The lower Cambrian (Series 2) White Point Conglomerate (WPC) on Kangaroo Island, South Australia contains exoticclasts representing a diverse array of lithologies, including metamorphics, chert, sandstone, and abundant carbonates,notably archaeocyath-rich bioclastic limestone. Acetic acid digestion of the WPC bioclastic limestone clasts reveals adiverse shelly fauna. This assemblage includes abundant organophosphatic brachiopods such as Cordatia erinae Brockand Claybourn gen. et sp. nov., Curdus pararaensis, Eodicellomus elkaniformiis, Eohadrotreta sp. cf. E. zhenbaensis,Eoobolus sp., Kyrshabaktella davidii, and Schizopholis yorkensis. Additional shelly taxa include the solenopleurid trilobiteTrachoparia? sp., the tommotiids Dailyatia odyssei, Dailyatia decobruta Betts sp. nov., Kelanella sp., and Lapworthellafasciculata, spines of the bradoriid arthropod Mongolitubulus squamifer, and several problematica, such as Stoibostrombuscrenulatus and a variety of tubular forms. The upper age limit for the WPC is constrained by biostratigraphic data fromthe overlying Marsden Sandstone and Emu Bay Shale, which are no younger than the Pararaia janeae Trilobite Zone(Cambrian Series 2, Stage 4). The shelly fossil assemblage from the WPC limestone clasts indicates an upper Dailyatiaodyssei Zone (= Pararaia tatei to lower P. janeae trilobite zones), equivalent to the Atdabanian–early Botoman of theSiberian scheme. This contrasts with the previously suggested late Botoman age for the limestone clasts, based on the diversearchaeocyath assemblage. The minor age difference between the WPC and its fossiliferous limestone clasts suggestsrelatively rapid reworking of biohermal buildups during tectonically-active phases of deposition in the Stansbury Basin.

The most successful chronostratigraphic correlation methods enlist multiple proxies such as biostratigraphy and chemostratigraphy to constrain the timing of globally important bio- and geo-events. Here we present the first regional, high-resolution shelly fossil biostratigraphy integrated with δ13C chemostratigraphy (and corresponding δ18O data) from the traditional lower Cambrian (Terreneuvian and provisional Cambrian Series 2) of South Australia. The global ZHUCE, SHICE, positive excursions II and III and the CARE are captured in lower Cambrian successions from the Arrowie and Stansbury basins. The South Australian shelly fossil biostratigraphy has a consistent relationship with the δ13C results, bolstering interpretation, identification and correlation of the excursions. Positive excursion II straddles the boundary between the Kulparina rostrata and Micrina etheridgei zones, and the CARE straddles the boundary between the M. etheridgei and Dailyatia odyssei zones, peaking in the lower parts of the latter zone. New CA-TIMS zircon dates from the upper Hawker Group and Billy Creek Formation provide geochronologic calibration points for the upper D. odyssei Zone and corresponding chemostratigraphic curve, embedding the lower Cambrian successions from South Australia into a global chronostratigraphic context. This multi-proxy investigation demonstrates the power of integrated methods for developing regional biostratigraphic schemes and facilitating robust global correlation of lower Cambrian successions from South Australia (part of East Gondwana) with coeval terranes on other Cambrian palaeocontinents, including South and North China, Siberia, Laurentia, Avalonia and West Gondwana.

Definition of early Cambrian chronostratigraphic boundaries is problematic with many subdivisions stillawaiting ratification. Integrated multi-proxy data from well-resolved regional-scale schemes are ultimately the key to resolving broader issues of global correlationwithin the Cambrian. In Australia, early Cambrian biostratigraphy has been based predominantly on trilobites. Phosphatic shelly fauna have great potential as biostratigraphic tools, especially in pre-trilobitic strata because they are widespread and readily preserved, but they have remained underutilised. Here we demonstrate their value in a new biostratigraphic scheme for the early Cambrian of South Australia using a diverse shelly fauna including tommotiids, brachiopods, molluscs and bradoriids. Biostratigraphic data are derived from ten measured stratigraphic sections across the Arrowie Basin, targeting Hawker Group carbonates including the Wilkawillina, Wirrapowie and Ajax limestones and the Mernmerna Formation. The stratigraphic ranges of shelly fossils are predictable and repeatable across the Arrowie Basin, allowing three discrete shelly biozones to be identified, spanning Terreneuvian, Stage 2 to Series 2, Stages 3–4. The Kulparina rostrata Zone (new) and part of the overlyingMicrina etheridgei Zone (new) are pre-trilobitic (predominantly Terreneuvian). The Cambrian Series 2, Stage 3 Dailyatia odyssei Zone (new) features a very diverse shelly fauna and will be described in detail in a separate publication. These zones provide robust means to correlate Terreneuvian–Series 2 successions in neighbouring coeval basins in Australia, particularly the Stansbury Basin. Wider correlation is possible throughout East Gondwana, and especially with South China.

A lack of well resolved biostratigraphic data has prevented robust regional and global correlation of lower Cambriansuccessions from South Australia. A new early Cambrian biostratigraphy, based on data derived from 21measured stratigraphic sections and drill cores (11 described herein) reveals the abundance and diversity ofshelly fauna from the Arrowie Basin, and the value of early Cambrian “small shelly fossils” (SSF) for biostratigraphicstudies. Here we examine shelly fauna associated with the youngest of three recently establishedbiozones, the Dailyatia odyssei Taxon Range Zone (hereafter D. odyssei Zone), and their correlative potential.The D. odyssei Zone features a diverse suite of tommotiids, organophosphatic brachiopods, bradoriid arthropods,molluscs and phosphatic problematica. This fauna permits strong correlation (often at species-level) with othermajor early Cambrian terranes, particularly Antarctica, South China and Laurentia, and suggest a Cambrian Series2, Stages 3–4 age for the D. odyssei Zone. Bradoriids have proven to be useful biostratigraphic tools. Four newspeciesand three new genera are described herein: Acutobalteus sinuosus gen. et sp. nov., Eozhexiella adnyamathanha gen. etsp. nov., Manawarra jonesi gen. et sp. nov. and Mongolitubulus descensus sp. nov. The description of Eohadrotreta sp.cf. zhenbaensis represents the first occurrence of the acrotretoid brachiopod Eohadrotreta from Australia.

A new assemblage of early Cambrian bivalved arthropods (Bradoriida) is described from the Arrowie Syncline in the northern Flinders Ranges, South Australia. The well preserved, largely endemic fauna comprises a total of six taxa (including five new species): Jiucunella phaseloa sp. nov., Jixinlingella daimonikoa sp. nov., Mongolitubulus anthelios sp. nov., Neokunmingella moroensis sp. nov., Phasoia cf. spicata ( Öpik, 1968), and Sinskolutella cuspidata sp. nov. This assemblage is derived from a carbonate sedimentary package representing a high energy, shallow water archaeocyath-Renalcis biohermal facies of Terreneuvian, Stage 2 age which transitions up-section to a more restricted, low energy, intra-shelf lagoonal environment that correlates with a Cambrian Series 2, Stage 3 age. The new taxon J. phaseloa sp. nov., has a first appearance datum (FAD) in shallow water biohermal facies of the Hideaway Well Member of the Wilkawillina Limestone at a level 47 m below the FAD of Pelagiella subangulata which is taken to approximate the base of Series 2, Stage 3 in South Australia. Along with Liangshanella circumbolina, this makes J. phaseloa sp. nov. amongst the oldest bivalved arthropods in South Australia and potentially greater Gondwana. The presence of 25 bradoriid taxa from the early Cambrian of South Australia suggests East Gondwana represents a major centre of origin for the Bradoriida.

Gnathobasic spines are located on the protopodal segments of the appendages of various euarthropod taxa, notably chelicerates. Although they are used to crush shells and masticate soft food items, the microstructure of these spines are relatively poorly known in both extant and extinct forms. Here we compare the gnathobasic spine microstructures of the Silurian eurypterid Eurypterus tetragonophthalmus from Estonia and the Cambrian artiopodan Sidneyia inexpectans from Canada with those of the Recent xiphosuran chelicerate Limulus polyphemus to infer potential variations in functional morphology through time. The thickened fibrous exocuticle in L. polyphemus spine tips enables effective prey mastication and shell crushing, while also reducing pressure on nerve endings that fill the spine cavities. The spine cuticle of E. tetragonophthalmus has a laminate structure and lacks the fibrous layers seen in L. polyphemus spines, suggesting that E. tetragonophthalmus may not have been capable of crushing thick shells, but a durophagous habit cannot be precluded. Conversely, the cuticle of S. inexpectans spines has asimilar fibrous microstructure to L. polyphemus, suggesting that S. inexpectans was a competent shell crusher. This conclusion is consistent with specimens showing preserved gut contents containing various shelly fragments. The shape and arrangement of the gnathobasic spines is similar for both L. polyphemusand S. inexpectans, with stouter spines in the posterior cephalothoracic or trunk appendages, respectively.This differentiation indicates that crushing occurs posteriorly, while the gnathobases on anterior appendages continue mastication and push food towards and into the mouth. The results of recent phylogenetic analyses that considered both modern and fossil euarthropod clades show that xiphosurans and eurypterids are united as crown-group euchelicerates, with S. inexpectans placed within more basalartiopodan clades. These relationships suggest that gnathobases with thickened fibrous exocuticle, if not homoplasious, may be plesiomorphic for chelicerates and deeper relatives within Arachnomorpha. This study shows that the gnathobasic spine microstructure best adapted for durophagy has remained remarkably constant since the Cambrian.

Abundant and diverse small shelly fossils have been reported from Cambrian Series 2 in North China, but the co-occurring brachiopods are still poorly known. Herein, we describe seven genera, five species and two undetermined species of organophosphatic brachiopods including one new genus and new species from the lower Cambrian Xinji Formation at Shuiyu section, located on the southern margin of North China Platform. The brachiopod assemblage comprises one mickwitziid (stem group brachiopoda), Paramickwitzia boreussinaensis n. gen. n. sp., a paterinide, Askepasma toddense Laurie, 1986, an acrotretoid, Eohadrotreta cf. zhenbaensis Li and Holmer, 2004, a botsfordiid, Schizopholis yorkensis (Holmer and Ushatinskaya in Gravestock et al., 2001) and three linguloids, Spinobolus sp., Eodicellomus cf. elkaniiformis Holmer and Ushatinskaya in Gravestock et al., 2001 and Eoobolus sp. This brachiopod assemblage suggests a late Age 3 to early Age 4 for the Xinji Formation and reveals a remarkably strong connection with coeval faunas from East Gondwana, particularly the Hawker Group in South Australia. The high degree of similarity (even at species level) further supports a close palaeogeographic position between the North China Platform and Australian East Gondwana during the early Cambrian as indicated by small shelly fossil data.

A succession of diverse hyolith assemblages comprising 10 genera and 14 species are reported from the lower Cambrian Shangwan and Sanjianfang sections of the Xinji Formation, and Xiaomeiyao section of the Houjiashan Formation, which crop out along the southern margin of the North China Platform. Most of the specimens are represented by both conchs and opercula. The identified orthothecids include Conotheca australiensis, Cupitheca holocyclata, C. costellata, Neogloborilus applanatus, N. spinatus, Tegminites hymenodes, Triplicatella disdoma, T. xinjia sp. nov. and Paratriplicatella shangwanensis gen. et sp. nov. The hyolithids comprise Protomicrocornus triplicensis gen. et sp. nov., Microcornus eximius, M. petilus, Parkula bounites and Parakorilithes mammillatus. Some anomalous taxa possess characteristics of both Hyolithida and Orthothecida, such as C. australiensis, Neogloborilus and P. triplicensis. Protomicrocornus may constitute a sister group of other hyolithids. The teeth of Parkula bounites and clavicles of Parakorilithes mammillatus are documented for the first time. The hyolith assemblages from North China are probably coeval, and can be correlated with the Cambrian upper Stage 3–lower Stage 4. Many taxa are also globally distributed and have significant potential for biostratigraphical correlations. In accordance, the hyoliths from North China reveal closest compositional similarities to faunas from eastern Gondwana, and especially South Australia. However, some taxa are shared with Laurentian assemblages suggesting cosmopolitanism, and possibly planktonic larval dispersal.

The Cambrian radiation is characterized by the emergence of diverse bilaterian animal phyla and theestablishment of complex marine ecosystems. The Guanshan Biota records an unusual ecological transition from trilobite- to brachiopod-dominated communities during Cambrian Stage 4. This community transition is accompanied by direct evidence of in situ biological interactions such as durophagous pre-dation and kleptoparasitism. Here we describe new material from the Guanshan biota, focusing on an association of palaeoscolecidomorphs and brachiopods with parasitic tube worms that occur on micro-bedding planes. The bedding plane assemblages are dominated by the organophosphatic brachiopod Neobolus wulongqingensis encrusted with kleptoparasitic tube-dwelling worms, along with infaunal palaeoscolecidans. Taphonomic and sedimentological evidence indicates that these specimens are com-monly preserved in life position, and thus the association between individuals represent potential biological interactions. This case study reveals that ecosystems during the early Cambrian exhibited a well-developed system of tiering and a complex trophic network, easily distinguished from the simple communities typical of precursor deposits in the Ediacaran. Brachiopods forming extremely dense concentrations on the sea floor are effectively acting as ecosystem engineers, not only to stabilize the soft-substrate seafloor, but also act as an alternative substrate for the oldest empirically demonstrated kleptoparasites.The in situ biological interactions preserved in the Guanshan Biota are critical for filling gaps in ourknowledge of ecosystem complexity in the Cambrian.

An assemblage of Cambrian Series 2, Stages 3–4, conchiferan mollusks from the Shackleton Limestone, Transantarctic Mountains, East Antarctica, is formally described and illustrated. The fauna includes one bivalve, one macromollusk, and 10 micromollusks, including the first description of the species Xinjispira simplex Zhou and Xiao, 1984 outside North China. The new fauna shows some similarity to previously described micromollusks from lower Cambrian glacial erratics from the Antarctic Peninsula. The fauna, mainly composed of steinkerns, is relatively low diversity, but the presence of diagnostic taxa, including helcionelloid Davidonia rostrata (Zhou and Xiao, 1984), bivalve Pojetaia runnegari Jell, 1980, cambroclavid Cambroclavus absonus Conway Morris in Bengtson et al., 1990, and bradoriid Spinospitella coronata Skovsted et al., 2006, as well as the botsfordiid brachiopod Schizopholis yorkensis (Ushatinskaya and Holmer in Gravestock et al., 2001), in the overlying Holyoake Formation correlates the succession to the Dailyatia odyssei Zone (Cambrian Stages 3–4) in South Australia

Brachiopods from Cambrian Series 2, Stage 4 carbonate strata of the Byrd Group in the Central Transantarctic Mountains, East Antarctica, are described for the first time. These include six lingulate, one paterinate, and one rhynchonelliform taxa, including the new lingulate brachiopod Plicarmus wildi gen. et sp. nov. The biostratigraphy correlates closely to the brachiopods recently reported from the Xinji Formation (Shuiyu section) in North China, as well as brachiopods recovered from the Dailyatia odyssei Zone across the Arrowie Basin of South Australia. These findings also support the previously identified close palaeobiogeography of these regions. The first unambiguous example of the acrotretid brachiopod Eohadrotreta zhenbaensis Li and Holmer outside South China is also identified in the context of its ontogenetic stages. Well preserved specimens of the acrotheloid Schizopholis yorkensis (Holmer and Ushatinskaya in Gravestock et al.) facilitates a new reconstruction of its musculature and visceral region. This data is synthesised into a new cladistic analysis that resolves Acrotheloidea as a well-supported monophyletic clade and supports previous hypotheses of a morphocline in acrotheloid evolution.

Cambrian Series 2 shelly fossils from thick carbonate successions in East Antarctica have received limited systematic treatment through the 20th century. Described here are the East Antarctic camenellan tommotiids from the Shackleton Limestone in the Central Transantarctic Mountains and the Schneider Hills limestone in the Argentina Range. This material comes from both newly sampled collections and incompletely described material from older collections. The assemblage supports correlation to the Dailyatia odyssei Zone and Pararaia janeae Trilobite Zone of South Australia, with the newly examined specimens of Dailyatia decobruta from the Shackleton Limestone providing direct correlation to the Mernmerna Formation of the Ikara-Flinders Ranges and White Point Conglomerate of Kangaroo Island. These East Antarctic assemblages include five species referred to Dailyatia, in addition to an undetermined kennardiid species and fragments of the problematic Shetlandia multiplicata. The results further corroborate the notion that fossiliferous carbonate clasts found on King George Island were sourced from the same carbonate shelf as the Shackleton Limestone, with the taxon S. multiplicata found in both units. The Schneider Hills limestone in the Argentina Range has yielded sclerites of Dailyatia icari sp. nov., currently only known from this location. 

The Cambrian Stage 4 upper Bastion Formation of Albert Heim Bjerge and CH Ostenfeld Nunatak, NE Greenland, yielded 34 excellently preserved sclerites of Lapworthella schodackensis among other small shelly fossils. Lapworthellids have been interpreted as members of the camenellans,a basal tommotiid group. Little is known about this group although the morphological andultrastructural features of their sclerites allow a potential reconstruction of a lophophorate body plan.The exquisite material from Greenland provides significant new data for the revision of the species taxonomy, but also for the comprehension of the scleritome structure of lapworthellids and the modeof formation of their sclerites. Two morphotypes of L. schodackensis sclerites are identified: one with asimple apex, occurring in sinistral and dextral forms; and one bilaterally symmetrical sclerite with twoapices. All bear a similar ornamentation constructed of repeated growth sets consisting of a reticulate inter-rib groove with tubercles, a densely denticulate rib and a striated sub-rib area. The new data onthe ornamentation and observations of the laminar shell microstructure of L. schodackensis enable us to improve the reconstruction of growth in lapworthellids. Finally, the morphological features of the two types of sclerites provide new information for the reconstruction of the bilaterally symmetrical multi-component lapworthellid scleritome with evidence of the fusion of adjacent sclerites duringearly ontogeny.

The Camenellan tommotiid Lapworthella bornholmiensis is systematically revised based on the original type material and collections of Small Shelly Fossils at the Swedish Museum of Natural History. The species is clearly differentiated from all other species of Lapworthella by its unique surface ornamentation. Variability in sclerite shape is analyzed among the 39 specimens and four sclerite morphotypes are recognized: bilaterally subsymmetrical A sclerites with straight apertural margin and subrectangular to subelliptical cross-section; asymmetrical B sclerites with oblique to curved apertural margin and subrectangular to subelliptical cross-section; asymmetrical C sclerites with one concave surface resulting in elliptical to sub-triangular cross section; elongated asymmetrical D sclerites with subcircular cross section. The C sclerites are further divided into two sub-types. The sclerite morphotypes of Lapworthella bornholmiensis allow a detailed comparison with other lapworthellid taxa and we identify a pattern of recurring sclerite morphotypes across a range of species, allowing a new understanding of the lapworthellid scleritome structure. The stratigraphic range of Lapworthella bornholmiensis includes the Cambrian Stage 4 Gislöv Formation, the Wuliuan-Drumian Forsemölla and Exsulans Limestones in the Alum Shale Formation and Guzhangian limestone fill in Proterozoic basement fissures in Bohuslän, making this long ranging species the youngest known tommotiid in the fossil record.

Correlation of lower Cambrian strata is often confounded by provincialism of key fauna. The widespread occurrence of themicromollusc Watsonella crosbyi Grabau, 1900 is therefore an important biostratigraphic signpost with potential for international correlation of lower Cambrian successions. Previous correlations of W. crosbyi from Australia (Normanville Group) suggested an Atdabanian- to Botoman-equivalent age. However, in the upper part of the Mount Terrible Formation, stratigraphic ranges of W. crosbyi and Aldanella sp. cf. golubevi overlap prior to the incoming of vertically burrowed ‘piperock’, which is indicative of an age no earlier than Cambrian Stage 2. The stratigraphic range of W. crosbyi in the Normanville Group, South Australia correlates with the ranges of the taxon in China, France, Mongolia and Siberia (though not Newfoundland). The new Australian data add further support for considering the first occurrence of W. crosbyi a good potential candidate for defining the base of Cambrian Stage 2. The stratigraphic range of W. crosbyi through the lower Cambrian Normanville Group has been determined based on collections from measured sections. Although rare, W. crosbyi is part of an assemblage of micromolluscs including Bemella sp., Parailsanella sp. cf. murenica and a sinistral form of Aldanella (A. sp. cf. A. golubevi). Other fauna present include Australohalkieria sp., Eremactis mawsoni, chancelloriids and Cupitheca sp.

The upper Lower Cambrian Forteau Formation in southern Labrador and the Great Northern Peninsula (GNP) is a successionof shale, limestone, siltstone and sandstone accommodated by rising sea levels during the early drift stages ofNewfoundland’s Laurentian passive margin. Its Dyeran trilobite fauna, characterized by Elliptocephala logani (Walcott, 1910)that ranges throughout the formation, indicates it was mostly deposited in the middle Bonnia‒Olenellus Zone. Its three lithostratigraphicdivisions, the Devils Cove member, Middle shale and Upper limestone, preserve a transgressive system tract(TST; Devils Cove and lower part of the Middle shale) and the early stages of a regressive high-stand system tract (HST; upper Middle shale and Upper limestone) that hosts a carbonate ramp shelf.A mudstone-dominated succession characterizes the TST, comprised of an inner belt of archeocyathid patch reef andcyclic, well-stratified, fine-grained mixed clastic and carbonate shelf rocks in southern Labrador. To the southeast on the GNP,the shale succession along with minor limestone and no reefs suggests a deeper water shelf basin in which shale accumulated across the GNP. Maximum flooding on the GNP is linked to dark shale midway through the basinal succession, and to athick shale bed that overlies the reefal strata in Labrador.Thin-bedded siltstone, sandstone and limestone in Labrador, and extensively bioturbated siltstone, sandstone and rare limestone on the GNP, support a shelf shallowing above storm-wave base as it prograded during the early stages of regression. Shallow-water carbonate of the Upper limestone supports a prograding shelf, at first dominated by an archeocyathid reefal tract and oolitic shoal complex. The reef tract and carbonate sand shoal complex prograded southeastward to justbeyond the northeast-trending Ten Mile Lake–Long Range fault system. Evidence of slumping in the underlying fine clastic sediment in the same area suggest that this fault zone may coincide with a hinge zone, beyond which the shelf steepened intomostly deep-water clastic sedimentation. The archeocyathid tract in southern Labrador is a broad biostromal complex confined within an erosional recess in the shelf. On the GNP, however, the tract is characterized by high-energy bioherms associated with crossbedded grainstone channels that can be traced for over 60 km along a northeast strike length. East of the reef tract‒shoal complex, the succession appears to be dominated by deeper water shelf mudrock, nodular carbonate and little evidence of shallow-water carbonate facies. The facies transition suggests the Forteau Formation insouthern Labrador and the GNP was laid down in a high-energy shallow-water, inner ramp setting that was up to 75 km wide. Above the carbonate sand shoal complex, the succession is marked by decametre-thick parasequences of intercalated carbonateand clastic intervals. The sequences support a shelf of fine grained to grainy carbonate deposited on an open shelframp overlain by intervals dominated by coarsening upward high-energy siliciclastics that suggest barrier complexes alongthe landward margin of the ramp. Thick units of crossbedded quartz arenite, in the upper half of the Upper limestone, suggest terrigenous sediments initially encroached along the inner part of the shelf, and eventually smothered the Forteau shelf leading to the low-stand deposits of the overlying Hawke Bay Formation. Trilobites recovered from this transition throughout the GNP, indicate that it occurred very late in the upper part of the Bonnia‒Olenellus Zone, likely between the Bristolia mohavensis Biozone and the top of the zone.

New assemblages of skeletal fossils chemically extracted from carbonates of the Cambrian Stage 2–Drumian Stage are reported from the lower reaches of the Lena River as well as from the Khorbusuonka, Malaya Kuonamka, and Bol’shaya Kuonamka rivers in northern part of the Siberian Platform. The fauna studied with scanning electron microscopy includes brachiopods, molluscs, hyoliths, halkieriids, chancelloriids, tommotiids, lobopodians, palaeoscolecidans, bradoriids, echinoderms, anabaritids, hyolithelminths, and sponges showing similarity to previously described fossil assemblages from Siberia, Laurentia, and Gondwana. The material includes emended descriptions of Halkieria proboscidea, Hadimopanella knappologica, Archaeopetasus typicus, and first descriptions of Hadimopanella foveata Kouchinsky sp. nov. and Archaeopetasus pachybasalis Kouchinsky sp. nov. Affinity of Archaeopetasus to chancelloriids is suggested. Finding of an in-place operculum in a planispiral shell of Michniakia minuta enables reinterpretation of this form as a hyolith, not a mollusc. The cambroclavids Cambroclavus sp. and Zhijinites clavus and the earliest echinoderms belonging to the Rhombifera and Ctenocystoidea are reported respectively from the lower Botoman stage and Botoman–Toyonian transitional beds, correlated with Cambrian Stage 4. Carbon isotopes are analysed from sections of the Chuskuna (upper Kessyusa Group), Erkeket, Kuonamka, Olenyok, Yunkyulyabit-Yuryakh, Tyuser and Sekten formations. A major part ofthe δ13C record is obtained from the Cambrian Stage 4–Drumian Stage strata which remain incompletely characterised by chemostratigraphy. The Lower Anomocarioides limbataeformis Carbon isotope Excursion (LACE) from the Drumian Stage of the Khorbusuonka River is introduced herein. New chemostratigraphic data are used for regional and global correlation and facilitate study of the evolutionary development of animals and faunas through the “Cambrian explosion”.

Data on the first appearances of major animal groups with mineralized skeletons on the Siberian Platform and worldwide are revised and summarized herein with references to an improved carbon isotope stratigraphy and radiometric dating in order to reconstruct the Cambrian radiation (popularly known as the ‘Cambrian explosion’) with a higher precision and provide a basis for the definition of Cambrian Stages 2 to 4. The Lophotrochozoa and, probably, Chaetognatha were first among protostomians to achieve biomineralization during the Terreneuvian Epoch, mainly the Fortunian Age. Fast evolutionary radiation within the Lophotrochozoa was followed by radiation of the sclerotized and biomineralized Ecdysozoa during Stage 3. The first mineralized skeletons of the Deuterostomia, represented by echinoderms, appeared in the middle of Cambrian Stage 3. The fossil record of sponges and cnidarians suggests that they acquired biomineralized skeletons in the late Neoproterozoic, but diversification of both definite sponges and cnidarians was in parallel to that of bilaterians. The distribution of calcium carbonate skeletal mineralogies from the upper Ediacaran to lower Cambrian reflects fluctuations in the global ocean chemistry and shows that the Cambrian radiation occurred mainly during a time of aragonite and high-magnesium calcite seas.

The tommotiid Paterimitra pyramidalis Laurie, 1986, is redescribed based on well-preserved material from the lower Cambrian Wilkawillina, Wirrapowie and Ajax limestones of the Flinders Ranges, South Australia. The material shows that the scleritome of Paterimitra pyramidalis includes three sclerite morphotypes (S1, S2 and L). Detailed shell microstructure studies show striking similarities with both the paterinid brachiopod Askepasma toddense and the tommotiid Eccentrotheca helenia, which strengthens the suggested evolutionary link between tommotiids and brachiopods. Based on the partly articulated specimens and similarities in shell microstructure and sclerite morphology with Eccentrotheca, Paterimitra pyramidalis is reconstructed as a tube-dwelling, epifaunal, sessile, filter-feeder with an organic pedicle-like attachment structure. The proposed reconstruction of the scleritome comprises a basal unit composed of one S1 and one S2 sclerite, as well as an unresolved number of L sclerites lining a coniform tubular structure.

A minute limpet-shaped shell, Qingjianglepas elegans n. gen. n. sp., is reported from the 518 million-year-old Qingjiang biota in Hubei, China. The new shell exhibits a typical limpet morphology with a shell sculpture of strongly developed radial ribs, an anterior shell fold, and distinctive shell shape shift. Feature combinations such as in Qingjianglepas are unique among Cambrian mollusk assemblages, but strikingly resemble that of some modern fissurellids (keyhole and slit limpets; Vetigastropoda). The biological affinity of Qingjianglepas is discussed, but temporarily remains uncertain. Regardless, the heavily sculptured and limpet-shaped Qingjianglepas represents a previously unrecognized shell form, and hence significantly extends the morphological diversity of Cambrian mollusks. The abrupt ontogenetic change in shell morphology suggests that Qingjianglepas might change microhabitat throughout the growth of the animal. This study reveals that the phenomenon of limpetization occurred within early Cambrian total-group conchiferans. Furthermore, Qingjianglepas might represent an evolutionary dead-end in course of helcionelloid mollusk evolution, which marks the end of the Cambrian explosion of animals at the beginning of Cambrian Series 2.

Hyoliths (hyolithids and orthothecids) were one of the most successful early biomineralizing lophotrochozoans, and were a key component of the Cambrian evolutionary fauna. However, the morphology, skeletogenesis and anatomy of earliest members of this enigmatic clade, as well as its relationship with other lophotrochozoan phyla remain highly contentious. Here we present a new orthothecid, Longxiantheca mira gen. et sp. nov. preserved as part of the secondarily phosphatized Small Shelly Fossil assemblage from the lower Cambrian Xinji Formation of North China. Longxiantheca mira retains some ancestral traits of the clade with an undifferentiated disc-shaped operculum and a simple conical conch with a two-layered microstructure of aragonitic fibrous bundles. The operculum interior exhibits impressions of soft tissues, including muscle attachment scars, mantle epithelial cells and a central kidney-shaped platform in association with its feeding organ. Our study reveals that the muscular system and tentaculate feeding apparatus in orthothecids appear to be similar to that in hyolithids, suggesting a consistent anatomical configuration among the total group of hyoliths. The new finding of shell secreting cells demonstrates a mantle regulating mode of growth for the operculum. Taking all these data into considerations, especially on the basis of shell microstructures, we argue that hyoliths were an extinct sister group of molluscs.

Aragonitic crossed-lamellar (CL) is one of themost commonly formed and extensively studied molluscanshell microstructures, yet its origin and early evolution within the Mollusca remains poorly understood. Here, a primitive CL microstructure from one of the oldest gastropods,Pela-giella  madianensis, and the problematic hyolith Cupitheca sp. of the Cambrian Series 2 Xinji Formation on the North China Platform, was investigated. In P. madianensis, detailed characterization has revealed a typical four-ordered hierarchical orga-nization of aragonitic crystallites, and a thick layer of organic membranes surrounding its first-order lamellae. A transitional fibrous microstructure was observed between the outer CL and inner foliated aragonite structural layers. In Cupitheca sp.,only the first and second-order lamellae were visible due to preservation limitations, and the first-order lamellae were extremely irregular in shape and size, which is consistent with modern representatives. This study demonstrates that the capability to construct highly-mineralized intricate shells was acquired in early Cambrian stem-group gastropods. The CL microstructure first emerged in the early Cambrian and as a basal synapomorphic trait in total-group molluscs. Moreover, presence of the CL microstructure in problematic lophotrochozoans (i.e.hyoliths) is confirmed. This study contributes to a more complete picture of the evolutionary origin andarchitectural diversity of biomineralized mollusc shells during the Cambrian explosion, and strengthens the phylogenetic links between hyoliths and molluscs.

The initial acquisition of calcium carbonate polymorphs (aragonite and calcite) at the onset of skeletal biomineralization by disparate metazoans across the Ediacaran-Cambrian transition is thought to be directly influenced by Earth’s seawater chemistry. It has been presumed that animal clades that first acquired mineralized skeletons during the so-called “aragonite sea” of the latest Ediacaran and earliest Cambrian (Terreneuvian) possessed aragonite or high-Mg calcite skeletons, while clades that arose in the subsequent “calcite sea” of Cambrian Series 2 acquired low-Mg calcite skeletons. Here, contrary to previous expectations, we document shells of one of the earliest helcionelloid molluscs from the basal Cambrian of southwestern Mongolia that are composed entirely of low-Mg calcite and formed during the Terreneuvian aragonite sea. The extraordinarily well-preserved Postacanthellashells have a simple prismatic microstructure identical to that of their modern low-Mg calcite molluscan relatives. High-resolution scanning electron microscope observations show that calcitic crystallites were originally encased within an intra- and interprismatic organic matrix scaffold preserved by aggregates of apatite during early diagenesis. This indicates that not all molluscan taxa during the early Cambrian produced aragonitic shells, weakening the direct link between carbonate skeletal mineralogy and ambient seawater chemistry during the early evolution of the phylum. Rather, our study suggests that skeletal mineralogy in Postacanthella was biologically controlled, possibly exerted by the associated prismatic organic matrix. The presence of calcite or aragonite mineralogy in different early Cambrian molluscan taxa indicates that the construction of calcium carbonate polymorphs at the time when skeletons first emerged may have been species dependent

A diverse group of molluscs from the Cambrian Series 2, Stages 3–4 Xinji Formation of the North China Block (NCB) is described, based on more than 4500 specimens from three well-studied sections in Shaanxi and Henan provinces, along the southern and southwestern margin of the NCB. Twenty molluscan species are identified, including one bivalve, three stem group gastropods, and 16 additional helcionelloids. Among these, six helcionelloid species are reported from the NCB for the first time, and one new species, Parailsanella luonanensis sp. nov. is proposed. This diverse molluscan fauna shares a large number of species with contemporaneous faunas of South Australia (15), Antarctica (7), Laurentia (6), Siberia (3) and South China (1). Faunal similarities are even greater on a generic level. The striking similarities of the molluscan faunas of North China, South Australia and Antarctica strongly support the hypothesis that the NCB was situated close to Eastern Gondwana, most likely close to South Australia in the Cambrian Epoch 2. In addition, well-preserved shell attachment muscle scars were observed in the helcionelloid Figurina figurina, with two pairs of symmetrical, continuous, band-like muscle scars, which are obviously different from the musculature of both gastropods and monoplacophorans. Because of this unique musculature, these characteristic Cambrian cap-like molluscs are assigned to the Helcionelloida rather than to the Gastropoda or Monoplacophora. A diverse group of molluscs from the Cambrian Series 2, Stages 3–4 Xinji Formation of the North China Block (NCB) is described, based on more than 4500 specimens from three well-studied sections in Shaanxi and Henan provinces, along the southern and southwestern margin of the NCB. Twenty molluscan species are identified, including one bivalve, three stem group gastropods, and 16 additional helcionelloids. Among these, six helcionelloid species are reported from the NCB for the first time, and one new species, Parailsanella luonanensis sp. nov. is proposed. This diverse molluscan fauna shares a large number of species with contemporaneous faunas of South Australia (15), Antarctica (7), Laurentia (6), Siberia (3) and South China (1). Faunal similarities are even greater on a generic level. The striking similarities of the molluscan faunas of North China, South Australia and Antarctica strongly support the hypothesis that the NCB was situated close to Eastern Gondwana, most likely close to South Australia in the Cambrian Epoch 2. In addition, well-preserved shell attachment muscle scars were observed in the helcionelloid Figurina figurina, with two pairs of symmetrical, continuous, band-like muscle scars, which are obviously different from the musculature of both gastropods and monoplacophorans. Because of this unique musculature, these characteristic Cambrian cap-like molluscs are assigned to the Helcionelloida rather than to the Gastropoda or Monoplacophora. 

Hyoliths were among the earliest biomineralizingmetazoans in Palaeozoic marine environments. They havebeen known for two centuries and widely assigned tolophotrochozoans. However, their origin and relationshipswith modern lophotrochozoan clades have been a longstand-ing palaeontological controversy. Here, we provide broadmicrostructural data from hyolith conchs and opercula fromthe lower Cambrian Xinji Formation of North China, includ-ing two hyolithid genera and four orthothecid genera as wellas unidentified opercula. Results show that most hyolithconchs contain a distinct aragonitic lamellar layer that is com-posed of foliated aragonite, except in the orthothecid Newtaxon 1 that has a crossed foliated lamellar microstructure.Opercula are mostly composed of foliated aragonite andoccasionally foliated calcite. These blade or lath-likemicrostructural fabrics coincide well with biomineralizationof Cambrian molluscs rather than lophophorates, as exempli-fied by the Cambrian members of the tommotiid-brachiopodlinage. Accordingly, we propose that hyoliths and molluscsmight have inherited their biomineralized skeletons from anon-mineralized or weakly mineralized common ancestorrather than as a result of convergence. Consequently, fromthe view of biomineralization, the homologous shellmicrostructures in Cambrian hyoliths and molluscs stronglystrengthen the phylogenetic links between the two groups.

Although various types of shell microstructures are documented from Cambrian molluscs, the precise organization and mineralogical composition of Terreneuvian molluscs are relatively unknown. Anabarella was one of the first helcionellid molluscs to appear in the Terreneuvian, with the genus surviving until the third epoch of the Cambrian. Here, shell microstructures of Anabarella australis have been studied based on new collections from the lower Cambrian (Series 2) Xinji Formation of the North China Block. Results show that A. australis has a laminar inner shell layer that consists of crossed foliated lamellar microstructure (CFL). Nacreous, crossed-lamellar and foliated aragonite microstructures previously documented in the older (Terreneuvian) species A. plana are here revised as preservational artefacts of the CFL layers. This complex skeletal organization of Anabarella suggests that mechanisms of molluscan biomineralization evolved very rapidly. Morphologically, specimens from the Chaijiawa section show a pattern of distinct ‘pseudo-dimorphism’ as external coatings are identical to Anabarella, while associated internal moulds are similar to the helcionelloid genus Planutenia. In contrast, internal moulds from the Shangzhangwan section show considerable morphological variation owing to preservational bias and show greater similarities to specimens from South Australia, Northeast Greenland and Germany. These observations demonstrate that the extensive morphological variation seen in the internal moulds of the cosmopolitan genus Anabarella are primarily preservational artefacts and are unlikely to represent the real intra- and interspecific variability of the animal. In these cases, Planutenia is here confirmed to be a subjective synonym of Anabarella.

The origin of the Brachiopoda has long been a hotly debated topic, and various models have been proposed following the latest finds of exceptionally preserved material. The lower Cambrian (Stage 3) Heliomedusa orienta from the Chengjiang Konservat‐Lagerstätte, eastern Yunnan of South China, is an important example of exceptional preservation. A wide variety of affinities have been proposed for Heliomedusa, but recently it has been suggested to reside within the mickwitziids, which may form a stem group to the Brachiopoda. Detailed studies of exceptionally preserved Heliomedusa have increased our knowledge of the soft‐part anatomy of this important early brachiopod, but unfortunately, almost nothing is known about its shell structure. Here, we describe new exceptionally preserved specimens from the Chengjiang biota to better reveal both shell structure and ornamentation. Its reticulate–pustulose ornament and tubular structure are reminiscent of traits seen in other mickwitziid brachiopods. In addition, two types of setae can be observed. Apart from the pyritized marginal mantle setae, some tubules are filled with iron oxides, potentially representing thinner and shorter penetrative setae. Both valves of H. orienta appear to have been less mineralized as compared to Mickwitzia monilifera, and the two species differ in diameter and density of tubules and pustules, and in terms of slightly less projected profile of ventral valve with lower umbo posteromedially placed. Although Heliomedusa clearly is closely related to Mickwitzia, their different preservational modes (compacted poorly mineralized/noncompacted mineralized) make detailed comparison difficult; they are provisionally kept as separate genera pending further studies of better‐preserved Chinese material.

New hyolith specimens from the early Cambrian (Stage 4) of the Three Gorges area, western Hubei Province are described and assigned to the species Doliutheca orientalis. Doliutheca are preserved in two taphonomic modes: casts in silty mudstone revealing gross morphology and some soft parts, and internal molds in calcareous pelites, which exhibit new morphological details of the conch and operculum. SEM and Micro-CT analyses show that Doliutheca preserve well-developed platy clavicles and cardinal processes on the interior of the operculum composed of rod-shaped tubular elements. This observation and the distinct cardinal and conical shields of the operculum indicate that Doliutheca could be placed within the Family Paramicrocornidae, most recently established as a group of hyoliths closely related to hyolithids.

Hyoliths, known globally from the Cambrian to the Permian are generally recognized as early lophotrochozoan animals of uncertain phylogenetic position. The calcareous exoskeleton of hyolithids, one of the two subgroups of hyoliths, consist of four elements including a conical conch, an external operculum, and a pair of curved enigmatic helens, but their ecology and internal morphology have long been considered controversial. Here we present for the first time, remarkably preserved hyolithids with helens and interior soft tissues from the Guanshan Biota (Cambrian, Stage 4) of Shijiangjun section, Wuding County in eastern Yunnan Province of South China. Preserved soft tissues and organs encompass the digestive tract, muscle scars and connective tissues between the operculum and conch. Three modes of soft part preservation in hyolithids from the Guanshan Biota are described: (1) preservation through pyritization, (2) sediment-infilling of guts, and (3) bacterial biofilm pseudomorphs (expressed as large internal hollow cavities), resulting from endogenous bacterial decay of soft parts. Such differential preservation of hyolith guts indicates different taphonomic modes for the soft parts of Cambrian animals in the Guanshan Biota. The newly collected material yields the oldest preserved guts in hyolithids from Konservat-biotas, and reveals new evidence for reconstructing hyolithid anatomy, such as the articulation mechanism of the opercula and the internal structures in the conch. Comparisons with younger wellpreserved hyolithid specimens indicate that the gut anatomy of hyolithids was evolutionarily conservative from the early Cambrian through to at least the Ordovician. 

The cap-shaped shells of Triplicatella are known almost exclusively from small shelly fossil assemblages with articulated specimens showing unequivocally that they represent the operculum of a hyolith. Abundant specimens of Triplicatella opimus from the fine-grained shales of the Chengjiang Lagerstätte of South China with soft-part preservation are documented herein. The soft tissues, including the feeding apparatus and complex digestive system, in T. opimus strongly suggest that Triplicatella was a deposit feeder. The digestive tract of T. opimus consists of two limbs, a spiral loop folded into a chevron-like structure and a slightly recurved to straight anal tube, which are preserved as reddish-black traces enriched in iron. The new anatomical information obtained from T. opimus in the Chengjiang Biota suggests an intermediate stage in the development of the characteristic folded gut of orthothecid hyoliths. The new anatomical information reported here shows that Triplicatella is one of the best-preserved early members of the Orthothecida and promotes our understanding of the general anatomy and evolution of the Hyolitha.

The phylogenetic position of hyoliths has long been unsettled, with recent discoveries of a tentaculate feeding apparatus (‘lophophore’) and fleshy apical extensions from the shell (‘pedicle’) suggesting a lophophorate affinity. Here, we describe the first soft parts associated with the feeding apparatus of an orthothecid hyolith, Triplicatella opimus from the Chengjiang biota of South China. The tuft-like arrangement of the tentacles of T. opimus differs from that of hyolithids, suggesting they collected food directly from the substrate. A reassessment of the feeding organ in hyolithids indicates that it does not represent a lophophore and our analysis of the apical structures associated with some orthothecids show that these represent crushed portions of the shell and are not comparable to the brachiopod pedicle. The new information suggests that hyoliths are more likely to be basal members of the lophotrochozoans rather than lophophorates closely linked with the Phylum Brachiopoda.

Background: A small hyolith, with a triangular operculum and a conical-pyramidal conch with a sharp apex, originally documented as Ambrolinevitus ventricosus, is revised based on new material from the Chengjiang biota. The operculum of ‘Ambrolinevitus’ ventricosus displays strong morphological similarities with the operculum of Paramicrocornus from the Shuijingtuo Formation (Cambrian Series 2), indicating that the species should be reassigned to Paramicrocornus.

Results: Based on the unusual morphology of Paramicrocornus, we herein propose a new family Paramicrocornidae fam. nov. A cladistic analysis of Cambrian and Ordovician hyoliths clearly delineates hyolithids as a monophyletic group which evolved from the paraphyletic orthothecids in the early Cambrian and with Paramicrocornidae as its closest relative.

Conclusions: The phylogenetic analysis, together with the distribution of hyoliths from the Cambrian to the Ordovician, reveals the presumptive evolution model of both the skeleton and soft-part anatomy of hyoliths. The Family Paramicrocornidae plays an intermediate role in hyolith evolution, representing the transitional stage in the evolution from orthothecids to hyolithids.

Graptolite reflectance was thought to be one of the most useful thermal maturity indicators for graptolite bearing sediments, however, the relationship between graptolite reflectance and vitrinite reflectance is not well established. Graptolites, especially in the Wufeng–Longmaxi Formations from the Ordovician to Silurian of South China, have been mistaken for vitrinite-like particles or solid bitumen, which results in inconsistent data on the thermal maturity. In this paper, we have employed optical microscope techniques to describe the detailed optical characteristics of graptolites and solid bitumen in the Wufeng–Longmaxi Formations. Laboratory simulation of maturation was used to determine the relationship between graptolite reflectance and vitrinite reflectance. The organic constituents in the Wufeng–Longmaxi Formations are mainly composed of graptolites and solid bitumen. Granular and non-granular graptolites were observed in the Wufeng–Longmaxi Formations, with nongranular as the most common texture. Solid bitumen can be distinguished from non-granular graptolites by its coarse surface, weaker anisotropy, and lower random reflectance. The combination of non-polarized and polarized light is very helpful to distinguish solid bitumen from graptolite. For comparison, organic material from the early Ordovician Alum Shale Formation of Sweden and Estonia was also studied. The macerals of the Alum shales are mainly composed of lamalginites, mineral-bituminous groundmasses, graptolites, and solid bitumen.The major textures of the graptolites in the Sweden and Estonia sediments are non-granular and granular, respectively.Both non-granular graptolite and vitrinite reflectances display a systematic increase with the increase of heating temperature and time. The granular graptolites in the Estonian sample were gradually changed to nongranular graptolites following laboratory simulated maturation, indicating that granular graptolites can transform into non-granular graptolites with maturation. Solid bitumen in the Wufeng–Longmaxi Formations was derived from the solid residue of kerogen and/or post-oil bitumen. The graptolite random reflectance is a better thermal maturity indicator than graptolite maximum reflectance and is more precise due to the smaller standard deviation. Several equations are proposed to determine the thermal maturity of the graptolite-bearing sediments based on graptolite random reflectance, graptolite maximum reflectance and solid bitumen random reflectance.

The Wufeng–Longmaxi Formations (Ordovician-Silurian) contain abundant graptolite-bearing shales, which have proven to be one of the most successful shale gas exploration targets in China and can provide an opportunity to study the organic petrology of graptolites. Organic petrology analysis has been conducted on these organic-rich shales in order to depict the shape of graptolite reflectance indicating surface (GRIS) and their organic maturity. Granular and non-granular graptolites have been observed in the studied sediments, and nongranular graptolites were the major graptolite species. The strong positive correlation between the maximum reflectance (Rmax) and bireflectance of graptolites indicates that the graptolites display stronger anisotropy with maturation. A linear coalification path of graptolite fragments was observed in the studied sediments. A remarkable decrease in the minimum reflectance (Rmin) values of graptolites was observed when Rmax exceeds 6%.The samples with lower thermal maturity (Rmax ≤2%) tend to be biaxial neutral, whereas higher rank samples display biaxial negative characteristics. The GRIS shape becomes more oblate with maturation. The Wufeng–Longmaxi shales from the southeastern Chongqing and Wuxi areas of the Chongqing municipality, China, are post mature and in the dry gas zone, whereas samples from the Chengkou area of the Chongqing municipality, China, are still in the oil window. Their different burial and thermal histories controlled their thermal maturity.

The thermal maturity of lower Paleozoic graptolite-bearing marine sediments, which host many hydrocarbon deposits worldwide, has long been difficult to determine due to the absence of wood-derived vitrinite particles for conventional vitrinite reflectance. In 1976, graptolite reflectance was introduced as a new indicator for organic maturity of these deposits and has been used since in many regional studies. The majority of these studies, however, were done on a limited sample set and a limited range of thermal maturity, which resulted in a number of controversial views concerning the usefulness of graptolite reflectance as an alternative paleothermal indicator and its correlation with vitrinite reflectance through various proxies. In this paper, we review previous studies and combine those analyses with new data to assess the physical and chemical characteristics of graptolite periderm with increasing thermal maturity. We conclude that graptolite random reflectance (GR_or) is a better parameter for the thermal maturity assessment than graptolite maximum reflectance (GR_omax) due to the better quality of available data. Combining published data with results of our study of both natural and heat-treated graptolites and vitrinite, we present a new correlation between GR_or and equivalent vitrinite reflectance (EqVR_o), as EqVR_o = 0.99GR_or + 0.08. Chemical composition of graptolite periderm is similar to vitrinite; graptolites are mainly kerogen Type II-III, are gas prone and have a substantial hydrocarbon potential. Lower Paleozoic graptolite-bearing organic-rich sediments are important shale gas source rocks and reservoirs globally and make a significant contribution to worldwide petroleum reserves

The overmature Precambrian to Lower Paleozoic marine shales from China contain relatively simple organic matter (OM) composition, which is dominated by in-source solid bitumen (i.e., solid bitumen in the hydrocarbon source rocks). The thermal evolution behavior of the original OM and the determination of thermal maturity in the Precambrian to Cambrian marine shales have been challenging for decades. The vitrinite-like maceral (VLM) is widely present in these marine shales, and its origin is still unknown. To address these issues, the immature Proterozoic Xiamaling shales from China and the immature Cambrian Alum shales from Sweden, and a Chinese immature Carboniferous coal were heat-treated at temperature range of 300 ◦C to 550 ◦C. The carbonized residue of the artificially matured samples was examined for their morphological and reflectance variation, and the results were compared with data on the other overmature natural shales from China and Sweden. OM components are similar in the Xiamaling and Alum immature oil shales, consisting of filamentous algae, matrix bituminite, bituminite, VLM and liptodetrinite, and rare thucholites are present in the Xiamaling shales. The algal-derived OM decomposed gradually due to hydrocarbon generation at 300–350 ◦C. OM is mainly composed of the in-source solid bitumen in the artificially heated shales after 350 ◦C, similar to the overmature Precambrian to Cambrian natural shales, and the in-source solid bitumen gradually loses its mass with increasing thermal maturity. The in-source solid bitumen is derived from the thermal cracking of the retained oil or the direct conversion of algal-derived liptinite macerals (e.g., the bituminite) or their mixture. VLM in the Xiamaling oil shales can not be observed after 350 ◦C, but VLM is still present in the Alum oil shales. It can be inferred that there is a different source of VLM in these shales, and VLM in the Xiamaling oil shales contains more volatile material. The VLM in the Xiamaling shales may be biodegradation products of liptinites under anoxic environments. The origin of VLM in the Cambrian Alum shales requires further study to be verified, although it is certain that graptolites are not its sources. The reflectance of in-source solid bitumen (SBRo) increases with heat-treated temperature in both the Xiamaling shales and the Alum shales, but at different rates, which may be due to the difference of the original kerogen composition in these shales. VLM reflectance (VLMRo) and SBRo in sections perpendicular to bedding can be used to determine the maturation level of the Precambrian-Cambrian sediments. Their relation to equivalent vitrinite reflectance (EqVRo) can be expressed by the following equations: EqVRo = 1.07 × VLMRo – 0.18, EqVRo = 0.87 × SBRo + 0.25 (in the Precambrian sediments) and EqVRo = 1.15 × SBRo + 0.01 (in the Cambrian sediments).

Nineteen ichnotaxa, together with algal and invertebrate remains, and various pseudo-traces and sedimentary structures are described from the Torneträsk Formation exposed near Lake Torneträsk, Lapland, Sweden, representing a marked increase in the diversity of biotic traces recorded from this unit. The “lower siltstone” interval of the Torneträsk Formation contains mostly simple pascichnia, fodinichnia and domichnia burrows and trails of low-energy shoreface to intertidal settings. The assemblage has very few forms characteristic of high-energy, soft-sediment, foreshore or upper shoreface environments (representative of the Skolithos ichnofacies).

Uranium-lead (U-Pb) LA-ICPMS analysis of zircon from a thin claystone layer within the “lower siltstone” interval yielded a maximum depositional age of 584 ± 13 Ma, mid-Ediacaran. Most of the zircon is represented by rounded detrital grains that yield dates between 3.3 and 1.0 Ga. Although the age of the basal sandstone-dominated interval of the Torneträsk Formation remains elusive owing to the absence of fossils, the ichnofossil suite from the overlying “lower siltstone” interval lacks deep arthropod trackways, such as Rusophycus and Cruziana, and is suggestive of a very early (Terreneuvian, possibly Fortunian) Cambrian age. The ichnofauna is otherwise similar to early Cambrian trace fossil assemblages from other parts of Baltica, regions further south in modern Europe, and from Greenland.

The eccentrothecimorph tommotiid Paterimitra pyramidalis Laurie, 1986, was previously only known from lower Cambrian rocks of the Northern Territory and South Australia. Herein, we document the first occurrence of P. pyramidalis from the Xinji Formation in the Shuiyu section at Ruicheng County, Shanxi Province, located at the southwestern margin of the North China Platform. This represents the first report of a tommotiid taxon from lower Cambrian strata of the North China Platform. All three sclerite types that characterise the scleritome of P. pyramidalis have been recovered and are described, permitting definitive identification to species level. The discovery of P. pyramidalis fromthe North China Platformnot only greatly extends the palaeogeographic range of this distinctive tommotiid taxon, but also supports planktotrophic development of larvae in Paterimitra as a stem group brachiopod. The discovery of P. pyramidalis supports a Cambrian, Epoch 2, late Age 3 to early Age 4 age for the shelly fossil fauna from the Xinji Formation and indicates a close palaeogeographic position between the North China Platform and Australian East Gondwana during the early Cambrian.

Disarticulated net-like plates of the lobopod Microdictyon had a near cosmopolitan distribution from the early to middle Cambrian, but are yet to be documented from the North China Platform. Here we report isolated plates of Microdictyon from the lower Cambrian Xinji Formation (Stage 4, Series 2) of the North China Platform, extending the paleogeographic distribution of Microdictyon in the early Cambrian. The plates of Microdictyon from the Xinji Formation are similar to those of other species established on the basis of isolated plates, but do bear some new characters, such as mushroom-shaped nodes with a single inclined platform-like apex, and an upper surface that displays radial lines. However, the plates documented here are left under open nomenclature due to an inadequate knowledge of intraspecific and ontogenetic variation and low specimen numbers. Through comparison of the node shapes of the isolated plates of different Microdictyon species, we consider that low mushroom-shaped nodes could be a primitive and conservative character of Microdictyon, while tall mushroom-shaped nodes may be derived character. Subtle differences in shape and number of node apices may also represent intraspecific or ontogenetic variation.

Three small assemblages of lower Cambrian (Cambrian Series 2, Stage 4) small shelly fossils are described from Laurentian strata astride Nares Strait. The fauna from the Humboldt Formation of Daugaard-Jensen Land, North Greenland, is derived from inner shelf sediments deposited on the stable craton of the Inglefield Land High. Fossils from Judge Daly Promontory, eastern Ellesmere Island, Nunavut, occur in strata of the Cambrian Ellesmere Group (Kane Basin Formation) that have been structurally juxtaposed against older strata; they were originally assigned to the Kennedy Channel Formation, which is now considered to be of Neoproterozoic age. A similar fauna from offshore environments of the Aftenstjernesø Formation in northern Nyeboe Land, North Greenland, reflects the regional structural and sedimentological continuity with the Canadian Cambrian succession. Pojetaia robsonae sp. nov. is described from Judge Daly Promontory. 

Tuberculate fragments referred to Nikolarites spasskyi are compared with a complete carapace and other material of Spinospitella from Cambrian Series 2, Stage 4 in North Greenland. The descriptions expand earlier brief records from Laurentia of the two bradoriids, originally described from Siberia and Australia, respectively. The mutually exclusive occurrences of the two taxa indicate ecological control of their distribution in North Greenland. Robust fragments of Nikolarites spasskyi occur in reworked higher energy deposits of the Aftenstjernesø Formation, while thin-walled carapaces of Spinospitella characterise deeper water mudstones within the Buen and Aftenstjernesø formations.

Ovatoryctocara granulata Chernysheva 1962, a key trilobite in discussions concerning intercontinental correlation and the placement of the Cambrian Series 2–Series 3 boundary, occurs together with more than 60 other species in a fossil assemblage from the Henson Gletscher Formation of Løndal, Peary Land, North Greenland. Most taxa are described from acid residues in which diverse protoconodonts, helcionellid molluscs and hyoliths occur together with linguliformean brachiopods, bradoriids and other fossils. Recognition of an earlier First Appearance Datum for O. granulata in Løndal consolidates the biostratigraphic relationship between the assemblage and other partly co-occurring trilobites valuable for correlation within Laurentia and beyond. The assemblage can be correlated with the Ovatoryctocara Biozone, the basal biozone of the Molodian Stage in the Republic of Sakha (Yakutia, north-eastern Siberia), proposed as the earliest stage of Cambrian Series 3. New taxa are Tulenicornus? frykmani sp. nov., Conotheca hensoni sp. nov., Capitoconus borealis sp. nov., ‘Hipponicharion’ pearylandica sp. nov. and Liangshanella? nivalis sp. nov.

Three specimens of a new eccentrothecimorph tommotiid are described from the Paralelldal Formation of North Greenland. The specimens are all tubular structures composed of a number of individual sclerites. The sclerites were arranged in rings which fused to form a rigid tube during ontogeny. The tube has a basal aperture presumably allowing attachment to a hard substrate. In morphology, both individual sclerites and the tubular scleritome are remarkably similar to specimens of Eccentrotheca from South Australia. However, the Greenland specimens are silicified and may have been either weakly mineralized or calcareous in original composition. In this respect they differ from all previously known tommotiids, considerably expanding the ultrastructural disparity of the Tommotiida and allowing comparison to a new range of possible lophotrochozoan fossils.

Internal moulds of complete shells, including conch and associated opercula, of the hyolith Triplicatella disdoma from Cambrian Series 2, Stages 3–4 of South Australia are described. The conch of T. disdoma is shown to be narrow and cone-shaped, and with a rounded triangular cross-section. The conch shows a gentle dorsal curvature in lateral view. The fossils lack evidence of helens, and the operculum was smaller than the apertural diameter of the conch and could be withdrawn a short distance into the conch. Triplicatella was probably closely related to orthothecid hyoliths.

The camenellan tommotiid Dailyatia is one of the most common fossils in shallow water carbonates from Cambrian Stages 2-4 in South Australia (Arrowie and Stansbury basins). Six species of Dailyatia are analysed and new terminology for describing camenellan sclerites is introduced. Dailyatia sclerites are found in three fundamental sclerite types (A-C), each of which may be present in one to three sub-types depending on species. The previously described species Dailyatia ajax Bischoff 1976 and D. macroptera (Tate 1892) are revised and four additional species are described for the first time from South Australia. These include D. odyssei Evans & Rowell 1990, previously known only from Antarctica, and two new species; D. bacata sp. nov. and D. helica sp. nov. as well as a species left under open nomenclature. Two of the recognised species (D. macroptera and D. helica) occur in two different ecophenotypic variants. Species and variants are defined by differences in sclerite types present in the scleritome, sclerite morphology and ornament. The sclerites of Dailyatia are finely laminated with distal expansion of laminae supporting the prominent concentric ribs. The external surface is covered by a fine reticulate network which indicates that the sclerites were at least partly embedded in soft integument. The pattern of incremental growth reveals specific initial and possible gerontic growth stages with unique surface sculptures. Evidence of physical damage and growth disturbances is common in Dailyatia sclerites and many specimens reveal preferential abrasion of the apex. Apical canals are present in all sclerites and are connected to specialised internal apical structures.

The internal surface of the sclerites in most species reveals raised platforms and depressed, scar like areas forming unique patterns in each sclerite type, presumably representing muscular attachment. Two specimens revealing ontogenetic fusion of Dailyatia sclerites have been recovered. Based on all available evidence, a new reconstruction of the Dailyatia scleritome is proposed. In the reconstruction, a central row of A and paired B sclerites is flanked on both sides by one or two lateral rows of C sclerites. The exact number of sclerites may vary between species. This reconstruction is based on an assumed slug-like bodyplan and the Dailyatia animal is considered to be a vagrant, benthic animal living in and around archaeocyathan-microbial buildups and in other shallow water carbonate environments.

The organophosphatic early Cambrian (Terreneuvian, Cambrian Stage 2) fossil Tumulduria incomperta has been problematic ever since its original description in 1969. Comparison of abundant specimens from the Lower Cambrian of Siberia with co-occurring brachiopod valves show that T. incomperta represents the central portion of the ventral interarea of a paterinid brachiopod similar to Cryptotreta neguertchenensis, and that the domed central portion of typical Tumulduria specimens represents the ridge-like pseudodeltidium of the interarea.

The extraordinary window of phosphatized and phosphatic small shelly fossils (SSF) during the early and middle Cambrian is an important testament to the radiation of biomineralizing metazoans. While SSF are well known from most Cambrian palaeocontinents during this time interval, western Laurentia has relatively few SSF faunas. Here we describe a diverse SSF fauna from the early Cambrian (Stages 3–4) Mural Formation at three localities in Alberta and British Columbia, Canada, complemented by carbon isotope measurements to aid in a potential future bio-chemostratigraphic framework. The fauna expands the recorded SSF assemblage diversity in western Laurentia and includes several brachiopods, four bradoriids, three chancelloriids, two hyoliths, a tommotiid and a helcionellid mollusc as well as echinoderm ossicles and specimens of Microdictyon, Volborthella and Hyolithellus. New taxa include the tommotiid genus Canadiella gen. nov., the new bradoriid species Hipponicharion perforata sp. nov. and Pseudobeyrichona taurata sp. nov. Compared with contemporaneous faunas from western Laurentia, the fauna is relatively diverse, particularly in taxa with originally phosphatic shells, which appear to be associated with archaeocyathid build-ups. This suggests that the generally low faunal diversity in western Laurentia may be at least partly a consequence of poor sampling of suitable archaeocyathan reef environments. In addition, the tommotiid Canadiella filigrana appears to be of biostratigraphical significance in Cambrian Stage 3 strata of western Laurentia, and the unexpected high diversity of bradoriid arthropods in the fauna also suggests that this group may prove useful for biostratigraphical resolution in the region. 

The operculum of the problematic tubular fossil Cupitheca holocyclata Bengtson in Bengtson et al., (1990) is described for the first time based on collections from South Australia and North China. The phosphatized sub-circular operculum exhibits well defined cardinal processes and a narrow cardinal shield unequivocally demonstrating that Cupitheca is a hyolith, probably an orthothecid. C. holocyclata has an almost global distribution in Cambrian Stages 3–4. The apical structure of the operculum is an elevated, disc-shaped platform with a concave base and a marginal rim that could represent the scar of a specialized larval attachment structure, perhaps anchoring the larval hyolith to a sediment grain, algae or other benthic substrate. Cupitheca probably had a pelagic larval stage and settled on the seafloor by attachment of the apical disc to suitable substrates before developing a free-living benthic adult lifestyle. This contrasting mode of life compared to other hyolith genera suggests that the group had already evolved a range of distinct lifestyles in the Cambrian, providing significant clues into their ecology and distribution.

A new cap-shaped mollusc, Emargimantus angulatus gen. et sp. nov. is described from the Arrowie Basin of South Australia. The new species is closely comparable to mollusc species from South China and North-East Greenland previously described under the generic name Ocruranus Liu, a genus recently reinterpreted as a multiplated, possibly polyplacophoran mollusc. Emargimantus is interpreted as a univalved helcionelloid mollusc and differs from Ocruranus in both morphology and function. E. angulatus represents the first discovery of Ocruranus-like helcionelloids in the lower Cambrian of eastern Gondwana and demonstrates that these molluscs had a global distribution during the early Cambrian.