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As a significant evolutionary innovation, multicellularity has independently evolved multiple times throughout the evolutionary history of eukaryotes, making a substantial contribution to their diversity. In retracing the multicellularity of eukaryotes, deep‐time fossil records play an irreplaceable role. In this paper, we report a new acritarch Concavaesphaera ornata gen. et sp. nov. from the early Cambrian Kuanchuanpu biota (535 Ma). These fossils are generally spherical, with diameters ranging from 450 to 950 μm, and feature an envelope with complex ornament. Inside, there are two groups of cells varying in size. Larger cells are relatively fewer in number and each resides within a sac‐like cavity distributed along the inner wall of the envelope. Statistical data show a positive correlation between the diameter and cavity volume during the development of large cells. Small cells are numerous, have a diameter of about 15 μm, and fill the entire interior of the specimens. We interpret the large cells as reproductive cells and the small cells as somatic cells. These observations show that Concavaesphaera evolved multicellular complexity equivalent to that of modern Volvox, with germ‐soma differentiation and separation. Available evidence suggests that the similarities between Concavaesphaera and Volvox are more likely to be the result of convergent evolution. Therefore, Concavaesphaera may represent an extinct multicellular eukaryote which evolved during the Cambrian explosion and probably documented an independent innovation of multicellularity in eukaryotic evolutionary history. This implies that multicellularity evolved many more times in eukaryotes than previously estimated. [ABSTRACT FROM AUTHOR]

Trilobites were notably flexible in the moulting behaviours they employed, producing a variety of moult configurations preserved in the fossil record. Investigations seeking to explain this moulting variability and its potential impacts are few, despite abundant material being available for study. We present the first quantitative study on moulting in a single trilobite species using a dataset of almost 500 moult specimens of Estaingia bilobata from the Cambrian (Series 2, Stage 4) Emu Bay Shale, South Australia. Specimens were categorized by moulting mode (Salter's or Sutural Gape) and their associated configurations, and their body proportions measured from both a museum collection (including a bycatch sample) and a randomly‐collected field sample. This enabled analysis of the proportion of E. bilobata specimens displaying the Sutural Gape and Salter's modes of moulting and their different configurations, and tests for association between moulting behaviour and body proportions. The results show a wide range of E. bilobata moulting configurations in all samples, suggesting that configurations represent definable instances in a largely continuous spectrum of variation. Analyses comparing body proportions of specimens showing the two modes of moulting were non‐significant, suggesting there is no true association between moulting behaviour and body proportion, except for a single significant result for body length. All results were relatively consistent between the museum and field samples. However, removing accessioned specimens from the museum sample brought results even further in line with the field sample, supporting the need for consideration of museum collection bias in palaeontological analyses. [ABSTRACT FROM AUTHOR]

Cnidarians are phylogenetically located near the base of the ‘tree of animals’, and their early evolution had a profound impact on the rise of bilaterians. However, the early diversity and phylogeny of this ‘lowly’ metazoan clade has hitherto been enigmatic. Fortunately, cnidarian fossils from the early Cambrian could provide key insights into their evolutionary history. Here, based on a scrutiny of the purported hyolith Burithes yunnanensis Hou et al. from the early Cambrian Chengjiang biota in South China, we reveal that this species shows characters distinct from those typical of hyoliths, not least a funnel-shaped gastrovascular system with a single opening, a whorl of tentacles surrounding the mouth, and the lack of an operculum. These characters suggest a great deviation from the original definition of the genus Burithes, and a closer affinity with cnidarians. We therefore reassign the material to a new genus: Palaeoconotuba. Bayesian inference of phylogeny based on new anatomical traits identifies a new clade, including Palaeoconotuba and Cambrorhytium, as a stem group of sessile medusozoan cnidarians that are united by the synapomorphies of developing an organic conical theca and a funnel-like gastrovascular system. This study unveils a stem lineage of medusozoans that evolved a lifelong conical theca in the early Cambrian. [ABSTRACT FROM AUTHOR]

The Cambrian Period, primarily known for animal life diversifying, experienced global extinctions. Pulses of extinction in Cambrian Series 2 are exemplified by the disappearance of archaeocyath sponges and olenelline and redlichiid trilobites. However, the effect of such extinctions on outer shelf organisms, as typify Burgess Shale‐type (BST) deposits, remains relatively unknown. The phylogeny of naraoiid arthropods, represented in BST deposits globally, has consequently been reconstructed from either side of the Series 2 – Miaolingian extinction event to evaluate the response of offshore marine organisms to Cambrian environmental perturbation. As soft anatomy is known for only a subset of naraoiid species, exoskeletal morphology has proven important. Misszhouia and Naraoia (Naraoia) are distinguished morphometrically by posterior shield length/width and anterior shield length/posterior shield length. Morphometry has also been used to strengthen the identification of some cryptic naraoiid species and revise stratigraphic ranges. A revised naraoiid phylogeny reveals Misszhouia as a monophyletic subgenus, the former genus Pseudonaraoia nests within Naraoia and is placed in synonymy, and the systematic position and status of the Subfamily Liwiinae are sensitive to character weighting. Ten species of Naraoiidae range across the Series 2 – Miaolingian boundary, all naraoiid lineages originating during the main BST window. The persistence of outer shelf naraoiids throughout the Series 2 extinctions suggests that deeper offshore marine environments were resilient to extinction during periods of environmental stress. This study provides novel empirical support for the asylum of BST communities, which may contribute to the taxonomic longevity and widespread geographic distribution of taxa in these biotas. [ABSTRACT FROM AUTHOR]

Reconstructing the function and behaviour of extinct groups of echinoderms is problematic because there are no modern analogues for their aberrant body plans. Cinctans, an enigmatic group of Cambrian echinoderms, exemplify this problem: their asymmetrical body plan differentiates them from all living species. Here, we used computational fluid dynamics to analyse the functional performance of cinctans without assuming an extant comparative model. Three‐dimensional models of six species from across cinctan phylogeny were used in computer simulations of water flow. The results demonstrate that cinctans with strongly flattened bodies produced much less drag than species characterized by dorsal protuberances or swellings, suggesting the former were more stable on the seafloor. However, unlike the flattened forms, cinctans with high‐relief bodies were able to passively direct flow towards the mouth and associated food grooves, indicating that they were capable of more efficient feeding on particles suspended in the water. This study provides evidence of a previously unknown evolutionary trade‐off between feeding and stability in Cambrian cinctan echinoderms. [ABSTRACT FROM AUTHOR]

The Cambrian coral Cambroctoconus occurs selectively in the crypts of calcimicrobe Epiphyton reefs of the Zhangxia Formation (Miaolingian) in Shandong Province, North China. These cryptobionts preferentially grow laterally and/or downward, in some cases showing pendent growth from the ceilings of the reef framework. The upside‐down growth is clearly shown by the downward‐oriented aperture and the presence of fork‐like holdfasts at its base. The coral aggregations reveal two cryptic growth strategies: one involves the prevalence of far smaller individuals in vertically narrow spaces; the other involves the predominance of modular individuals in large cavernous spaces. Offset individuals frequently appear projecting downward and are connected to each other in a chain‐like fashion. Individuals often undergo rejuvenescence laterally and downward, when the growth directions may change. Therefore, the size and modularity of individuals and the direction of budding depend largely upon the cryptic microenvironments available at the time. The cryptobionts make the best use of vacant spaces by modifying the location of budding and altering their growth directions through rejuvenescence. The cryptic growth strategies found in Cambroctoconus indicate a certain degree of morphological variability in the modules. More importantly, they indicate that the individuals flexibly and exquisitely utilized the microbial crypts that were still predominant even during the middle Cambrian. These modes of cryptic growth demonstrate the age‐specific exploitation of niches by sessile skeletal organisms that was facilitated by the development of a firm attachment structure. [ABSTRACT FROM AUTHOR]

Exceptionally well‐preserved impressions of two bundles of bristles protrude from the apertures of small, spiral shells of Pelagiella exigua, recovered from the Kinzers Formation (Cambrian, Stage 4, 'Olenellus Zone', c. 512 Ma) of Pennsylvania. These impressions are inferred to represent clusters of chitinous chaetae, comparable to those borne by annelid parapodia and some larval brachiopods. They provide an affirmative test in the early metazoan fossil record of the inference, from phylogenetic analyses of living taxa, that chitinous chaetae are a shared early attribute of the Lophotrochozoa. Shells of Pelagiella exhibit logarithmic spiral growth, microstructural fabrics, distinctive external sculptures and muscle scars characteristic of molluscs. Hence, Pelagiella has been regarded as a stem mollusc, a helcionelloid expressing partial torsion, an untorted paragastropod, or a fully torted basal member of the gastropod crown group. The inference that its chaeta‐bearing appendages were anterior–lateral, based on their probable functions, prompts a new reconstruction of the anatomy of Pelagiella, with a mainly anterior mantle cavity. Under this hypothesis, two lateral–dorsal grooves, uniquely preserved in Pelagiella atlantoides, are interpreted as sites of attachment for a long left ctenidium and a short one, anteriorly on the right. The orientation of Pelagiella and the asymmetry of its gills, comparable to features of several living vetigastropods, nominate it as the earliest fossil mollusc known to exhibit evidence of the developmental torsion characteristic of gastropods. This key adaptation facilitated an evolutionary radiation, slow at first and rapid during the Ordovician, that gave rise to the remarkable diversification of the Gastropoda. [ABSTRACT FROM AUTHOR]

Hyoliths were among the earliest biomineralizing metazoans in Palaeozoic marine environments. They have been known for two centuries and widely assigned to lophotrochozoans. However, their origin and relationships with modern lophotrochozoan clades have been a longstanding palaeontological controversy. Here, we provide broad microstructural data from hyolith conchs and opercula from the lower Cambrian Xinji Formation of North China, including two hyolithid genera and four orthothecid genera as well as unidentified opercula. Results show that most hyolith conchs contain a distinct aragonitic lamellar layer that is composed of foliated aragonite, except in the orthothecid New taxon 1 that has a crossed foliated lamellar microstructure. Opercula are mostly composed of foliated aragonite and occasionally foliated calcite. These blade or lath‐like microstructural fabrics coincide well with biomineralization of Cambrian molluscs rather than lophophorates, as exemplified by the Cambrian members of the tommotiid‐brachiopod linage. Accordingly, we propose that hyoliths and molluscs might have inherited their biomineralized skeletons from a non‐mineralized or weakly mineralized common ancestor rather than as a result of convergence. Consequently, from the view of biomineralization, the homologous shell microstructures in Cambrian hyoliths and molluscs strongly strengthen the phylogenetic links between the two groups. [ABSTRACT FROM AUTHOR]

Macro‐organisms of the Ediacaran period (635–541 Ma) were large and morphologically complex, with some living in aphotic habitats, presenting the possibility that they were early animals. However, 'bizarre' Ediacaran morphologies and mouldic preservation have frustrated comparison to later taxa. Consequently, both the positions of Ediacaran biota in the tree of life and the origins of the Metazoa have remained disputed. Here we provide phylogenetic evidence to identify Ediacaran macro‐biota as animals, based on 206 new fossils of Stromatoveris psygmoglena from the lower Cambrian Chengjiang Lagerstätte. Exceptionally preserved soft‐tissue anatomy shows that Stromatoveris was a soft‐bodied, radially symmetric animal with multiple, sub‐branched petaloids and a differentiated holdfast. Photo‐referenced morphological character analysis enables phylogenetic reconstruction of a monophyletic clade designated Petalonamae, that unites Stromatoveris with iconic Ediacaran genera (Rangea, Pteridinium, Ernietta, Swartpuntia, Arborea, Pambikalbae and Dickinsonia) and is placed as sister‐group to the Eumetazoa. Therefore, based on phylogenetic bracketing within the Metazoa, the Ediacaran petalonamids are established as animals. From these findings, it follows that petalonamids remained an important component of Cambrian marine ecosystems and that the metazoan radiation can be dated to a minimum age of between 558 and 571 myr. [ABSTRACT FROM AUTHOR]

The Swedish Cambrian 'Orsten'‐type fossil sites have yielded diverse secondarily phosphatized three‐dimensionally preserved microfossils, mainly of arthropod affinities. Similar material has also been recorded from Canada, the UK, Poland, Siberia, China and Australia. Only one other non‐arthropod group, the Cycloneuralia, is commonly reported from any of these sites, leading to the general assumption that 'Orsten'‐type preservation is largely restricted to animals with a chitin‐containing cuticle. We describe here secondarily phosphatized, originally unmineralized, thread‐shaped fossils etched out of Cambrian 'Orsten'‐type deposits from the Agnostus pisiformis Biozone of the Alum Shale Formation in Sweden. These fossils strikingly resemble specimens previously described from Precambrian deposits, with at least two different morphotaxa identified (Siphonophycus kestron Schopf and Oscillatoriopsis longa Timofeev & Hermann) as well as the modern Oscillatoria. This leads us to interpret the new fossils as unbranched, uniseriate filamentous cyanobacteria. Our morphological investigations, combined with morphometrics, allow grouping the specimens assigned to O. longa into two size classes, suggesting an even higher diversity within the 'Orsten' assemblages. The lack of cyanobacterial material in any sample younger than the A. pisiformis Biozone indicates that rather drastic changes occurred in the environment, that is, substrate conditions on the Alum Shale seafloor. This coincided with a significant change in the composition of the trilobite communities and onset of the globally recognized Steptoean Positive Isotope Carbon Excursion (SPICE) in Scandinavia. [ABSTRACT FROM AUTHOR]

Abstract: Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators. [ABSTRACT FROM AUTHOR]

Chaetognaths (arrow-worms) are enigmatic in terms of their phylogenetic position, while the existence of Protosagitta spinosa from the Chengjiang Lagerstätte suggests minimal change in their unique bodyplan since at least the early Cambrian. Apart from rare (and sometimes controversial) soft-bodied remains, the fossil record of chaetognaths is otherwise almost entirely dependent on early Palaeozoic phosphatic microfossils, some of which are placed amongst so-called protoconodonts. Fused spine clusters are strikingly similar to the cephalic grasping apparatus of extant forms and are assumed to have had a comparable configuration. Here we report a new chaetognath, Ankalodous sericus gen. et sp. nov., coeval with Protosagitta but with a complex feeding apparatus consisting of multiple bundles of recurved spines whose principal function appears to have been grasping. Like all other chaetognaths a predatory mode of life is likely, but its position relative to the sediment-water interface is less certain. Reduction of the feeding apparatus, from the multi-jawed arrangement of A. sericus to the grasping spines and associated smaller teeth seen in other chaetognaths, was probably a subsequent development and conceivably was linked with a shift to a pelagic mode of life. We also report a new specimen of Protosagitta. This confirms earlier observations but it possesses hitherto unrecognized features, including a cephalic tentacle and fin rays. [ABSTRACT FROM AUTHOR]

Singuuriqia simoni gen. et sp. nov. represents the first record of a priapulid worm from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia). It is defined by an unusually broad, longitudinally folded, foregut which tapers through the pharynx towards the anterior mouth; posteriorly, the same longitudinal folding is evident in the narrow gut. The slender, smooth, trunk in the unique specimen passes anteriorly into an oval proboscis which culminates in a smooth, extensible, pharynx with pharyngeal teeth. The capacity for substantial expansion of the foregut permitted rapid ingestion of food prior to digestion at leisure. Cololites suggest both carnivorous and deposit feeding behaviour, indicating that Singuuriqia, like the present day Priapulus, was probably omnivorous. [ABSTRACT FROM AUTHOR]

Despite many major advances in recent years, three key challenges remain in bringing clarity to the early history of the phylum: (1) identifying the origin, morphology and life modes of the first brachiopods; (2) understanding the relationships of the major groups to each other and higher sister taxa; and (3) unravelling the roles of the Cambrian and Ordovician radiations that set the agenda for much of subsequent brachiopod evolution. Since some 95% of all brachiopod taxa are extinct, the fossil record is the primary source of data to frame and test models for the evolution of the phylum. The acquisition of new, and the redescription of existing faunas, in precise spatial and temporal frameworks, using new and well-established analytical and investigative techniques, are as important as ever. [ABSTRACT FROM AUTHOR]

Trilobites were capable of enroling in different ways based on the flexible articulation of thoracic segments and associated interlocking devices; the type of enrolment (spiral or sphaeroidal) is thought to have largely depended on the coaptative devices that each trilobite used to enclose the body. Based on X-ray microtomography scans of complete enrolled specimens from the Cambrian, we created three-dimensional (3D) computer models to assess the kinematics needed to achieve both enrolment types. We demonstrate that closely related trilobites with little morphological variation ( Bailiaspis?, Conocoryphe and Parabailiella) developed different enrolment types as a result of small variations in the number of thoracic segments and the angle between adjacent segments. Moreover, our models indicate that sphaeroidal enrolment, which is associated with a smaller number of thoracic segments, enabled faster encapsulation. This supports the hypothesis that there was a trend in the evolution of trilobites towards reduction in the number of thoracic segments in phylogenetically derived taxa in order to enhance the efficiency of enrolment. [ABSTRACT FROM AUTHOR]

Pengia Geyer & Corbacho is a Cambrian burlingiid trilobite with fused trunk segments devoid of any articulation in the anamorphic and epimorphic phases of development. The type species is Pengia fusilis (Peng et al.) from the Wanshania wanshanensis Zone of China. Here we describe a second species, Pengia palsgaardia sp. nov., from the Lejopyge laevigata Zone of the Paradoxides forchhammeri Superzone. It comes from a glacial erratic in Denmark which probably originated in the Alum Shale Formation of Västergötland, Sweden. Pengia palsgaardia is a large burlingiid (~10 mm in length), with 14 fused segments in the trunk whose boundaries are marked by ridges. The axis is narrow, with the axial furrows faintly indicated or effaced across the median. Laterally along the axis and the tapering glabella, symmetrical globular lobes are developed that are pinched at their base. During ontogeny the glabellar furrows are pit-like adaxially but shallow towards the axial furrow as the globular lobes develop. Their pit-like appearance in Pengia palsgaardia and some other burlingiid species is not considered similar to the condition seen in oryctocephalid trilobites. A median preglabellar ridge resembling that of Schmalenseeia Moberg develops late in ontogeny but in early ontogeny the preglabellar field resembles that of Burlingia Walcott, Alumenella Geyer & Corbacho and Niordilobites Geyer & Corbacho. This gives Pengia a more basal position in the schmalenseeid lineage, outside the derived Schmalenseeia. In mature specimens the facial sutures in P. palsgaardia are fused, but an ocular suture may have been present. During ontogeny Pengia would have gone through the anamorphic and protomeric protaspid segmental conditions, but articulation between either the cephalon and pygidium, or pygidium and thoracic segments of the trunk never developed so it did not progress beyond the protaspid phase. This extreme protomeric development is considered to be a derived feature in Pengia. [ABSTRACT FROM AUTHOR]

Identification of the primary constituents of small shelly fossil ( SSF) shells is important for explaining the evolution of SSF faunas. The characteristics and constituents of Terreneuvian tubular SSFs found in north-east Sichuan, China, are revealed using light microscopy, scanning electron microscopy and energy dispersive X-ray spectroscopy. Petrographic thin sections indicate that the chemical composition of the shells is mainly calcium carbonate with smaller amounts of phosphorus, silica and pyrite. Most of the tubular shells composed of calcium carbonate have a distinct layered structure. Evidence of replacement of the original shell by phosphatization, pyritization and silicification, and recrystallization of calcium carbonate have been found, all of which destroyed the shell's original layered structure. Most fossils treated with acetic acid are phosphatic casts or steinkerns, with some preserving organic textures of the shell as phosphatic casts. We conclude that the Terreneuvian tubular SSFs from north-east Sichuan were originally composed mainly of calcium carbonate; indeed, most Terreneuvian small skeletal fossils appear to have had an originally calcareous composition. The fossil casts or internal core fossils are composed of phosphate, which is related to local taphonomic processes. [ABSTRACT FROM AUTHOR]

Arthropods are characterized by a rigid, articulating, exoskeleton operated by a lever-like system of segmentally arranged, antagonistic muscles. This skeletomuscular system evolved from an unsegmented body wall musculature acting on a hydrostatic skeleton, similar to that of the arthropods' close relatives, the soft-bodied onychophorans. Unfortunately, fossil evidence documenting this transition is scarce. Exceptionally-preserved panarthropods from the Cambrian Lagerstätte of Sirius Passet, Greenland, including the soft-bodied stem-arthropod Pambdelurion whittingtoni and the hard-bodied arthropods Kiisortoqia soperi and Campanamuta mantonae, are unique in preserving extensive musculature. Here we show that Pambdelurion's myoanatomy conforms closely to that of extant onychophorans, with unsegmented dorsal, ventral and longitudinal muscle groups in the trunk, and extrinsic and intrinsic muscles controlling the legs. Pambdelurion also possesses oblique musculature, which has previously been interpreted as an arthropodan characteristic. However, this oblique musculature appears to be confined to the cephalic region and first few body segments, and does not represent a shift towards arthropodan myoanatomy. The Sirius Passet arthropods, Kiisortoqia and Campanamuta, also possess large longitudinal muscles in the trunk, although, unlike Pambdelurion, they are segmentally divided at the tergal boundaries. Thus, the transition towards an arthropodan myoanatomy from a lobopodian ancestor probably involved the division of the peripheral longitudinal muscle into segmented units. [ABSTRACT FROM AUTHOR]

Omnidens is a large feeding apparatus composed of circlets of teeth, first documented from the early Cambrian of China. Originally interpreted as the oral cone of a radiodontan, it was later reinterpreted as the introvert of a priapulan. In both cases the Omnidens mouthparts underpinned estimates of gigantic ( c. 2 m) body size. Recent evidence has been used to suggest that pharyngeal teeth and radially-arranged oral plates in the stem-group onychophoran Hallucigenia and the lower stem-group euarthropod Jianshanopodia are homologous to structures of the introvert in priapulans and other scalidophorans, and are thus primitive characters for moulting animals (Ecdysozoa) as a whole. Here we show that the early Cambrian gilled lobopodian Pambdelurion whittingtoni from Sirius Passet, Greenland, possesses a mouth apparatus identical to Omnidens, being composed of the same three zones with detailed similarities of sclerites in each zone. An oral cone with rings of pharyngeal teeth, radial plates and outer scalid-like plates are ecdysozoan characters retained in the euarthropod stem group. Omnidens from China probably belongs to an unrecognized Pambdelurion-like animal rather than being part of a giant priapulan. [ABSTRACT FROM AUTHOR]

There were multiple origins of metamorphosis-undergoing protaspides in trilobite evolution: within the superfamilies Remopleuridioidea, Trinucleoidea, and within the Order Asaphida. Recent studies have revealed that the protaspides of the Cambrian representatives of the Remopleuridioidea and the Trinucleoidea did not undergo metamorphosis. However, ontogeny of the Cambrian members of the Order Asaphida has remained unknown. This study documents the ontogeny of the Furongian asaphoidean ceratopygid trilobite, Proceratopyge cf. P. lata Whitehouse, from northern Victoria Land, Antarctica. Two stages for the protaspid phase, five developmental stages for the post-protaspid cranidia, and ten stages for the post-protaspid pygidia have been identified. Interestingly, the protaspis directly developed into a meraspis without metamorphosis. A new cladistic analysis resulted in a single most parsimonious tree, according to which the presence of the bulbous commutavi protaspis turns out to be a synapomorphy for Asaphidae + Cyclopygoidea, not a synapomorphy for the Order Asaphida as previously suggested. In addition, it is inferred that there was convergent evolution of indirectly-developing commutavi protaspides during the Furongian and Early Ordovician. Metamorphosis-entailing planktonic larvae evolved in many different metazoan lineages near the Cambrian- Ordovician transition, due to the escalating ecological pressure of the Great Ordovician Biodiversification Event. Since the bulbous commutavi protaspid morphology is thought to be an adaptation for a planktonic life mode, the convergent evolution of the indirect development in the three trilobite lineages at this period might have been a result of adaptation to the early phase of the Great Ordovician Biodiversification Event. [ABSTRACT FROM AUTHOR]

The Emu Bay Shale Lagerstätte ( Cambrian Series 2, Stage 4) occurs on the north coast of Kangaroo Island, South Australia. Over 50 species are known from here, including trilobites and non-biomineralized arthropods, palaeoscolecids, a lobopodian, a polychaete, vetulicolians, nectocaridids, hyoliths, brachiopods, sponges and chancelloriids. A new chelicerate, Wisangocaris barbarahardyae gen. et sp. nov., is described herein, based on a collection of some 270 specimens. It is up to 60 mm long, with the length of the cephalic shield comprising about 30% that of the exoskeleton. The cephalic margin has three pairs of bilaterally-symmetrical small triangular spines. A pair of small eyes is placed well forwards on the ventral margin of the cephalic shield. The trunk comprises 11 segments that increase in length while narrowing posteriorly, each possibly bearing a pair of biramous appendages; the most posterior segment is almost square whereas the others are transversely elongated. The spatulate telson is proportionately longer than in taxa such as Sanctacaris, Utahcaris and Leanchoilia. Up to eight (?four pairs) of 3 mm-long elements bearing alternating inward-curving short and long spines beneath the cephalic shield are interpreted as basipodal gnathobases, part of a complex feeding apparatus. A well-developed gut includes a stomach within the cephalic shield; it extends to the base of the telson. In a few specimens there are shell fragments within the gut, including those of the trilobite Estaingia bilobata (the most common species in the biota); these fragments have sharp margins and extend across the gut lumen. The species may have been a predator or a scavenger, ingesting material already broken up by a larger predator/scavenger. The morphology of this taxon shares many overall body features with Sanctacaris, and some with Sidneyia, particularly its gnathobasic complex. These chelicerate affinities are corroborated by phylogenetic analyses. [ABSTRACT FROM AUTHOR]

Fossilized embryos afford direct insight into the pattern of development in extinct organisms, providing unique tests of hypotheses of developmental evolution based in comparative embryology. However, these fossils can only be effective in this role if their embryology and phylogenetic affinities are well constrained. We elucidate and interpret the development of Olivooides from embryonic and adult stages and use these data to discriminate among competing interpretations of their anatomy and affinity. The embryology of Olivooides is principally characterized by the development of an ornamented periderm that initially forms externally and is subsequently formed internally, released at the aperture, facilitating the direct development of the embryo into an adult theca. Internal anatomy is known only from embryonic stages, revealing two internal tissue layers, the innermost of which is developed into three transversally arranged walls that partly divide the lumen into an abapertural region, interpreted as the gut of a polyp, and an adapertural region that includes structures that resemble the peridermal teeth of coronate scyphozoans. The anatomy and pattern of development exhibited by Olivooides appears common to the other known genus of olivooid, Quadrapyrgites, which differs in its tetraradial, as opposed to pentaradial symmetry. We reject previous interpretations of the olivooids as cycloneuralians, principally on the grounds that they lack a through gut and introvert, in embryo and adult. Instead we consider the affinities of the olivooids among medusozoan cnidarians; our phylogenetic analysis supports their classification as total-group Coronata, within crown- Scyphozoa. Olivooides and Quadrapyrgites evidence a broader range of life history strategies and bodyplan symmetry than is otherwise commonly represented in extant Scyphozoa specifically, and Cnidaria more generally. [ABSTRACT FROM AUTHOR]

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian ( Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian ( Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral-like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube-like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian ( Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef-building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well-developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle. [ABSTRACT FROM AUTHOR]

Due to inadequate preservation, pterobranchs are often difficult to identify in the fossil record, and a better understanding of preservational modes and diagenetic and metamorphic effects is needed for their recognition. Pterobranch hemichordates are common in Cambrian Stage 5 and younger sedimentary rocks, but are frequently overlooked. Often, pterobranch hemichordate colonies have been considered to be algal remains or hydroids. Re-examination of Cambrian Burgess Shale algae reveals that the genera Yuknessia and Dalyia can be recognized as putative early representatives of pterobranch hemichordates. Distinct fusellar construction of the individual zooidal tubes and branching of the creeping proximal part of the colonies are found in the morphologically similar rhabdopleurid pterobranch genus Sphenoecium. The erect tubes of Sphenoecium do not branch and can reach a length of several centimetres. The development of the fusellar construction in this taxon shows a highly irregular development of the suture patterns, but a fairly consistent height of the individual fuselli. The taxon is widely distributed in the Cambrian Series 3, but has regularly been identified as a hydroid or an alga. Sphenoecium wheelerensis from the Cambrian Wheeler Shale of Utah is described as new. [ABSTRACT FROM AUTHOR]

Palaeoscolecid worms are a ubiquitous group of Early Palaeozoic ecdysozoans that are curiously lacking in the archetypal Cambrian Lagerstätten, the Burgess Shale. Here I describe Scathascolex minor gen. et sp. nov, the first unequivocal palaeoscolecid from this site. Scathascolex is armoured with simple Hadimopanella-like plates, but lacks smaller platelets, pointing to a close affinity with the Palaeoscolecida sensu stricto. Neither preservational nor environmental factors account for the scarcity of palaeoscolecids in the Burgess Shale, which presumably represents an ecological phenomenon. [ABSTRACT FROM AUTHOR]

Diploblastic eumetazoans of the phylum Cnidaria originated during the Neoproterozoic Era, possibly during the Cryogenian Period. The oldest known fossil cnidarians occur in strata of Ediacaran age and consist of polypoid forms that were either nonbiomineralizing or weakly so. The oldest possible anthozoans, including the genus Ramitubus, may be related to tabulate corals and occur in the Doushantuo Lagerstätte (upper Doushantuo Formation, South China), the age of which is poorly constrained (approximately 585 Ma?). Conulariid scyphozoans may first appear as early as 635-577 Ma ( Lantian Formation, South China). A definite conulariid, most similar to Palaeozoic species assigned to the genus Paraconularia, occurs in association with the possible scyphozoan, Corumbella werneri, in the latest Ediacaran ( c. 543 Ma) Tamengo Formation of Brazil. Basal Cambrian ( c. 540 Ma) phosphorites in the upper Kuanchuanpu Formation (South China) yield solitary polyps of the oldest probable anthozoan ( Eolympia pediculata), which appears to have been a stem hexacorallian. This same formation contains fossils interpreted by some authors as pentaradial cubozoan polyps; however, both the oldest known cubozoans and the oldest hydrozoans, all medusae, may actually occur in the Cambrian (Series 3, c. 505 Ma) Marjum Formation (Utah, USA). Although these recently published palaeontological data tend to corroborate the hypothesis that Cnidaria has a relatively deep Neoproterozoic history, the timing of major internal branching events remains poorly constrained, with, for example, the results of some molecular clock analyses indicating that the two cnidarian subphyla ( Anthozoaria and Medusozoa) may have originated as many as one billion years ago. Further progress towards elucidating the evolution and early fossil record of cnidarians may accrue from: (1) an intensive search for phosphatized soft parts in possible anthozoans from the Ediacaran Doushantuo Formation; (2) an expanded search for Ediacaran conulariids; and (3) additional detailed analyses of the taphonomy and preservation of Ediacaran and Cambrian cnidarians, including possible pentaradial cubozoan polyps from the Fortunian upper Kuanchuanpu Formation. [ABSTRACT FROM AUTHOR]

The ability to enrol effectively evolved several times among trilobites. Here, we show that, unlike most redlichiid trilobites that could not enrol, both morphotypes of Eccaparadoxides pradoanus from the middle Cambrian of Spain enrolled so as to enclose most of the ventral surface beneath the exoskeleton and possessed specialized articulating devices that facilitated this behaviour. The holaspid thorax of all E. pradoanus was divided into two principal regions. The boundary between these marked a transition from anterior segments with short pleural spines, fulcra and ridge-and-groove inner pleural regions to posterior segments with longer, acuminate pleural spines that lack fulcra and inner pleural regions. Devices that aid articulation, such as fulcra with short articulating pleural surfaces, the petaloid articulating facet and long articulating half rings, are concentrated in the anterior region. These features, and the large number of specimens preserved in various degrees of enrolment, suggest an enrolment procedure in which the rear part of the trunk, containing both the posterior thorax and the pygidium, rotated as a single unit without significant internal flexure. As these posterior trunk articulations were apparently not required to permit enrolment, concentrating flexure in the anterior may have presaged the caudalized condition seen in many derived trilobite groups that encapsulated, in which a larger proportion of the trunk segments were allocated to the mature pygidium, and therefore unable to articulate. [ABSTRACT FROM AUTHOR]

The ontogeny of the trilobite Protopeltura aciculata (Angelin, 1854) is described on the basis of material from the upper Cambrian (Furongian) of Andrarum (Skåne) and Hjelmsäter (Västergötland), Sweden. P. aciculata is present in the Parabolina brevispina and Parabolina spinulosa zones. Protopeltura aciculata is represented by all stages of growth, from early protaspides to holaspides, although most of the specimens are disarticulated and precise degrees are unknown. The cranidia have therefore been allocated to five morphological groups. Cuticular sculpture of the cranidia changes throughout ontogeny. Large tubercles are present in earlier stages, disappear gradually in middle meraspid stages and are replaced with a very faint granulation. The transitory pygidium, relatively large and shield-shaped with upwardly and backwardly directed marginal spines in early meraspides, later becomes very small, triangular-shaped and lacking spines as a late meraspid and holaspid. The development of hypostomes and librigenae is also described. Protopeltura aciculata shows major intraspecific variations throughout development, especially regarding the pygidium where variation is much less constrained than in many other olenids. This high developmental plasticity may be a survival strategy for a trilobite living in a stressed environment. Protopeltura inhabited a dysoxic environment, possibly unusually prone to localised spreading of anoxic or toxic water. Some morphs may have been less vulnerable than others to such stresses, surviving by chance and thus enabling the species to continue. [ABSTRACT FROM AUTHOR]

The complex history of description of the anomalocaridids has partly been caused by the fragmentary nature of these fossils. Frontal appendages and mouth parts are more readily preserved than whole-body assemblages, so the earliest work on these animals examined these structures in isolation. After several decades of research, these disarticulated elements were assembled together to reconstruct the anomalocaridid body plan, and a total of three Burgess Shale genera, Anomalocaris, Laggania and Hurdia, were described in full. Here we present new frontal appendage material of additional anomalocaridid taxa from the ‘Middle’ Cambrian (Series 3) Burgess Shale Formation in Canada, showing that the diversity of anomalocaridids in this locality is even higher than previously thought. Material includes Amplectobelua stephenensis sp. nov. , the first known occurrence of this genus outside of China; Caryosyntrips serratus gen. et sp. nov., which is similar to the Anomalocaris appendage but has a straighter outline and a different arrangement of spines; and an appendage that may be either the Laggania appendage or a third morph of the Hurdia appendage. The new anomalocaridid material is contemporaneous with the previously described taxa Anomalocaris, Laggania, and Hurdia, and the differences in morphology between the frontal appendages may reflect different feeding strategies. The stratigraphically lowest locality, S7 on Mount Stephen, yields material from all anomalocaridid taxa, but the assemblages in the younger quarries on Fossil Ridge are dominated by Anomalocaris and Hurdia only. [ABSTRACT FROM AUTHOR]

Cambrian trilobites mainly lived on the sea floor, and up till now few, if any, unequivocally planktonic trilobites have been reported from earlier than the Ordovician. The late Cambrian (Furongian) to late Ordovician olenids are a distinctive group of benthic (sea-floor dwelling) or nekto-benthic trilobites. Here we show, however, that one recently described, miniaturized and very spiny olenid species, Ctenopyge ceciliae must have been planktonic (passively drifting or feebly swimming in the upper waters of the sea). This interpretation is based not only upon body form but also on the analysis of its visual system and may be one of the earliest records of the planktonic realm being invaded by trilobites. [ABSTRACT FROM AUTHOR]

Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation ( Ptychagnostus atavus/ Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi-nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite-shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre-like microstructures in many of these fossils reveals how widespread these strong (fracture-resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov. [ABSTRACT FROM AUTHOR]

Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft-bodied features previously unknown from this Burgess Shale-type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better-known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada. [ABSTRACT FROM AUTHOR]

The Ocruranus– Eohalobia group, whose members were variously considered to be brachiopods, bivalves, chitons, tommotiids and coeloscleritophorans, are difficult to classify because of lack of morphological detail and evidence for skeletal reconstruction. New specimens from South China reveal more information about Ocruranus– Eohalobia and allow progress towards deciphering the skeletal reconstruction and phylogenetic affinity of this enigmatic group. Many specimens have a phosphatic inner and outer coat (mould) with empty space in between that resulted from dissolution of the original shell. Moreover, many of the internal moulds show a previously unknown type of shell microstructure that consisted of stacked layers of highly organized, acicular crystallites that radiated from the apex of the shell towards the aperture. The dissolved shell and needle-like crystals suggest an original calcareous, probably aragonitic, shell mineralogy. A few specimens also show a polygonal texture in regions that suggests the shell had a thin, prismatic inner shell microstructure. Ocruranus and Eohalobia belong to the same skeleton, and we herein synonymize Eohalobia with the older Ocruranus. Moreover, new specimens from Meishucun reveal a third type of shell plate, similar in form and inferred placement to intermediate valves of chitons. Ocruranus is likely a mollusc, and possibly a member of the chiton stem lineage. If so, then the beginning of the known record of chitons would be extended back from late Cambrian ( Saukia Zone; Furongian) to early Cambrian (Meishucunian; Series 1). [ABSTRACT FROM AUTHOR]

Extinct clades whose higher-level relationships are problematic or disputed pose a particular problem for phylogenetic analysis because of the difficulty of establishing a convincing root. Cinctans are a problematic group of primitive preradiate echinoderms whose morphology is so distinct that few characters can be usefully homologized with any other group. Here we use a two stage approach, first constructing an unrooted network using the totality of phylogenetically informative characters in which individual plates are homologized, and then determining the root position with a much smaller set of recast characters that can be polarized by outgroup and/or ontogenetic criteria. Furthermore, when rooting is based on stratigraphic criteria the same two major derived clades are identified, although basal relationships are somewhat different. We contend that the congruence of results obtained using our various approaches provides confidence that we have correctly rooted the cladogram. A revised taxonomy for Cincta is presented. [ABSTRACT FROM AUTHOR]

A new arthropod, Synophalos xynos gen. et sp. nov., is described from the Lower Cambrian of the Chengjiang Lagerstätte, Yunnan, China. It is provisionally assigned to Crustacea, Waptiida and Waptiidae. Just one example of an isolated individual of S.  xynos is known. Typically, individuals of the new species occur together in similar, robustly integrated chain-like associations. Such a strongly linked configuration is apparently unique for any arthropod, fossil or living. The chain-like association is suggested to represent an example of collective behaviour, possibly associated with reproduction, or more likely migration. [ABSTRACT FROM AUTHOR]

Continuing debate over the evolution and morphology of the arthropod head has led to considerable interest in the relevance of the evidence from the fossil record. However, dispute over homology and even presence of appendages and sclerites in Cambrian arthropods has resulted in widely differing views of their significance. The head structures of several important taxa, Fuxianhuia, Canadaspis, Odaraia, Chengjiangocaris and Branchiocaris are redescribed, revealing the essential similarity between them. In particular, all possessed an anterior sclerite, probably followed by a large posterior, ventral sclerite that is likely to be homologous to the hypostome of trilobites. The presence of a similar feature in Sanctacaris is also possible, but less well-supported. An anterior sclerite, usually bearing eyes, as in Fuxianhuia, appears to be a widespread feature of basal arthropods. Whether or not this sclerite represents an original articulating protocerebral segment on its own is, however, open to debate. [ABSTRACT FROM AUTHOR]

The linguliform brachiopod Eoobolus from the Early Cambrian Mural Formation (Jasper National Park, Canadian Rocky Mountains) exhibits various calcitic features in its otherwise apatitic shell. It is argued here that the decomposition of the organic matter within the shell led to a microenvironment similar to those resulting in the phosphatization of soft tissues. This diagenetic regime encouraged the initial precipitation of apatite cements followed by calcite cements. By fully coating primary structures early apatite cements separate primary structures from the later precipitation of calcite cement. Round calcareous grains, about 3 µm in size, that occur in the centre of apatite botryoids must therefore represent original components of the shell. The equivalent pits of such calcareous granules are seen in the larval shells of many Palaeozoic linguliform brachiopods. This suggests that mixed organophosphatic-calcareous shells were relatively common at that time but that they have been overlooked owing to the obliteration of original calcareous structures by traditional acid preparation methods for the extraction of phosphatic fossils. The Eoobolus shell structure is intermediate between purely organophosphatic and calcitic shells. Although one such genus is not sufficient to reconstruct the ancestral composition of the brachiopod shell, it provides a means of recognizing other transitional forms that are needed to understand fully the shift in shell mineralogy. [ABSTRACT FROM AUTHOR]

The collection, since 1975, of over 1500 specimens of Leanchoilia Walcott by the Royal Ontario Museum has prompted reassessment of the genus and its species from the Middle Cambrian Burgess Shale. Among new characters in Leanchoilia superlata, the type species, are the presence of two pairs of eyes, a dorsal double carina bracketing the axis of the body segments, segmentation of the gill branch of the appendages, and serration along the body edges from the posterior third of the cephalic shield to the last body segment. Leanchoilia persephone Simonetta, previously synonymized with L. superlata, is also well represented in the Burgess Shale, and is re-established as a valid species, owing to conspicuous differences from the type species. These are the absence of the diagnostic up-curving snout of the cephalic shield, the absence of carina, the shorter ‘great appendages’, the smooth edges of the body, and its overall shape in dorsal aspect. Leanchoilia superlata and L. persephone may be sexual dimorphs of each other. The ROM collections extend considerably the geographical distribution and stratigraphic range of Leanchoilia in western Canada. [ABSTRACT FROM AUTHOR]

Current models for the exceptional preservation of Burgess Shale fossils have focused on either the HF-extractable carbonaceous compressions or the mineral films identified by elemental mapping. BSEM, EDX and microprobe analysis of two-dimensionally preserved Marpolia, Wiwaxia and Burgessia identifies the presence of both carbonaceous and aluminosilicate films for most features, irrespective of original lability. In the light of the deep burial and greenschist facies metamorphism documented for the Burgess Shale, the aluminosilicate films are identified as products of late-stage volatilization and coincident mineralization of pre-existing compression fossils, whereas the three-dimensionally preserved gut-caecal system of Burgessia is interpreted as an aluminosilicate replacement of a pre-existing carbonate phase. The case for late diagenetic emplacement of aluminosilicate minerals is supported by the extensive aluminosilicification of trilobite shell and (originally) calcareous veinlets in the Burgess Shale, as well as documentation of other secondarily aluminosilicified compression fossils. By distinguishing late diagenetic alteration from the early diagenetic processes responsible for exceptional preservation, it is possible to reconcile the range of preservational modes currently expressed in the Burgess Shale. [ABSTRACT FROM AUTHOR]

Vetulicolians have variously been considered to be unusual arthropods, stem-group deuterostomes or relatives of the tunicates. They are known from a number of Cambrian Lagerstätten, and are particularly diverse in the Chengjiang biota of Yunnan Province, China. We recognize two classes, Vetulicolida and Banffozoa, which together form a monophyletic group. Within the Chinese collections we also identify two new species and recognize one new genus: Vetulicola monile sp. nov. and Bullivetula variola gen. et sp. nov. The evidence from new and previously described specimens is used to undertake a phylogenetic analysis and to evaluate a range of hypotheses for the affinities of vetulicolians. Given the difficulties of interpreting features in enigmatic fossils and the apparently contradictory set of characters possessed by vetulicolians, it is not possible on current evidence to reach an unequivocal conclusion regarding the phylogenetic position of the group. One possibility is that they are a sister group of arthropods that lost limbs but gained gill structures analogous to those of deuterostomes, but several features remain unexplained by this model. If they are protostomes, a more generally parsimonious position is close to the kinorhynchs. An alternative is that they are deuterostomes, although a placement at the base of the clade is not supported by the evidence. If they are deuterostomes, it is more likely that they are close to the tunicates. [ABSTRACT FROM AUTHOR]

The fossil record documents two mutually exclusive macroevolutionary modes separated by the transitional Ediacaran Period. Despite the early appearance of crown eukaryotes and an at least partially oxygenated atmosphere, the pre-Ediacaran biosphere was populated almost exclusively by microscopic organisms exhibiting low diversity, no biogeographical partitioning and profound morphological/evolutionary stasis. By contrast, the post-Ediacaran biosphere is characterized by large diverse organisms, bioprovinciality and conspicuously dynamic macroevolution. The difference can be understood in terms of the unique escalatory coevolution accompanying the early Ediacaran introduction of eumetazoans, followed by their early Cambrian (Tommotian) expansion into the pelagic realm. Eumetazoans reinvented the rules of macroecology through their invention of multitrophic food webs, large body size, life-history trade-offs, ecological succession, biogeography, major increases in standing biomass, eukaryote-dominated phytoplankton and the potential for mass extinction. Both the pre-Ediacaran and the post-Ediacaran biospheres were inherently stable, but the former derived from the simplicity of superabundant microbes exposed to essentially static, physical environments, whereas the latter is based on eumetazoan-induced diversity and dynamic, biological environments. The c. 100-myr Ediacaran transition (extending to the base of the Tommotian) can be defined on evolutionary criteria, and might usefully be incorporated into the Phanerozoic. [ABSTRACT FROM AUTHOR]

The Scandinavian trilobites described in the classical papers by Moberg and Möller (1898) and Moberg (1898) are reviewed, based on the original type collections. The ten species and subspecies originally described are here considered to represent 12 species and subspecies, ranging in age from the latest Furongian (late Cambrian) through the Tremadocian (Early Ordovician). Lectotypes are selected for Parabolina ( Parabolina) acanthura (Angelin), Acerocare ecorne Angelin and Westergaardia lata (Matthew). Leptoplastus claudicans (Moberg and Möller) is regarded as a junior subjective synonym of L. stenotus Angelin. These, and lectotypes formerly only figured by hand-drawings, are illustrated by high-resolution photographs. [ABSTRACT FROM AUTHOR]

Ontogenies are described for the first time for three species of Early Cambrian oryctocephalid trilobites: the oryctocephaline Changaspis elongata Lee, in Chien and the oryctocarines Duyunaspis duyunensis Chang and Chien, in Zhou et al. and Balangia balangensis Chien from the Balang Formation in eastern Ghuizhou Province, south-west China. The complete protaspid to holaspid ontogeny for Duyunaspis duyunensis and incomplete meraspid to holaspid ontogenies for Changaspis elongata and Balangia balangensis are described. The relative degree of ontogenetic variation within the lineage Duyunaspis duyunensis– Balangia balangensis is analysed and indicates that Balangia may have evolved from Duyunaspis by paedomorphosis. Decrease in thoracic segment number and concomitant increase in pygidial segment number in the later genus mirrors the pattern seen in the evolution of another oryctocephalid genus, Arthricocephalus. In both cases, selection was probably targeting an increase in cephalic convexity. This resulted in the evolution of a larger pygidium due to increased elevation of the axis above the substrate. The ontogenetic development of Changaspis provides evidence supporting the view that the Oryctocarinae may be considered as paedomorphic descendants of the Oryctocephalinae. [ABSTRACT FROM AUTHOR]

The Cambrian trilobite originally described as Bathyurus gregarius by Billings in 1865 is revised in the light of a new collection from what is judged to be its type locality in the Big Gully Marl Member of the Chamberlain's Brook Formation on the Cape St. Mary's Peninsula of south-east Newfoundland. Here assigned to Parasolenopleura gregaria, it is regarded as the senior synonym of Parasolenopleura cristata (Linnarsson, 1877) in the Swedish Paradoxides ( Eccaparadoxides) insularis and P. ( Plutonides) pinus zones, and Parasolenopleura lemdadensis Geyer, 1998 in the Moroccan Ornamentaspis frequens-Kymataspis arenosa zonal boundary beds. Among its features, important details of the relationships of the genal caeca to the eye and cephalic borders are given, as well as illustrations of the significant range of biotic and abiotic morphological variation in holaspid growth stages of the cephalon that will have some bearing on the revision of the Solenopleuridae. [ABSTRACT FROM AUTHOR]

A new metazoan, Skeemella clavula gen. et sp. nov., is described from the Middle Cambrian Pierson Cove Formation of the Drum Mountains, Utah, USA. Skeemella is similar to vetulicolians, but differs from other examples of this group in the relative proportions of the anterior and posterior sections, the large number of divisions, and the elongate bifid termination. The posterior section is arthropodan in character. The similarity of this fossil to vetulicolians throws hypotheses of their deuterostome affinity into question and highlights their problematic status. [ABSTRACT FROM AUTHOR]

Taxonomy of the Cambrian trilobite family Nepeidae is revised. Morphometric analyses of the generaNepeaandPenarosademonstrate that use of proportions of linear dimensions of the cranidium to differentiate species is invalid, and that infrageneric variation is continuous. The taxonomy and biostratigraphy of all Australian species of Nepeidae is revised. Species considered valid herein include:Nepea narinosa(type species),N. tonsillata,N. nans,Penarosa retifera(type species),P. elaticeps,P. rhinodelphis,P. netenta,Loxonepea loxophrys(type species) andFerenepea hispida(type species).Follicepsis considered to be a junior subjective synonym ofNepea,Trinepeais regarded as a junior subjective synonym ofPenarosa, andAscionepeais considered to be a junior subjective synonym ofFerenepea. [ABSTRACT FROM AUTHOR]

A diverse assemblage of late Early Cambrian brachiopods is described from the Bastion and Ella Island formations of North-East Greenland. The fauna includes nine species, representing all three extant brachiopod subphyla in addition to the stem group brachiopodMickwitziacf.occidens. Four linguliforms:Eoobolus priscus,Botsfordia caelata,Micromitra bella,Vandalotretasp., three rynchonelliforms:Obolella crassa,Kutorgina reticulata, and an unidentified chileid plus a possible craniiform species occur. The fauna shows similarities to late Early Cambrian (Dyeran Stage) brachiopod faunas of eastern Canada and the United States, but also to faunas from the late Early Cambrian (Botomian–Toyonian equivalent) of Australia, Antarctica and Siberia. [ABSTRACT FROM AUTHOR]

The genus Ctenopyge is known mainly from disarticulated sclerites and from rare complete specimens flattened in shales. Hitherto, very few specimens have been found preserved intact and in three dimensions. In a recently discovered fauna, however, in the Peltura minor Subzone in Västergötland, central Sweden, there occur several species of Ctenopyge, of which many are complete and superbly preserved; moreover they occur at all stages of growth. Of these the abundant Ctenopyge (Eoctenopyge) angusta Westergård, 1922 is described and reconstructed here as an adult, and the entire ontogeny is documented for all post–protaspid growth stages. Many characters typical of the adult, such as the long genal spines and the caudal spine, develop very early in ontogeny, and the relative dimensions of the cranidium do not greatly change during growth. Macropleural spines, however, develop later. The transitory pygidium, relatively large and shield–shaped in the early meraspid, later becomes very small as the ten thoracic segments are liberated; a median spine develops on the last thoracic segment only at the holaspid stage. Instar groupings can be clearly distinguished for the early stages. Recurrent associations of sclerites are interpreted as moulting configurations. As reconstructed, the genal spines are horizontal and parallel with the extended thorax; an adaptation which presumably allowed the trilobite to rest on the sea floor. [ABSTRACT FROM AUTHOR]