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.

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.

Sphenothallus is a tubular organism that is one of the most widely distributed and longest-ranging genera through the Palaeozoic. Despite its apparent cosmopolitan distribution, the genus has never been reported from North China. New specimens of Sphenothallus sp. have been discovered in the upper part of the Houjiashan and base of the Mantou formations (early to middle Age 4, Epoch 2, Cambrian) in Jiangsu Province, North China. The specimens are small tubes (up to 5 mm long) and have typical Sphenothallus characteristics, such as a multilayered lamellar structure, and subcircular to elliptical transverse cross-section with a pair of longitudinal thickenings situated at the widest diameter. Our material shows that both the rate of apertural expansion and the curvature of the tubes are significantly larger in early growth stages than in the later growth stages. As the diameter of the aperture increases, the transverse cross-section of the Sphenothallus sp. tube changes from subcircular at the proximal end to elliptical or lenticular at the distal end, and its wall thickness changes from uniform to thickening longitudinally. The discovery of Sphenothallus sp. from the North China Platform represents an extension of its palaeogeographic range during the Cambrian.

New material of disarticulated paleoscolecid remains have been found in “Small Shelly/Skeletal Fossils” assemblages from Cambrian Stage 3 extracted from a section in the Khubsugul Lake region of northern Mongolia. The current material is composed of isolated phosphatic plates, rendering the whole-body reconstruction and comparisons difficult. However, the morphology of the plates is unique enough to warrant description of a new genus and species Floraconformis egiinensis. The new taxon is characterised by a stellate depression network spreading from the middle that separates numerous elevations. Floraconformis egiinensis gen. et sp. nov. represents one of the oldest records of isolated palaeoscolecid plates.

The occurrence of a series of thin but persistent early Cambrian (Cambrian Age 4) brachiopod shell concentrations extending along a transect of some 150 km in NE Greenland, indicates the ability of the group to form widespread, skeletal pavements very early in the history of the phylum, its gregarious behaviour and ability to harness available nutrients. These extensive shell pavements within the Bastion Formation mark biological events within the basin, the abundance of shells perhaps associated with oscillating redox conditions prompting the dissolution of phosphate and its spread across the shelf. The shells were subsequently reworked and deposited by episodic distal storm surges with some winnowing.

The lower Cambrian Lagerstätte of Sirius Passet, Peary Land, North Greenland, is one of the oldest of the Phanerozoic exceptionally preserved biotas. The Lagerstätte evidences the escalation of numbers of new body plans and life modes that formed the basis for a modern, functionally tiered ecosystem. The fauna is dominated by predators, infaunal, benthic and pelagic, and the presence of abundant nekton, including large sweep-net feeders, suggests an ecosystem rich in nutrients. Recent discoveries have helped reconstruct digestive systems and their contents, muscle fibres, and visual and nervous systems for a number of taxa. New collections have confirmed the complex combination of taphonomic pathways associated with the biota and its potentially substantial biodiversity. These complex animal-based communities within the Buen Formation were associated with microbial matgrounds, now preserved in black mudstones deposited below storm wave base that provide insight into the shift from late Neoproterozoic (Ediacaran) to Cambrian substrates and communities. Moreover, the encasing sediment holds important data on the palaeoenvironment and the water-column chemistry, suggesting that these animal-based communities developed in conditions with very low oxygen concentrations.

Two major, extended diversifications punctuated the evolution of marine life during the Early Palaeozoic. The interregnum, however, between the Cambrian Explosion and the Great Ordovician Biodiversification Event, is exemplified by the Furongian Gap when there was a marked drop in biodiversity. It is unclear whether the gap is apparent, due to sampling failure or lack of rock, or real — associated with unique and fluctuating environments, a distinctive palaeogeography and extreme climates during the late Cambrian. Indications suggest that there has been little attention paid to this interval compared with those below and above, while some of the classical areas for Cambrian research, such as Bohemia, have poor coverage through the Furongian. Moreover, based on information available in databases and the literature, together with the ghost ranges of many higher taxa through the Furongian, data suggest that biodiversity in this stage has been significantly underestimated. Emphasis, to date, has been placed on widespread, deeper-water dark shale facies of the interval, with generally low diversity faunas, whereas shallow-water communities have often been neglected.

The Precambrian–Cambrian interval saw the first appearance of disparate modern metazoan phyla equipped with a wide array of mineralized exo- and endo-skeletons. However, the current knowledge of this remarkable metazoan skeletonization bio-event and its environmental interactions is limited because uncertainties have persisted in determining the mineralogy, microstructure, and hierarchical complexity of these earliest animal skeletons. This study characterizes in detail a previously poorly understood fibrous microstructure—the lamello-fibrillar (LF) nacre—in early Cambrian mollusk and hyolith shells and compares it with shell microstructures in modern counterparts (coleoid cuttlebones and serpulid tubes). This comparative study highlights key differences in the LF nacre amongst different lophotrochozoan groups in terms of mineralogical compositions and architectural organization of crystals. The results demonstrate that the LF nacre is a microstructural motif confined to the Mollusca. This study demonstrates that similar fibrous microstructure in Cambrian mollusks and hyoliths actually represent a primitive type of prismatic microstructure constituted of calcitic prisms. Revision of these fibrous microstructures in Cambrian fossils demonstrates that calcitic shells are prevalent in the so-called aragonite sea of the earliest Cambrian. This has important implications for understanding the relationship between seawater chemistry and skeletal mineralogy at the time when skeletons were first acquired by early lophotrochozoan biomineralizers.

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

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 Postacanthella shells 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.

Morphology usually serves as an effective proxy for functional ecology,1,2,3,4,5 and evaluating morphological, anatomical, and ecological changes permits a deeper understanding of the nature of diversification and macroevolution.5,6,7,8,9,10,11,12 Lingulid (order Lingulida) brachiopods are both diverse and abundant during the early Palaeozoic but decrease in diversity over time, with only a few genera of linguloids and discinoids present in modern marine ecosystems, resulting in them frequently being referred to as “living fossils.”13,14,15 The dynamics that drove this decline remain uncertain, and it has not been determined if there is an associated decline in morphological and ecological diversity. Here, we apply geometric morphometrics to reconstruct global morphospace occupation for lingulid brachiopods through the Phanerozoic, with results showing that maximum morphospace occupation was reached by the Early Ordovician. At this time of peak diversity, linguloids with a sub-rectangular shell shape already possessed several evolutionary features, such as the rearrangement of mantle canals and reduction of the pseudointerarea, common to all modern infaunal forms. The end Ordovician mass extinction has a differential effect on linguloids, disproportionally wiping out those forms with a rounded shell shape, while forms with sub-rectangular shells survived both the end Ordovician and the Permian-Triassic mass extinctions, leaving a fauna predominantly composed of infaunal forms. For discinoids, both morphospace occupation and epibenthic life strategies remain consistent through the Phanerozoic. Morphospace occupation over time, when considered using anatomical and ecological analyses, suggests that the limited morphological and ecological diversity of modern lingulid brachiopods reflects evolutionary contingency rather than deterministic processes.

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.

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.

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.

New fossil discoveries are reported from the Grammajukku Formation at Luobákte south of Lake Torneträsk in northern Swedish Lapland, including a fauna of Small Shelly Fossils (SSF) from a limestone bed in the uppermost part of the formation and new occurrences of brachiopods and trilobites in siltstones of the lower part of the formation. The moderately diverse SSF fauna is the first of its kind reported from the Swedish Caledonides and includes the first record of the tommotiid Lapworthella schodackensis and the bradoriid spine Mongolitubulus spinosus from Baltica, together with fragmentary specimens of Bradoria sp. and remains of one additional bradoriid arthropod, a protoconodont and a helcionelloid mollusc. In addition, the limestone bed yields abundant specimens of the brachiopods Botsfordia cf. caelata and Eoobolus cf. priscus and an unidentified ellipsocephalid trilobite. Lower down in the Grammajukku Formation, specimens of both brachiopod taxa, orthothecid hyoliths, the trilobite Ellipsocephalus cf. gripi and an unidentified holmiid trilobite were found at several levels in a siltstone, previously regarded as unfossiliferous. These discoveries markedly increase the known diversity of the palaeobiota from the Grammajukku Formation in northern Lapland and provide new insights into the biostratigraphy and palaeoenvironment of the lower Cambrian in Scandinavia and the palaeobiogeography of Cambrian faunas in general.

The diversification of animals during the Cambrian Period is one of the most significant evolutionary events inEarth’s history. However, the sequence of events leading to the origin of ‘modern’ ecosystems and the exacttemporal relationship between Ediacaran and Cambrian faunas are uncertain, as identification of the Ediacaran–Cambrian boundary and global correlation through this interval remains problematic. Here we review thecontroversies surrounding global correlation of the base of the Cambrian and present new high-resolutionbiostratigraphic, lithostratigraphic and δ13C chemostratigraphic data for terminal Ediacaran to basal Cambrianstrata in the Zavkhan Basin of Mongolia. This predominantly carbonate sequence, through the Zuun-Arts andBayangol formations in southwestern Mongolia, captures a distinct, negative δ13C excursion close to the top ofthe Zuun-Arts Formation recognized as the BAsal Cambrian carbon isotope Excursion (BACE). In this location,the nadir of the BACE closely coincides with first occurrence of the characteristic early Cambrian protoconodontProtohertzina anabarica. Despite recent suggestions that there is an evolutionary continuum of biomineralizinganimals across the Ediacaran–Cambrian transition, we suggest that this continuum is restricted to tubular forms,and that skeletal taxa such as Protohertzina depict ‘true’ Cambrian representatives that are restricted entirely tothe Cambrian. Employing the first appearance of the trace fossil Treptichnus pedum to define the base of theCambrian suffers significant drawbacks, particularly in carbonate settings where it is not commonly preserved.As T. pedum is the only proxy available to correlate the Cambrian Global boundary Stratotype Section and Point(GSSP) defined at Fortune Head, Newfoundland, we suggest that the GSSP be redefined elsewhere, in a newstratigraphic section that contains secondary markers that permit global correlation. We propose the nadir of theBACE as the favored candidate to define the base of the Cambrian. However, it is essential that the BACE becomplemented with secondary markers. In many global sections the nadir of the BACE and the first occurrence ofthe genus Protohertzina are closely juxtaposed, as are the BACE and T. pedum. Hence these taxa provide essentialbiostratigraphic control on the BACE and increase potential for effective global correlation. We also recommendthat an Auxiliary boundary Stratotype Section and Point (ASSP) be simultaneously established in order toincorporate additional markers that will aid global correlation of the Ediacaran–Cambrian boundary. The BAY4/5 section through the upper Zuun-Arts and Bayangol formations yields key shelly fossils and δ13C values and istherefore an ideal candidate for consideration as the GSSP for the Ediacaran–Cambrian boundary.

Recently we documented a bilaterally symmetrical, solitary organism, Yanjiahella biscarpa from the early Cambrian (Fortunian) of China¹. We interpreted that Y. biscarpa possessed an echinoderm-like plated theca, a muscular stalk similar to hemichordates and a pair of long, feeding appendages. Our interpretation and our phylogenetic analysis suggest that Y. biscarpa is a stem-echinoderm, which would confirm that echinoderms acquired plates before pentaradial symmetry and that their history is firmly rooted in bilateral forms. Zamora et al.² however, have criticized our interpretation, arguing against an echinoderm affinity, instead suggesting that the phylogenetic placement of Y. biscarpa is dubious and its significance for understanding deuterostome evolution is uncertain.

This criticism² seems to stem from our interpretation of particular morphological features in Y. biscarpa¹ and the perceived lack of echinoderm synapomorphies. Echinoderms possess a calcitic skeleton with a distinctive three-dimensional mesh-like microstructure called stereom, that is considered a major synapomorphy of the Echinodermata³. Zamora et al.² highlighted the absence of stereom in Y. biscarpa, additionally stating that we had omitted appropriate methods, specifically latex casting, that may confirm the presence of stereom in our specimens. We concede that initially we did not latex cast any specimens of Y. biscarpa, predominantly due to the fragile nature and the associated risk of damaging the specimens in question. In lieu of latex casting we employed Scanning Electron Microscopy (SEM) to investigate the surface and the details of Y. biscarpa specimens. SEM has been extensively used in the past to study stereom microstructure^4,5,6 and if such a microstructure was preserved in Y. biscarpa it would have been detected using this technique.

The Ediacaran–Cambrian transition is characterized by an unprecedented change in biosphere, lithosphere, and atmosphere. However, the identification of this transition and its global correlation remains debated. East Yunnan is a region of utmost importance in the study of the Ediacaran-Cambrian transition. Although strata from this region have been well studied, recognition of the base of the Cambrian continues to be controversial. This paper presents, a carbon chemostratigraphic isotope and trace element profile through the Dengying and Zhujiaqing formations in the Sujiawa section in East Yunnan. Through carbon isotope and trace element analyses of the Dengying and Zhujiaqing formations we attempt to regionally correlate the section and discuss the paleo-marine redox environment changes during this period. The Fe content of samples is low and the Mn/Sr ratios of the majority of the Sujiawa section samples is lower than 10 and shows no correlations with δ13Ccarb and δ18Ocarb values, indicating that the samples retain near-primary δ13Ccarb characteristics. The carbon isotope profile from the Sujiawa section, shows a prominent negative δ13Ccarb in the basal Zhongyicun Member (N1’, −7.3‰) that most likely represents the prominent negative excursion recorded in Ediacaran-Cambrian strata across the world. REE patterns and Y/Ho ratios were used to screen samples, to ensure that the primary seawater REE features and Ce anomalies were preserved. The samples satisfying this screening process show seawater-like REE distribution patterns in leached carbonates and may have recorded Ce anomalies of the seawater from which the carbonates precipitated. These data show that the Baiyanshao Member has Ce/Ce* values between 0.57 and 0.88, the Daibu Member between 0.56 and 0.83, and the Zhongyicun Member between 0.60 and 0.96. The Ce anomalies through the section indicates that the region experienced suboxic-oxic-anoxic conditions throughout the Ediacaran-Cambrian period. Carbon isotope and trace element analyses indicate that the strata record a negative carbon isotope excursion event and anoxic conditions during the period of deposition. This study complements the carbon chemostratigraphic information and trace element data in the region and provides valuable clues for understanding the lithofacies changes through the Ediacaran and Cambrian in different regions of South China.

Parasite–host systems are pervasive in nature but are extremely difficult to convincingly identify in the fossil record. Here we report quantitative evidence of parasitism in the form of a unique, enduring life association between tube-dwelling organisms encrusted to densely clustered shells of a monospecific organophosphatic brachiopod assemblage from the lower Cambrian (Stage 4) of South China. Brachiopods with encrusting tubes have decreased biomass (indicating reduced fitness) compared to individuals without tubes. The encrusting tubes orient tightly in vectors matching the laminar feeding currents of the host, suggesting kleptoparasitism. With no convincing parasite–host interactions known from the Ediacaran, this widespread sessile association reveals intimate parasite–host animal systems arose in early Cambrian benthic communities and their emergence may have played a key role in driving the evolutionary and ecological innovations associated with the Cambrian radiation.

Acrotretides are extinct micromorphic brachiopods that exhibited considerable morphological variation during their rapid evolution in the early Palaeozoic. The plano-conical shells of acrotretides are distinct in comparison to other brachiopod groups and despite their diversity and abundance in early Palaeozoic communities, their origins, early evolution, life history and phylogeny are poorly understood. Here, we employ advanced geometric morphometrics to quantitatively investigate ontogenetic variation and allometry in the ventral valve of the oldest known acrotretide species from the early Cambrian of South China. Our results identify substantial shape variation for Eohadrotreta zhenbaensis, along with a parabolic morphological trajectory through ontogeny, demonstrating a remarkable reversal to PC1 values equivalent to those obtained for juveniles, during later ontogenetic stages. The evolutionary novel body plan (diminutive and plano-conical) of Acrotretida was established gradually during two phases of allometry, formed initially during the final stage of the Cambrian evolutionary radiation from an ancestral low, equivalved lingulide body plan. The development of a conical shaped valve seems to have resulted in an overall smaller body size, when compared with non-conical forms. The heterochronic processes responsible for generating these ontogenetic modifications at different allometric phases may have facilitated the evolutionary diversification of acrotretide brachiopods during the early Palaeozoic.

Biologically-controlled mineralization producing organic-inorganic composites (hard skeletons) by metazoan biomineralizers has been an evolutionary innovation since the earliest Cambrian. Among them, linguliform brachiopods are one of the key invertebrates that secrete calcium phosphate minerals to build their skeletons. One of the most distinct shell structures is the organo-phosphatic cylindrical column exclusive to phosphatic-shelled brachiopods, including both crown and stem groups. However, the complexity, diversity and biomineralization processes of these microscopic columns are far from clear in brachiopod ancestors. Here, exquisitely well-preserved columnar shell ultrastructures are reported for the first time in the earliest eoobolids. The hierarchical shell architectures, epithelial cell moulds, and the shape and size of cylindrical columns are scrutinised in Latusobolus xiaoyangbaensis gen. et sp. nov. and Eoobolus acutulus sp. nov from the Cambrian Series 2 Shuijingtuo Formation of South China. The secretion and construction of the stacked sandwich model of columnar architecture, which played a significant role in the evolution of linguliforms, is highly biologically controlled and organic-matrix mediated. Furthermore, a continuous transformation of anatomic features resulting from the growth of columnar shells is revealed between Eoobolidae, Lingulellotretidae and Acrotretida, shedding new light on the evolutionary growth and adaptive innovation of biomineralized columnar architecture among early phosphatic-shelled brachiopods during the Cambrian explosion.