaran, i.e. cloudinids and related tubicolous organisms, together with Cambrian SSFs including protoconodonts, anabaritids and siphogonuchitids from South China and Maly Karatau (Kazakhstan). The sediments yielding the transitional fauna are characterized by siliceous rocks in both regions and were previously considered to be unfossiliferous. Their chronostratigraphic assignment in South China has been debated for years. Based on the new fossil assemblage, the siliceous strata of the Daibu Member (Northeast Yunnan, South China) and the basal Kuanchuanpu Formation (South Shaanxi, South China) can be assigned to the earliest Cambrian SSF biozone (Anabarites trisulcatus–Protohertzina anabarica Assemblage Zone) and thus can be considered of early Cambrian in age. A new subzone of the earliest SSF zone in eastern Yunnan (South China) is proposed herein defined as Ganloudina symmetrica–Rugatotheca typica Interval Subzone. The new fauna demonstrates that the cloudinids, originally confined to the late Ediacaran, persisted into the Cambrian Fortunian, and no major extinction event occurred at the Pc–C transition.

Extraordinary Claim: Venus COuld be the Source of Complex Life on Earth

The Venus Hypothesis

Author:

Cartwright

Abstract:

Current models indicate that Venus may have been habitable. Complex life may have evolved on the highly irradiated Venus, and transferred to Earth on asteroids. This model fits the pattern of pulses of highly developed life appearing, diversifying and going extinct with astonishing rapidity through the Cambrian and Ordovician periods, and also explains the extraordinary genetic variety which appeared over this period.

Evidence of Multiple Increases in Paleoocean Oxygen Levels Across the Ediacaran/Cambrian Boundary

REE and trace element patterns from organic-rich rocks of the Ediacaran-Cambrian transitional interval

Authors:

Guo et al

Abstract:

It has been established that important changes in the marine environment and the biosphere occurred during the Cambrian. However the relationships between the so-called “ Cambrian Explosion” and the concomitant environmental changes are not yet fully understood. This study presents new geochemical data from the black shale successions from different facies belts of the Yangtze Platform in South China. Variations in the concentrations of REE and trace elements (varying Ce/Ce*, Th/U, V/Sc and V/Cr ratios) in kerogen as well as in bulk rocks from different depositional environments along a transect from platform to basin indicate two oxidation events, which led to the oxygenation of the water column in shallow-marine environments and euxinic conditions (weak correlation between TOC, V, U and Mo) in the deeper sea. During the first oxidation event in the late Terreneuvian, anoxic conditions in bottom waters rapidly changed to euxinic conditions. Subsequently, the second oxidation event during the early Epoch 2 of the Cambrian led to oxic-suboxic conditions in deeper seawater.

First Report of Fuxianospira from Cambrian North America

Reexamination of Yuknessia from the Cambrian of China and first report of Fuxianospira from North America

Authors:

LoDuca et al

Abstract:

Yuknessia Walcott, 1919 recently was transferred from the green algae to the Phylum Hemichordata on the basis of new details observed for the type species, Y. simplex, from the Burgess Shale Formation (Cambrian Stage 5) of British Columbia. This has prompted reexamination of material attributed to Yuknessia from various Cambrian localities in South China. Findings preclude both a Yuknessia and a hemichordate affinity for all of the Chinese study material, and most of this material is formally transferred to Fuxianospira Chen and Zhou, 1997, a taxon common in the Chengjiang biota. Comparable material from the Cambrian Marjum, Wheeler, and Burgess Shale formations of North America is also assigned to Fuxianospira, and this reassignment expands both the paleogeographic and stratigraphic range of this taxon. All aspects of the study specimens, including details obtained from scanning electron microscopy, are consistent with an algal affinity, as proposed in the original descriptions of the Chinese material.

The Earliest Known Pelagic Jellyfish Found in Stage 3 Cambrian China

The earliest pelagic jellyfish with rhopalia from Cambrian chengjiang Lagerstätte

Authors:

Han et al

Abstract:

Modern cnidarian medusae generally show a triphasic life cycle with a succession of a larva, a sessile polyp and a pelagic medusa stage. The debate around the metagenesis of sessile polyps into pelagic medusae has lasted for more than 100 years. When pelagic forms originated is not clear. Hitherto the earliest crown-group medusae have been found at Cambrian Stage 5 (traditional Middle Cambrian, 509 Ma) in Utah, while diverse stem-group medusozoans were found in the basal Cambrian Fortunian Stage. No reliable medusae have been found from Cambrian Series 2 Stage 3 (ca. 521 Ma), although the marine benthic community teemed with many phyla of bilaterians, sponges, and ctenophores. Here we reinterpret Yunnanoascus haikouensis Hu et al., 2007, originally described as a ctenophore, as a pelagic, predatory, crown-group medusozoan, based on the presence of rhopalia, possible radial canals, and marginal tentacles. The medusae were a predatory member of the pelagic food web at the middle level of the ocean at Cambrian Stage 3.

Cambrian Burgess Shale Ecologies Were Stable (!) Temporally and Geographically

Paleocommunity Analysis of the Burgess Shale Tulip Beds, Mount Stephen, British Columbia: Comparison with the Walcott Quarry and Implications for Community Variation in the Burgess Shale

Authors:

O'Brien et al

Abstract:

The Tulip Beds locality on Mount Stephen (Yoho National Park, British Columbia) yields one of the most abundant and diverse (~10,000 specimens in 110 taxa) Burgess Shale fossil assemblages in the Canadian Rockies. Detailed semi quantitative and quantitative analyses of this assemblage suggest strong similarities with the Walcott Quarry on Fossil Ridge. Both assemblages are dominated by epibenthic, sessile, and suspension feeding taxa, mostly represented by arthropods and sponges and have comparable diversity patterns, despite sharing only about half the genera. However, the Tulip Beds has a higher relative abundance of suspension feeders and taxa of unknown affinity compared to the Walcott Quarry. These biotic variations are probably largely attributable to ecological and evolutionary differences between the two temporally distinct communities that adapted to similar, but not identical, environmental settings. For instance, the Tulip Beds is farther away from the Cathedral Escarpment than the Walcott Quarry. The Tulip Beds and Walcott Quarry assemblages are more similar to each other than either one is to the assemblages of the Chengjiang biota, although the relative diversity of major taxonomic groups and ecological patterns are similar in all assemblages. The conserved diversity patterns and ecological structures among sites suggest that the ecological composition of Cambrian Burgess Shale-type communities was relatively stable across wide geographic and temporal scales.

Cambrian Arthropod Waptia From Burgess Shale Brooded its Eggs

Long before kangaroos carried their joeys in their pouches and honey bees nurtured their young in hives, there was the 508-million-year-old Waptia. Little is known about the shrimp-like creature first discovered in the renowned Canadian Burgess Shale fossil deposit a century ago, but recent analysis by scientists from the University of Toronto, Royal Ontario Museum, and Centre national de la recherche scientifique has uncovered eggs with embryos preserved within the body of the animal. It is the oldest example of brood care in the fossil record.

"As the oldest direct evidence of a creature caring for its offspring, the discovery adds another piece to our understanding of brood care practices during the Cambrian Explosion, a period of rapid evolutionary development when most major animal groups appear in the fossil record," said Jean-Bernard Caron, curator of invertebrate palaeontology at the Royal Ontario Museum and associate professor in the Departments of Earth Sciences and Ecology & Evolutionary Biology at the University of Toronto.

Caron, along with Jean Vannier at the Centre national de la recherche scientifique in Lyon, France, describe the findings in a study published December 17 in Current Biology.

Waptia fieldensis is an early arthropod, belonging to a group of animals that includes lobsters and crayfish. It had a two-part structure covering the front segment of its body near the head, known as a bivalved carapace. Caron and Vannier believe the carapace played a fundamental role in how the creature practised brood care.

"Clusters of egg-shaped objects are evident in five of the many specimens we observed, all located on the underside of the carapace and alongside the anterior third of the body," said Caron.

The clusters are grouped in a single layer on each side of the body with no or limited overlapping among the eggs. In some specimens, eggs are equidistant from each other, while in others, some are are closer together, probably reflecting variations in the angle of burial and movement during burial. The maximum number of eggs preserved per per individuals probably reached 24.

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The Implications of the Cambrian Brachiopod Fauna From the Three Gorges area of China

The Cambrian brachiopod fauna from the first-trilobite age Shuijingtuo Formation in the Three Gorges area of China

Authors:

Zhang et al

Abstract:

The Yangtze platform of South China offers evidence within its Ediacaran–Cambrian geological record of the Cambrian explosion and diversification events in metazoan history. To understand the explosive radiation of animals and the environments in which it took place, the basal Cambrian fauna succession of the Aijiahe section in the Three Gorges area, western Hubei Province, has been studied, revealing the earliest brachiopod fauna (Tsunyidiscus trilobite Zone) in this region, which was dominated numerically by Acrotretoids. This is accompanied by abundant skeletal fossils including minute well-preserved phosphatized archaeocyath cups and an assortment of abundant sponge spicules, chancelloriids, mollusks, hyoliths, and bradoriids, retrieved by acid-etching limestone interbeds in the black shale-dominated Shuijingtuo Formation (Series 2). The brachiopods comprise two species of acrotretoids, two types of botsfordiids (Botsfordiidae gen. et sp. indet. A and B), and four species of linguloids. Of the latter, Spinobolus popovi n. gen. n. sp. is strikingly distinctive and typified by spine-like ornamentation seen for the first time in the Lower Cambrian; the remaining three linguloid genera, Palaeobolus, Eoobolus, and Lingulellotreta, have a trans-paleocontinental distribution. The Three Gorges Shuijingtuo brachiopod assemblage differs from that of the upper Atdabanian Stage (Cambrian Stage 3) in Siberia and South China, but shows great similarities with those discovered in the Tsanglangpuan (equivalent to Botoman or Stage 4) Stage of eastern Yunnan Province, Siberia, and South Australia, suggesting a much more prolonged sedimentary hiatus in basalmost Shuijingtuo Formation of the Three Gorges area than previously expected. The presence of such unconformities provides a caveat to stable isotope-based correlations that involve a number of discussions of global ocean geochemical changes across the time interval that witnessed Cambrian explosion of metazoans.

The BRAAAAAAAAAAAINNNNNSSSSS of Cambrian Euarhropod Fuxianhuia protensa

Preservational Pathways of Corresponding Brains of a Cambrian Euarthropod

Authors:

Ma et al

Abstract:

The record of arthropod body fossils is traceable back to the “Cambrian explosion,” marked by the appearance of most major animal phyla. Exceptional preservation provides crucial evidence for panarthropod early radiation. However, due to limited representation in the fossil record of internal anatomy, particularly the CNS, studies usually rely on exoskeletal and appendicular morphology. Recent studies [ 1–3 ] show that despite extreme morphological disparities, euarthropod CNS evolution appears to have been remarkably conservative. This conclusion is supported by descriptions from Cambrian panarthropods of neural structures that contribute to understanding early evolution of nervous systems and resolving controversies about segmental homologies [ 4–12 ]. However, the rarity of fossilized CNSs, even when exoskeletons and appendages show high levels of integrity, brought into question data reproducibility because all but one of the aforementioned studies were based on single specimens [ 13 ]. Foremost among objections is the lack of taphonomic explanation for exceptional preservation of a tissue that some see as too prone to decay to be fossilized. Here we describe newly discovered specimens of the Chengjiang euarthropod Fuxianhuia protensa with fossilized brains revealing matching profiles, allowing rigorous testing of the reproducibility of cerebral structures. Their geochemical analyses provide crucial insights of taphonomic pathways for brain preservation, ranging from uniform carbon compressions to complete pyritization, revealing that neural tissue was initially preserved as carbonaceous film and subsequently pyritized. This mode of preservation is consistent with the taphonomic pathways of gross anatomy, indicating that no special mode is required for fossilization of labile neural tissue.

Persistent Oceanic Anoxia, Mass (?) Extinction Separate Cambrian, Ordovician Explosions

Persistent oceanic anoxia and elevated extinction rates separate the Cambrian and Ordovician radiations

Authors:

Saltzman et al

Abstract:

Recurrent mass extinction events (at "biomere"—a biostratigraphic unit—boundaries) characterize the middle Cambrian to Early Ordovician (Tremadocian) time interval that is between the major Cambrian and Ordovician radiations of animal life. A role for anoxia in maintaining elevated extinction rates in the late Cambrian has been proposed based on coincidence of an extinction with positive excursions in δ13Ccarb and δ34SCAS (CAS—carbonate-associated sulfate). Here we examine an Early Ordovician extinction event at the base of the North American Stairsian Stage (upper Tremadocian), and demonstrate concurrent onset of positive excursions in δ13C and δ34S inferred to reflect enhanced organic matter burial under anoxic waters. Sea-level rise may have brought anoxic waters onto the shelf to initiate extinctions. The evidence for δ13C excursions and elevated extinction rates appears to wane in the Tremadocian, consistent with progressive oxygenation of the oceans reaching a threshold that helped facilitate initial stages of the Great Ordovician Biodiversification Event.

Extinct Cambrian Comb Jellies Had Skeletons

A vanished history of skeletonization in Cambrian comb jellies

Authors:

Ou et al

Abstract:

Ctenophores are traditionally regarded as “lower” metazoans, sharing with cnidarians a diploblastic grade of organization. Unlike cnidarians, where skeletonization (biomineralization and sclerotization) evolved repeatedly among ecologically important taxa (for example, scleractinians and octocorals), living ctenophores are characteristically soft-bodied animals. We report six sclerotized and armored ctenophores from the early Cambrian period. They have diagnostic ctenophore features (for example, an octamerous symmetry, oral-aboral axis, aboral sense organ, and octaradially arranged ctene rows). Unlike most modern counterparts, however, they lack tentacles, have a sclerotized framework, and have eight pairs of ctene rows. They are resolved as a monophyletic group (Scleroctenophora new class) within the ctenophores. This clade reveals a cryptic history and sheds new light on the early evolution of this basal animal phylum. Skeletonization also occurs in some other Cambrian animal groups whose extant members are exclusively soft-bodied, suggesting the ecological importance of skeletonization in the Cambrian explosion.

Collinsium ciliosum: a new Filter Feeding, Armored Velvet Worm From Cambrian Paleozoic China

A new species of 'super-armoured' worm, a bizarre, spike-covered creature which ate by filtering nutrients out of seawater with its feather-like front legs, has been identified by palaeontologists. The creature, which lived about half a billion years ago, was one of the first animals on Earth to develop armour to protect itself from predators and to use such a specialised mode of feeding.

The creature, belonging to a poorly understood group of early animals, is also a prime example of the broad variety of form and function seen in the early evolutionary history of a modern group of animals that, today, are rather homogenous. The results, from researchers at the University of Cambridge and Yunnan University in China, are published today (29 June) in the journal PNAS.

The creature has been named Collinsium ciliosum, or Hairy Collins' Monster, named for the palaeontologist Desmond Collins, who discovered and first illustrated a similar Canadian fossil in the 1980s. The newly-identified species lived in what is now China during the Cambrian explosion, a period of rapid evolutionary development around half a billion years ago, when most major animal groups first appear in the fossil record.

A detailed analysis of its form and evolutionary relationships indicates that the Chinese Collins' Monster is a distant early ancestor of modern velvet worms, or onychophorans, a small group of squishy animals resembling legged worms that live primarily in tropical forests around the world.

"Modern velvet worms are all pretty similar in terms of their general body organisation and not that exciting in terms of their lifestyle," said Dr Javier Ortega-Hernández of Cambridge's Department of Earth Sciences, one of the paper's lead authors. "But during the Cambrian, the distant relatives of velvet worms were stunningly diverse and came in a surprising variety of bizarre shapes and sizes."

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paper link.

Better Fossil Makes Cambrian Ecdysozoan Hallucigenia’s Head Seem Even More Alien

Hallucigenia’s head and the pharyngeal armature of early ecdysozoans

Authors:

Smith et al

Abstract:

The molecularly defined clade Ecdysozoa comprises the panarthropods (Euarthropoda, Onychophora and Tardigrada) and the cycloneuralian worms (Nematoda, Nematomorpha, Priapulida, Loricifera and Kinorhyncha). These disparate phyla are united by their means of moulting, but otherwise share few morphological characters—none of which has a meaningful fossilization potential. As such, the early evolutionary history of the group as a whole is largely uncharted. Here we redescribe the 508-million-year-old stem-group onychophoran Hallucigenia sparsa from the mid-Cambrian Burgess Shale. We document an elongate head with a pair of simple eyes, a terminal buccal chamber containing a radial array of sclerotized elements, and a differentiated foregut that is lined with acicular teeth. The radial elements and pharyngeal teeth resemble the sclerotized circumoral elements and pharyngeal teeth expressed in tardigrades, stem-group euarthropods and cycloneuralian worms. Phylogenetic results indicate that equivalent structures characterized the ancestral panarthropod and, seemingly, the ancestral ecdysozoan, demonstrating the deep homology of panarthropod and cycloneuralian mouthparts, and providing an anatomical synapomorphy for the ecdysozoan supergroup.

If that doesn't look like an sfnal alien critter, IDK what would.

Royal Tyrrell Museum Lecture Series: On the Origin of Chordates in the Cambrian Explosion

Cambrian Exoplosion Substrate Adaptations of Sessile Benthic Animals

Substrate adaptations of sessile benthic metazoans during the cambrian radiation

Authors:

Kloss et al

Abstract:

Many marine benthic metazoans must stabilize themselves upon the seafloor for survival, and as a result their morphologies are controlled in part by local substrate conditions. The Agronomic Revolution (AR), spurred by increasing vertical bioturbation during the Ediacaran—Cambrian transition, permanently altered the nature of shallow marine substrate conditions and led to a major shift in adaptive strategies among benthic metazoans. These ecological and evolutionary changes, known as the Cambrian Substrate Revolution (CSR), are generally understood from observations of benthic metazoan fossils across the Ediacaran/Cambrian boundary, but the timing and geographic extent of this transition are less well known. This analysis attempts to constrain the temporal and spatial pattern of the AR and CSR by performing a global-scale paleoecological analysis of the adaptive strategies of benthic fauna living during the Cambrian. This analysis focused on Burgess Shale-type (BST) faunas because of their exceptional preservation, and was conducted through direct observation of fossil specimens, analysis of data compiled from the Paleobiology Database, and literature review. From these analyses, faunal groups are assigned a metric, the Substrate Adaptability Index (SAI), that relates the overall affinity the fauna demonstrates toward either Proterozoic-style (SAI=0) or Phanerozoic-style (SAI= 1) substrate conditions. The results of this analysis demonstrate that most early and middle Cambrian faunas were mixtures of Phanerozoic- and Proterozoic-style adaptive strategists, suggesting that Proterozoic-style substrates were still influential in controlling adaptive strategies in marine environments until at least that time. This is further supported by ichnofabric analysis of many of these localities, where overall bioturbation levels are exceedingly low, indicating a lack of mixed-layer development and the prevalence of firm Proterozoic-style substrates well into the Cambrian.

Yawunik kootenayi: a new Leanchoiliid Arthropod From Cambrian British Columbia

A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny

Authors:

Aria et al

Abstract:

Characterized by atypical frontalmost appendages, leanchoiliids are early arthropods whose phylogenetic placement has been much debated. Morphological interpretations have differed, some of which concern critical characters such as the number of eyes and head appendages, but methodological approaches also have diverged. Here, we describe a new leanchoiliid, Yawunik kootenayi gen. et sp. nov., based on 42 specimens from the newly discovered Marble Canyon locality of the Burgess Shale (Kootenay National Park, British Columbia; middle Cambrian). This new morphotype demonstrates the presence of a four-segmented head in leanchoiliids, along with two small antero-median eyes in addition to lateral eyes. Yawunik is characterized by a 12-segmented trunk and a carinate, lanceolate telson adorned with minute spines. The ‘great appendages’ of the animal bear teeth on their two distal rami, which would have enhanced their ability to grasp prey. Attitudes of specimens, resulting from burial at multiple aspects of bedding, suggest the ‘great appendages’ were flexible and capable of antero-posterior rotation. We also discuss the nature of intersegmental tissues and filaments present within the ‘great appendages’. Our phylogenetic analyses extend the monophyly of leanchoiliids to include Haikoucaris and Yohoia in a new clade, the Cheiromorpha nom. nov. (within Heptopodomera nom. nov.). Other nodes are poorly resolved unless implied weights are used, and in this case, the topology is critically sensitive to the coding prerogative of inapplicable states (NAs). Both the traditional ‘Arachnomorpha’ hypothesis (NAs as additional states) and the more recently favoured ‘Artiopoda + Crustacea’ (NAs as uncertainties) were obtained using the same data set and outgroup. This result stresses, first, the historical importance of polarization over data content in scenarios of early arthropod evolution, and second, a pressing need to investigate the impacts of coding inapplicables, especially given the inflating effect of implied weights.

Diversity Partitioning During the Cambrian Explosion

Diversity partitioning during the Cambrian radiation

Authors:

Na et al

Abstract:

The fossil record offers unique insights into the environmental and geographic partitioning of biodiversity during global diversifications. We explored biodiversity patterns during the Cambrian radiation, the most dramatic radiation in Earth history. We assessed how the overall increase in global diversity was partitioned between within-community (alpha) and between-community (beta) components and how beta diversity was partitioned among environments and geographic regions. Changes in gamma diversity in the Cambrian were chiefly driven by changes in beta diversity. The combined trajectories of alpha and beta diversity during the initial diversification suggest low competition and high predation within communities. Beta diversity has similar trajectories both among environments and geographic regions, but turnover between adjacent paleocontinents was probably the main driver of diversification. Our study elucidates that global biodiversity during the Cambrian radiation was driven by niche contraction at local scales and vicariance at continental scales. The latter supports previous arguments for the importance of plate tectonics in the Cambrian radiation, namely the breakup of Pannotia.

Events of the Cambrian Period

Global climate, sea level cycles, and biotic events in the Cambrian Period

Authors:

Babcock et al

Abstract:

The developing high-resolution chronostratigraphy of the Cambrian provides an updated age model for various geologic and biotic events that occurred during this critical period of Earth history. Broad, time-specific patterns of lithofacies, such as organic-rich deposits, and biofacies appear to be consistent across all Cambrian paleocontinents. Records of important evolutionary events including first appearances of certain metazoan taxa, migrations, and extinctions, tend to coincide with changes in eustatic sea level, as do the positions of many Konservat-Lagerstätten, concretion horizons, agnostoid-rich beds, and other sedimentary features. Most of these events or horizons also show a relationship to perturbations in the global carbon cycle. The positions of organic-rich deposits bear strong relationship to both paleogeographic position and sea level history. Cambrian strata show evidence of cyclicity at multiple scales. Synchronous or near-synchronous global cyclicity is inferred to be associated with oceanographic and climatic cycles characteristic of glacial expansion and deglaciation.

Was the Ediacaran/Cambrian Radiation Unique?

Was the Ediacaran—Cambrian radiation a unique evolutionary event?

Author:

Erwin

Abstract:

The extent of morphologic innovation during the Ediacaran—Cambrian diversification of animals was unique in the history of metazoan life. This episode was also associated with extensive changes in the redox state of the oceans, in the structure of benthic and pelagic marine ecosystems, in the nature of marine sediments, and in the complexity of developmental interactions in Eumetazoa. But did the phylogenetic and morphologic breadth of this episode simply reflect the unusual outcome of recurrent evolutionary processes, or was it the unique result of circumstances, whether in the physical environment, in developmental mechanisms, or in ecological interactions? To better characterize the uniqueness of the events, I distinguish among these components on the basis of the extent of sensitivity to initial conditions and unpredictability, which generates a matrix of possibilities from fully contingent to fully deterministic. Discriminating between these differences is important for informing debates over determinism versus the contingency in the history of life, for understanding the nature of evolutionary theory, and for interpreting historically unique events.