Integrated biostratigraphy and carbon isotopes from the Cambrian Tangwangzhai section,North China

The Tangwangzhai section, western Shandong Province, North China, the type section for the Cambrian Kushan and Chaomitian formations, yielded a diverse and relatively well-preserved conodont fauna, in which we recognize the Westergaardodina orygma, Westergaardodina matsushitai, Muellerodus? erectus, and Westergaardodina aff. fossa–Prooneotodus rotundatus zones of the North China conodont zonation. The Tangwangzhai conodont succession can be correlated not only with the polymerid trilobites occurring in the section but also with the conodont zones established for South China. The first occurrence of Furnishina longibasis and Furnishina quadrata in the upper part of the Westergaardodina matsushitai Zone allows the recognition of the base of the Paibian Stage and Furongian Series in the upper part of the Kushan Formation. The base of the Jiangshanian Stage, in the uppermost Muellerodus? erectus Zone, can be recognized by the presence of Westergaardodina cf. calix close to the base of the Chaomitian Formation. Chemostratigraphic analyses of the Tangwangzhai section show the onset of a positive carbon isotope excursion, referred to the SPICE event, in the upper part of the Kushan Formation at a level corresponding to the first occurrence of F. longibasis and F. quadrata. The base of the Jiangshanian Stage in the section is close to the demise of the SPICE positive excursion.     

Ontogeny of the eodiscoid trilobite Tsunyidiscus acutus from the Lower Cambrian of South China

Abstract: The morphology and ontogeny of the eodiscid trilobite Tsunyidiscus acutus Sun is described on the basis of numerous calcified specimens collected from the Lower Cambrian Shuijingtuo Formation in Yichang and Changyang, Hubei Province, South China. An ontogenetic series is established based on the articulated material including the previously unknown protaspides and meraspides (degrees 0 and 1). The material, revealing some prominent morphological changes such as the number of pygidial axial rings, genal spines retained throughout ontogeny and pygidial pleurae from furrowed to unfurrowed, enables a discussion on the trunk segmentation schedule, indicating that somitogenesis and tagmosis occurred independently during the ontogenetic development of T. acutus.     

Morphology and ontogeny of the eodiscoid trilobite Sinodiscus changyangensis from the lower Cambrian of South China

Abstract: A large number of complete specimens together with numerous disarticulated sclerites of the eodiscinid trilobite Sinodiscus changyangensis Zhang in Zhou et al., 1977 have been collected from the lower Cambrian Shuijingtuo Formation in Changyang, Hubei Province, South China. An ontogenetic series is established based on the immature and mature exoskeletons including the previously unknown protaspides and meraspides, in particular. No further substages can be differentiated in the protaspid specimens herein. Changes that took place during the meraspid period include the addition of postcephalic segments and prominent pygidial larval notches in early meraspid development which became progressively less distinct and disappeared in degree 2. Two holaspid stages are recognized based on the addition of a new pygidial segment, indicating that the start of the holaspid phase preceded the onset of the epimorphic phase and accordingly, its developmental mode is attributed to the protarthrous pattern. The trunk segmentation schedule of S. changyangensis is discussed, which is similar to other primitive eodiscoid trilobites, that is, as the boundary between the thorax and pygidium migrated posteriorly, there is no change in the number of the trunk segments. The processes of liberation of the thoracic segment and segment insertion into the pygidium are separated from one another, and the two different mechanisms, somitogenesis and tagmosis, progress independently during the ontogenetic development of the postcephalic region of these primitive eodiscinids.     

Evidence for the development of local anoxia during the Cambrian SPICE event in eastern North America

The later Cambrian Steptoean Positive Carbon Isotope Excursion (SPICE) event was an episode marked by pronounced changes to the global biogeochemical cycles of carbon and sulfur and significant extinctions on several paleocontinents including Laurentia (North America). While the exact cause(s) of these events remains debated, various lines of evidence suggest an increase in the areal extent of anoxia at the seafloor was a likely feature. Here, we explore whether changes in local oxygenation accompanied the onset of the SPICE in southern Laurentia using cores of the Nolichucky and Eau Claire Formations from Ohio and Kentucky, USA, that represent a transect into the Rome Trough/Conasauga intrashelf basin. At our study locations, the initial positive δ¹³C shift of the SPICE occurs in conjunction with increases in the abundance and δ³⁴S of sedimentary pyrite. Further local redox conditions, tracked using iron speciation analysis, indicate anoxic conditions developed at the two proximal locations after the start of the paired isotopic excursions. However, the location near the basin center shows no indication for anoxia before or during the onset of the SPICE. While this signal may reflect the structure of local redox conditions within the basin, with the development of anoxia limited to the basin margins, we argue that authigenic iron enrichments were muted by sedimentary dilution and/or the enhanced authigenesis of iron‐bearing sheet silicates near the basin center, masking the signal for anoxia there. Regardless of the areal extent of anoxia within the basin, in either scenario the timing of the development of anoxic bottom waters was concurrent with local faunal turnover, features broadly consistent with a global expansion of anoxia playing a role in driving the isotopic trends and extinctions observed during the event.     

A new oryctocephalid trilobite from the Balang Formation (Cambrian Stage 4) of northwestern Hunan,South China,with remarks on the classification of oryctocephalids

A new trilobite, Longaspis paiwuensis n. gen. n. sp., from the Balang Formation (Cambrian Stage 4) in northwestern Hunan, South China, is described. This rare trilobite adds to an expanding taxonomic list of organisms recognized from the Balang Lagerstätte, a deposit of exceptional preservation from the Cambrian. Longaspis paiwuensis is an unusually large-sized, micropygous oryctocephalid trilobite that has proparian facial sutures, pit-like lateral glabellar furrows, up to 17 thoracic segments, and a distinct medial notch in the pygidium; it lacks marginal spines.The classification of the family Oryctocephalidae is reviewed. Three subfamilies are recognized, and Longaspis n. gen. is assigned to the subfamily Oryctocarinae.     

Gut content fossilization and evidence for detritus feeding habits in an enrolled trilobite from the Cambrian of China

Enrolment was a major defensive strategy for trilobites that significantly contributed to their evolutionary success. The ability to enrol also helped to constrain the morphological evolution of trilobites, which in part was driven by the need to improve this capability to encapsulate the soft parts of the body within the mineralized dorsal exoskeleton. Here, we describe a unique example of gut content fossilization in an enrolled trilobite from the Cambrian of China, and we propose a taphonomic scenario that considers the possible implication of enrolment in this exceptional preservation. A micro‐facies analysis indicates that the specimen was entombed during an obrution event and remained intact due to limited infaunal activity. The encapsulation of the body did not prevent the decay of soft tissues, but it permitted the delicate gut content to be protected during diagenesis. In isolating the decaying soft tissues from the external environment, enrolment might also have favoured the establishment of microenvironmental conditions conducive to the precipitation of pyrite framboids. Within the gut, the formation of such crystals may have led to the consolidation of the ingested material. These results suggest that fossilized gut contents might be quite common in enrolled trilobites. Textural and compositional analyses reveal that the gut content is similar to the sediment surrounding the fossil except for the presence of pyrite framboids that indicate higher initial organic matter content. The complete enrolment of the body argues against an accidental ingestion of this material or a diagenetic origin for it. Accordingly, detritus feeding habits are inferred for this ptychopariid trilobite.     

Unique lowermost Cambrian small shelly fossils (SSFs) from eastern Yunnan,South China: implications for the early diversification history of animals

With a particular focus on the earliest Cambrian diversification of small shelly fossils (SSFs), stratigraphic analysis was conducted on the lower Cambrian Zhongyicun Member at the Hongjiachong section in the Chengjiang area, Yunnan, China. From ca. 3-m-thick bedded phosphorites (Unit A) in the lower part of the member, we recovered unique SSFs. This interval with 4 SSF-bearing horizons is characterized by the dominance in blade-shaped SSFs, including Halkieria spp., Brushenodus prionodes, Sinosachites delicatus, and Pteronus sp., and the absence of typical tube/cap-shape SSFs common in the Fortunian. This interval tentatively named “Halkieria-dominant interval” is stratigraphically positioned ca. 3 m below the previously known level of the oldest mollusks in the continuous bedded phosphorite sequence between the Anabarites trisulcatus-Protohertzina anabarica Assemblage Zone and the overlying Paragloborilus subglobosa-Purella squamulosa Assemblage Zone of the Fortunian (the earliest Cambrian). The “Halkieria-dominant interval” yields some SSFs likely affiliated with ostracods in view of size and morphology, suggesting that arthropod body fossils appeared in the Fortunian, considerably earlier than previously believed.     

Chancelloriid sclerites from the lowermost Cambrian of North China and discussion of sclerite taxonomy

Chancelloriid sclerites from the lowermost Cambrian Xinji Formation (Series 2, Stage 3), southern margin of the North China platform, are systematically described. Thousands of isolated sclerites from three sections are assigned to three genera and four species, including Chancelloria cf. eros, Allonnia tripodophora, Archiasterella pentactina, and Ar. tetraspina. To accurately document the taxonomic significance of the sclerite structure, modified formulas (m+nC, m+nA, and m+0) are put forward to represent the full series and variation of sclerite forms. Based on the sclerite construction, statistical analysis on the proportions of different sclerite forms in the rock samples and the composition of sclerites in previously described chancelloriid scleritomes, a new scheme for identification and classification of isolated sclerites reconciled within the framework of the complete scleritome, is proposed.     

Intraspecific variation of trunk segmentation in the oryctocephalid trilobite Duyunaspis duyunensis from the Cambrian (Stage 4, Series 2) of South China

A detailed exploration of growth and trunk segmentation of the oryctocephalid trilobite, Duyunaspis duyunensis Chang & Chien in Zhou et al. 1977, from the lower Cambrian (Stage 4, Series 2) Balang Formation in western Hunan Province, South China, is presented. Because of the excellent preservation, the complete post‐protaspid ontogenetic series from merapsid degree 0 to the holaspid phase is described. The ontogenetic series reveals new information on morphological changes such as the migration of the posterior branch of the facial sutures (from proparian to opisthoparian) and contraction of the posteromedial notch in the pygidium. The abundance of articulated specimens available from a narrow stratigraphical interval makes this material singularly useful for studying the morphogenesis and post‐embryonic growth of D. duyunensis in comparison with other oryctocephalids. Strong evidence that multiple numbers of pygidial segment are recognized in each meraspid degree as well as in the holaspid period showed unusual intraspecific variability in the rate of trunk segmentation, providing insights into how Cambrian subisopygous trilobites controlled their body patterning, including size, shape and trunk segment number in both thorax and caudal plate during growth.     

End-Permian mass extinction of calcareous algae and microproblematica from Liangfengya,South China

A new assemblage of calcareous algae and microproblematica is reported from the Changxing Formation at the Liangfengya section in Chongqing, South China. This assemblage comprises eighteen species of seven genera, including three genera of gymnocodiaceaens (Gymnocodium, Permocalculus, and Tauridium), three genera of dasycladaleans (Epimastopora, Macroporella, and Mizzia), and one genus of microproblematica (Pseudovermiporella). A new algal species, Tauridium elongatum nov. sp., is described. Quantitative analysis indicates that the last occurrences of 22% of the species (4 out of 18) fall into a 52 cm thick interval of the uppermost Changhsingian. No calcareous algae are found in the Permian-Triassic boundary (P-T boundary) beds and the overlying Feixianguan Formation. Four out of 10 species have a stratigraphic abundance greater than 10% and all vanished in the uppermost Changhsingian. An abrupt extinction for calcareous algae occurred in the Clarkina yini conodont Zone.     

Cambrian Series 3 lithistid sponge–microbial reefs in Shandong Province,North China: reef development after the disappearance of archaeocyaths

The Cambrian Series 3 Zhangxia Formation in Shandong Province, North China, includes small‐scale lithistid sponge–microbial reefs. The lithistid sponges grew on oolitic and bioclastic sediments, which were stabilized by microbial activities. The relative abundances of microbial components (e.g. calcimicrobe Epiphyton and stromatolites) vary among the reefs. However, the microbial components commonly encrusted or bound the lithistid sponges, formed remarkable encrustations on the surfaces of the sponges. Epiphyton especially grew upward and downward. The lithistid sponges thus provided substrates for the attachment and development of microbes, and the microbes played essential roles as consolidators, by encrusting reef‐building sponges. Additionally, the lithistid sponges were prone to degradation via microbial activities and diagenetic processes, and were thus preserved as micritic bodies, showing faint spicular networks or abundant spicules. Such low preservation potential within the reef environment obscured the presence of the sponges and their widespread contribution as reef‐building organisms during the Cambrian. During the prolonged interval after the demise of archaeocyaths, purely microbial reefs, such as stromatolites and thrombolites have been considered to be the principal reef builders, in association with rare lithistid sponge–microbial associations. However, recent findings, including those from Shandong Province and Korea, suggest that the lithistid sponge‐bearing reefs were more extensive during the Epoch 3 to the Furongian than previously thought. These lithistid sponge–microbial reefs were precursors of the sponge–microbial reefs that dominated worldwide in the Early Ordovician.     

The evolutionary relationships of the earliest known cycloneuralians and a new record from the Cambrian Fortunian of South China

The Cambrian Fortunian Kuanchuanpu Formation and coeval strata in southern Shaanxi and northern Sichuan Provinces contain phosphatic carbonate that has yielded unambiguous body fossils of cycloneuralians. To date, six genera and species as well as nine indeterminate forms have been reported, and they provided information on the earliest known cycloneuralians and ecdysozoans. Among them, an indeterminate form is critical for understanding the evolution of the Fortunian cycloneuralians, and here I formally name it as Xinliscolex intermedius n. gen. n. sp. It has a series of at least seven longitudinal spinose sclerites on the dorsal side, and sensory structures such as pores and tubules distributed randomly among the annuli. The trunk end has two pairs of bilaterally arranged caudal spines. Micro-CT analysis reveals the preservation of a pharynx with inverted introvert with internally hollow scalids, an esophagus, midgut and hindgut, and a terminal anus. The digestive tract of Xinliscolex intermedius resembles that of modern priapulids, implying that the earliest known cycloneuralians have evolved a relatively derived digestive system, and this lays a foundation for their further diversification and evolution.     

A new assemblage of dabashanellids (Crustacea: Phosphatocopida) from Cambrian Stage 3 of western Hubei Province,South China

Dabashanellids are univalved phosphatocopid crustaceans, and are among the earliest crustaceans known so far. They were reported mainly from the Cambrian Stage 3 of China, but only a single species was established. Here I report a new assemblage of dabashanellids from the Cambrian Stage 3 Shuijingtuo Formation in western Hubei Province, South China. All specimens are sub-millimetre in length, and are preserved as three-dimensional carbonaceous hollow shields. Five species of Dabashanella were recognized, including the previously reported D. hemicyclica, three new species, D. longa n. sp., D. semiorbiculata n. sp., D. unispinata n. sp., and an indeterminate form Dabashanella sp. In addition, five indeterminate forms of arthropods were recovered. These dabashanellids represent part of a meiofauna that lived in the off-shore, deeper environments of mid-northern Yangtze Sea during Cambrian Age 3. This study shows greater morphological diversity and ecologic disparity of dabashanellid crustaceans, and supports the previous suggestion that the invasion of meiofaunal ecdysozoans into the off-shore, deeper environments probably occurred simultaneously with their radiation in the shallow water.     

Potential global stratotype sections and points in China for defining Cambrian stages and series

Three stratigraphic sections in the Cambrian of China that contain complete successions across important biohorizons having chronostratigraphic value, and that may be useful for developing Cambrian stage or series boundaries, are reviewed. The Wuliu section (Guizhou, South China), contains the FAD of Oryctocephalus indicus at 58.2 m above the base of the Kaili Formation. The Wangcun North section (Hunan, South China), contains the FADs of Ptychagnostus (or Acidusus) atavus, Ptychagnostus punctuosus, and Lejopyge laevigata at 1.2 m, 56.7 m, and 111.3 m, respectively, above the base of the Huaqiao Formation. The Xiaoyangqiao section (Jilin, Northeast China), contains the FAD of Cordylodus proavus in the Fengshan Formation at 10.89 m above the zero point of the section. A fourth potential stratotype section, the Wangcun South section (Hunan, South China), which contains the P. atavus, P. punctuosus, and L. laevigata zones, is discussed briefly.     

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Reassessment of the mode of life of Pagetia Walcott, 1916 (Trilobita: Eodiscidae) based on a cluster of intact exuviae from the Kaili Formation (Cambrian) of Guizhou, China

Pagetia is the most abundant trilobite in the Kaili Formation (lower to middle Cambrian). During the course of studying the museum collections of Kaili trilobites ( n  > 1000), a cluster containing 22 pagetiid individuals at various growth stages is noted. Specimens record the growth range from the degree 0 meraspid to late holaspid phases. Based on the relative completeness of the moults except for missing free cheeks, these specimens are interpreted as intact exuviae that had undergone minimal transport prior to burial. If Kaili pagetiids had pelagic or planktonic living habits as previously suggested, it would be difficult to explain the presence of intact exuviae in clusters. Therefore, Kaili pagetiids are interpreted here as having a benthic mode of life after the onset of the meraspid phase.     

Thoracic structure and enrolment style in middle Cambrian Eccaparadoxides pradoanus presages caudalization of the derived trilobite trunk

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

The North China Steptoean positive carbon isotope excursion and its global correlation with the base of the Paibian Stage (early Furongian Series), Cambrian

The use of carbon isotope excursion in Cambrian stratigraphical correlation is a standard practice at both the intercontinental and intracontinental scales. The Steptoean positive carbon isotope excursion (SPICE) is one of the prime examples in this regard in correlating the base of the Paibian Stage and Furongian Series. A lack of definite SPICE evidence in the North China craton has been a challenge in precision correlation between North China and other palaeo‐continents. This study provides new carbonates carbon isotope data from the type locality of the Changshan Formation in Hebei Province, North China. Our new δ13C_carb data provide new objective evidence for the presence of the SPICE in North China. The sampling section is relatively condensed, and the interval of the SPICE curve is less than one and half‐metres after analysing 64 samples (the sampling interval within the SPICE is less than 10 cm). The onset of the SPICE curve in Tangshan, Hebei, occurs in the barren interval between the Neodrepanura and Chuangia trilobite zones. Based on this study and previous work, this could imply the middle part of the Prochuangia‐Paracoosia trilobite Zone in North China and can be correlated with the base of the Paibian Stage and Furongian Series.     

A new scyphozoan from the Cambrian Fortunian Stage of South China

Animals with radial symmetry are abundant in the Cambrian Fortunian Stage of South China, but with relatively low diversity: representatives include Olivooides, Quadrapyrgites, carinachitiids, hexangulaconulariids and Pseudooides. Here, we report a new radial animal, Qinscyphus necopinus gen. et sp. nov., from the Fortunian small shelly fauna of southern Shaanxi Province, South China. Qinscyphus necopinus has a cup‐shaped profile, with slightly raised annuli and five groups of triangular thickenings in pentaradial symmetry. This organism has a comparable morphology to, and thus a close affinity with, Olivooides and Quadrapyrgites, and is interpreted as a coronate scyphozoan. This discovery adds a new crown‐group cnidarian to the Cambrian Explosion.