The tommotiid Kelanella and associated fauna from the early Cambrian of southern Montagne Noire (France): implications for camenellan phylogeny

A carbonate bed of the Pardailhan Formation, early Cambrian, southern Montagne Noire (southern France), provided microfossils such as Hyolithellus sp., Torellella cf. mutila and Torellella sp. along with numerous disarticulated pieces of composite skeletons such as valves of the brachiopod Eoobolus priscus and of the bradoriid Monceretia erisylvia, and chancelloriid sclerites (Chancelloria sp.). The assemblage also furnished a rich set of sclerites from the tommotiid Kelanella altaica. Five morphological variations of the latter have been identified. The presence of concentric ribs formed by distal inflation of selected shell laminae in Kelanella supports its assignment to the camenellans. More particularly, the presence of transverse structures within the internal cavity (septa) of Kelanella suggests a close relationship with the Lapworthellidae. However, the latter differ from Kelanella by the continuous morphological variation along their scleritome which is also composed of simple conical elements with uniform ornamentation. Several forms of Kelanella are similar to mitral and sellate sclerites of Camenella, whereas some other forms are comparable to Kennardia. The new material suggests that Kelanella occupies a transitional position between Lapworthellidae and the grouping of Tommotiidae and Kennardiidae. Such a phylogenetic position also implies that the number of sclerite morphotypes tends to decrease within the camenellan scleritome during evolution.     

Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France)

The disused quarry east of Castelsec offers a view of shallow-marine carbonates of the poorly known Uppermost Mississippian of the Montagne Noire. At Castelsec, sections are studied in two characteristic facies types (bioclastic wackestone and microbial dominated boundstone) of the Upper Mississippian. The succession is rich in rugose corals and carbonate microfossils. Six genera with seven species belonging to a rugose coral fauna consisting of at least eight genera with several species are described herein; Dibunophyllum castelsecensis sp. nov. is described as new. Twenty-seven carbonate microfossils of different groups have been identified. The Castelsec succession is Brigantian in age, based on the stratigraphic occurrence of rugose corals, foraminifers, and calcareous algae observed in both sections. The rugose coral fauna shows relationships with the well-known fauna of northwestern Europe and the Ouralian-Asian Province. Typical elements of northwestern Europe are missing at Castelsec and vice versa. This differentiation between north and south is interpreted as responses to different palaeolatitudes and tectonic settings.     

Organic-walled microfossils from Cambrian Stage IV in the Jiaobang section,eastern Guizhou,China

The lower Cambrian succession in the Jiaobang section, Jianhe County, eastern Guizhou, China, includes, in ascending order, the Bianmachong, Balang, and Tsinghsutung formations, with a total thickness of about 645 m. Twenty-six morphological genera (including one new genus) are identified from the Balang and the underlying Bianmachong formations, many of which are common and widely distributed. Six acritarch assemblages are discerned in the Balang Formation. They are, in ascending order, the Adara alea‒Skiagia ornata, the Acrum radiale‒Pterospermella velata, the Comasphaeridium molliculum‒Solisphaeridium baltoscandium, the Corrugasphaera perfecta n. sp.‒Pterospermella vinctusa n. sp., the Acrum novum‒Heliosphaeridium oligum, and the Acrum membranosum‒Adarve diafanum acritarch assemblages. An obvious change of organic-walled microfossil assemblages occurred in the interval between 84 m and 98 m from the bottom of the Balang Formation which roughly corresponds to the boundary between the Oryctacarella duyunensis trilobite Zone and the overlying Arthricocephalus chauveaui trilobite Zone. In addition, organic-walled microfossils are scarce in about 24 m thick from the bottom of the Balang Formation. One new genus and five new species including Plagasphaera balangensis n. gen. n. sp., Asteridium tubulus n. sp., Cymatiosphaera spina n. sp., Corrugasphaera perfecta n. sp., and Pterospermella vinctusa n. sp. are described.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

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

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

Biostratigraphy of the Peri-Gondwana Cambrian trilobite fauna (northern Kerman,Iran) and correlation with other countries

Polymeroid trilobite fauna from two stratigraphic sections (Godbondar and Kuhbanan) of the Kuhbanan Formation in northern Kerman (central Iran) were studied and subjected to biostratigraphic analysis. Eleven genera and species are recognised from the latest Early Cambrian and Middle Cambrian Peri-Gondwanian successions of the study sections. The recognised fauna includes Afghanocare lategenatum, Blountia blountia, Iranoleesia sp., Iranoleesia pisiformis, Kermanella kuhbananensis, Kermanella lata lata, Kermanella lata minuta, Kermanella sp., Redlichia chinensis, Redlichia noetlingi and Redlichia sp. Based on trilobite distribution, three trilobite biozones were recognised in the study sections, namely Redlichia noetlingi biozone, Kermanella kuhbananensis biozone and Iranoleesia pisiformis biozone. The age of the study sections is late Early Cambrian to late Middle Cambrian based on the recognised trilobite biozones. The recognised late Early Cambrian trilobite assemblages (especially Redlichia and Kermanella) from northern Kerman are similar to those found from some other parts of Gondwana or Peri-Gondwana terrains (north India, Pakistan, northwestern Kashmir, Tajikistan, South Australia, South China and Afghanistan) and show affinities with fauna found in some other parts of Iran (Alborz, northern Iran; Tabas, eastern Iran; southeast Karman).     

Lower Cambrian Small Shelly Fossils of northern Sichuan and southern Shaanxi (China), and their biostratigraphic importance

Extensive information has been acquired on the occurrence of Lower Cambrian Small Shelly Fossils (SSF) in the shallow water sediments of the western fringe of the Yangtze Platform (eastern Yunnan and central Sichuan Provinces) during the last two decades. However, less is known on the temporal and spatial distribution of SSFs on the northern margin of the Yangtze Platform. Here we report SSF associations from Kuanchuanpu (Ningqiang County, southern Shaanxi Province), Shatan, and Xinli (both Nanjiang County, northern Sichuan Province) representing the Southwest Shaanxi region and from Sanlangpu, and Xiaowan (Xixiang County, southern Shaanxi Province) of the Southeast Shaanxi - Northwest Hubei region. The thickness of the Lower Cambrian SSF-bearing strata strongly decreases in this region from West to East and the stratigraphic hiatus increases. An earliest Cambrian hiatus is widely distributed on the northern margin of the Yangtze Platform. At the same time, a syngenetic brecciation occurred in wide shallow shelf settings. A tectonic mobilization, possibly related to the assembly of Gondwana following the breakup of Rodinia, may be responsible for this. In sections near Kuanchuanpu (Ningqiang County), the Meishucunian (equivalent to the Nemakit-Daldynian to Tommotian of Siberia) succession comprises ca. 55 m of dark phosphatic carbonates, siltstones, and cherts. The SSF associations are similar to those of East Yunnan and, in ascending order, can be discriminated into the Anabarites trisulcatus - Protohertzina anabarica Zone, the Watsonella yunnanensis Zone (previously mentioned as Heraultipegma yunnanensis Zone), and the Sinosachites flabelliformis - Tannuolina zhangwentangi Zone. The Siphogonuchites triangularis - Paragloborilus subglobosus Zone cannot be unambiguously detected in the Southwest Shaanxi region. The occurrence of embryos and eggs within the Anabarites trisulcatus - Protohertzina anabarica Zone, which has been reported earlier, is remarkable. The previously reconstructed life cycle and taxonomic treatment of Olivooides / Punctatus are re-discussed. The earliest SSF associations (Anabarites trisulcatus - Protohertzina anabarica Zone to Watsonella yunnanesis Zone) are missing in the Southeast Shaanxi - Northwest Hubei region and the Ninella tarimensis - Cambroclavus fangxianensis Assemblage Zone and Rhombocorniculum cancellatum Taxon Range Zone are introduced herein.     

Oldhamia radiata: the trace of an undermat miner from the Cambrian Series 2 Xinji Formation,western Henan,North China

Oldhamia trace fossils represent a complex horizontal burrow system usually associated with microbial mats, occurring in the upper part of the Cambrian Stage 2 of the Terreneuvian Series, with a peak in abundance during Stage 3 and 4 of Series 2, and then rapidly disappearing in the Wuliuan Stage of the Miaolingian Series. We here report Oldhamia radiata from the Cambrian Series 2 Xinji Formation in western Henan of North China, associated with microbial mats, as evidenced by the presence of wrinkle structures. The distinctive patterns indicate the exploitation of microbial mats by the tracemakers of O. radiata, featuring a specific feeding strategy under the mats where possibly anoxic and sulfidic conditions prevailed during the early Cambrian.     

Morphology and ultrastructure of epilithic versus cryptic,microbial growth in lower Cambrian phosphorites from the Montagne Noire,France

The lower Cambrian grainy phosphorites of the northern Montagne Noire occur interbedded with grey to black, laminated to massive shales and limestones deposited along the edge of a continental shelf, associated with slope‐related facies and unstable substrates. The concentration of phosphate took place by repeated alternations of low sedimentation rates and condensation (hardgrounds), in situ early‐diagenetic precipitation of fluorapatite, winnowing and polyphase reworking of previously phosphatized skeletons and hardground‐derived clasts. The succession of repeated cycles of sedimentation, phosphate concentration, and reworking led to multi‐event phosphate deposits rich in allochthonous particles. Phosphogenesis was primarily mediated by microbial activity, which is evidenced by the abundance of phosphatized putative microbial remains. These occur as smooth and segmented filaments, sheaths, and ovoid‐shaped coccoids. These simple morphologies commonly form composite frameworks as a result of their aggregation and entanglement, leading to the record of biofilms, microbial mats, and complex networks. These infested the calcitic skeletonized microfossils that littered the substrate. Microbial activity evidences epilithic (anisotropic coatings on skeletons), euendolithic (perforating skeletal walls), and cryptoendolithic (lining inter‐ and intraparticulate pores) strategies, the latter dominated by bundles of filaments and globular clusters that grew along the cavities of helcionellids and hyoliths. According to their epilithic versus cryptic strategies, microbial populations that penetrated and dwelled inside hard skeletal substrates show different network and colonial morphologies. These early Cambrian shell concentrations were the loci of a stepwise colonization made by saprophytic to mutualistic, cyanobacterial–fungal consortia. Their euendolithic and cryptoendolithic ecological niches provided microbial refugia to manage the grazing impact mainly led by metazoans.     

A new rhynchonelliform brachiopod Longtancunella with soft-part preservation from the Hongjingshao Formation (Cambrian Stage 3) in Yunnan,South China

Rhynchonelliform brachiopods made their first appearance in early Cambrian, and became a major group within the palaeozoic evolutionary fauna since late Cambrian. Exceptionally preserved fossils from the early Cambrian Lagerstätten provide valuable chances to investigate their phylogeny and ecology. Longtancunella is one of the most interesting early rhynchonelliforms, and has been mainly recovered from the Chengjiang Lagerstätte (Series 2, Stage 3). Here, we report a new rhynchonelliform Longtancunella xiazhuangensis n. sp. from the lower Hongjingshao Formation (upper Stage 3) in Yunnan Province, China. These specimens were well preserved with soft parts, including pinnate mantle canal system and a pedicle. It is identified as a new species based mainly on its difference in shell ornamentations, pinnate mantle canals and pedicle morphology from the type species. Its pedicle looks unusually stout with distinct annulated lamellae on the surface, and reveals crucial evidence in illustrating its ecology and settling strategy as an early marine epifauna. The ecological interaction between L. xiazhuangensis and other marine animals also provides insights into the food web structure in the early Cambrian.     

Redescription of Hexaconularia He and Yang, 1986 (Lower Cambrian, South China): implications for the affinities of conulariid-like small shelly fossils

Abstract:  Hexaconularia , a Lower Cambrian small shelly fossil (SSF) that has been allied with conulariids and scyphozoan cnidarians, is redescribed and refigured. A salient feature of this monospecific genus is the presence of distinct apical and abapical regions. The apical region probably represents an embryonic shell that apparently lacked a basal attachment structure. Comparisons of this feature with the apical end of the smallest known conulariids and with conulariids terminating in an apical wall (schott) reveal substantial differences in structure and ornamentation. Differences in apical anatomy between conulariids and Arthrochites , possibly the nearest SSF relative of Hexaconularia , are also apparent. Comparisons of Hexaconularia with Punctatus , an SSF taxon showing distinct apical and abapical regions in both posthatching specimens and prehatching embryos, suggest that the early development of Hexaconularia was direct. These results have important implications for hypotheses of a conulariid/scyphozoan affinity for Hexaconularia and its possible SSF relatives, and they suggest that Hexaconularia -bearing strata may yield prehatching embryos of this genus.     

Pelecyogyra (Mollusca) from the Early Ordovician of the Montagne Noire,France

A new occurrence of the onychochilid mollusc Pelecyogyra is reported from the late Floian Landeyran Formation (Apatokephalus incisus Biozone) in the Montagne Noire, southern France, where it is associated with the two tergomyans Carcassonnella courtessolei and Thoralispira laevis, and a mollusc indet. With a low cap-shaped shell. The Montagne Noire specimens compare morphologically with Pelecyogyra fezouataensis from the late Tremadocian Fezouata Formation (Araneograptus murrayi Biozone) in Morocco and are tentatively referred to this species. Each occurrence of Pelecyogyra is from one or two localities only, but separated in time. They differ further in the deeper facies, association with Carcassonnella and Thoralispira, and non-gregarious occurrence in Montagne Noire compared to the shallower facies, lack of other molluscs and gregarious occurrence in the Fezouata Formation. In the Lower Ordovician, Pelecyogyra, Carcassonnella courtessolei, the rare C. vizcainoi and Thoralispira laevis are taxa that occur only in Montage Noire and the Anti-Atlas, while Lesueurilla prima also occurs in the Prague basin of the Czech Republic. Contrary to other faunal elements, like echinoderms and trilobites, these genera seem to have a wide environmental tolerance and are long ranging taxa. The main occurrence of the three key taxa Carcassonnella, Thoralispira and Lesueurilla in the Montagne Noire is in the lower part of the La Maurerie Formation (Taihungshania miqueli acme Zone), while the same taxa appear just slightly later in Morocco (?Cymatograptus protobalticus Biozone). Migration to the Anti-Atlas during the early Floian seems to have been possible during a period of general faunal exchange. Pelecyogyra may either have migrated from the Anti-Atlas to Montagne Noire during the same interval or possibly during the late Floian as indicated by shared faunal elements between these two regions at that time.     

A new mobergellan (small shelly fossils) from the early middle cambrian of morocco and its significance

A new mobergellan genus and species,Tateltella ranoculata, is described from the early Middle Cambrian (Agdzian Stage) of Morocco. The new taxon is characterized by only four pairs of muscle scars and is furthermore distinguished from other mobergellans by its strongly concave shape and its distally rising muscle scars. The individual specimens ofTateltella ranoculata distinctly vary in size and display different ontogenetic stages. Juvenile, intermediate, and adult stages can be distinguished by means of the development of the muscle scars that differ in Position relative to the apex, size, and distinctness between individual stages. The shell ofT. ranoculata is composed of a succession of thin phosphatic lamellae separated by interlamellar gaps, presumably originally filled by organic material. The interlamellar gaps may be divided by septum-like structures producing discrete cavities. The specimens are the youngest mobergellans known so far and correlation of their stratigraphic position suggests a correspondence with the lower part of the Amgan stage of the Siberian Platform. In addition, they are the first reported mobergellans from the present day continent Africa. Other mobergellan taxa and mobergellan-like species are briefly reviewed and the genusHippoklosma Missarzhevsky, previously assigned to the Mobergellidae, is excluded from the family due to its different shell structure. An evolutionary trend of reduction in the number of muscle scars from 14 in the early Early Cambrian to only eight in the early Middle Cambrian is apparent among mobergellans.      

龙凤山藻在澄江早寒武世生物群中的发现

主要报道了产自云南昆明海口耳材村早寒武世筇竹寺组玉案山段澄江生物群中可能为漂浮生长的宏观藻类化石-心型龙凤山藻（新种）（Longfengshania cordata sp.nov.)和中华豆芽藻（新属、新种）（lantulaformis sinensis gen.et sp.nov)。这些化石的发现进一步显示了澄江生物群种物多样性,为揭示该生物群爆发性演化提供了新的证据。通过对龙凤山藻属亲缘关系的深入比较研究,进一步证明该类化石应归属宏观藻类以及它们遗传上的稳定性、演化上的保守性和环境上的适应性。     

龙凤山藻在澄江早寒武世生物群中的发现

主要报道了产自云南昆明海口耳材村早寒武世筇竹寺组玉案山段澄江生物群中可能为自由漂浮生长的宏观藻类化石--心型龙凤山藻(新种)(Longfengshania cordata sp. nov.)和中华豆芽藻(新属、新种)(Plantulaformis sinensis gen. et sp. nov.).这些化石的发现进一步显示了澄江生物群物种多样性,为揭示该生物群爆发性演化提供了新的证据.通过对龙凤山藻属亲缘关系的深入比较研究,进一步证明该类化石应归属宏观藻类以及它们遗传上的稳定性、演化上的保守性和环境上的适应性.     

Early Cambrian Teichichnus-dominated ichnofabrics and palaeoenvironmental analysis of the Caerfai Group, Southwest Wales, UK

The early Cambrian Caerfai Group of Pembrokeshire in Southwest Wales yields an intensely bioturbated Teichichnus-dominated ichnofabric, and provides evidence for the early evolution of infaunal organisms. Three units of the Caerfai Group comprise the focus of this study in sections exposed at Caerfai Bay. In the St. Non's Sandstone few primary sedimentary structures are observed, bedding being predominantly massive with subordinate parallel and current ripple cross-lamination. The topmost St. Non's Sandstone contains common soft-sediment deformation phenomena, and collectively deposition is believed to have taken place in the proximal subaqueous reaches of a steep-fronted delta system. Teichichnus is the dominant trace, with subordinate Planolites, Palaeophycus and possible Rhizocorallium.The overlying Caerfai Bay Shales were deposited from mass flows with evidence of downslope creep suggesting deposition on a steep delta front. Sandstone debrites are common, as are tuffaceous beds with evidence of reworking by bottom currents. Sandstone and tuff event beds contain common Skolithos and Arenicolites with less frequent Planolites. Interbedded mudstones contain infrequent Teichichnus.The Caerfai Bay Shales coarsen and thicken-upwards into the Caerbwdy Sandstone. Debrite sandstone beds dominate, and bed thickening upsection indicates progradation of the delta system. The dominant trace fossil within the unit is Teichichnus which is constrained to the top few centimetres of individual sandstone beds. Less common are Planolites and Palaeophycus.The size and abundance of Teichichnus in the Caerfai Group are similar to, or exceed those in contemporaneous sections in Avalonia and Baltica. Local environmental factors were probably the primary control on trace fossil distribution. Substrate and salinity are proposed as the main influences on ichnodiversity and size in the Caerfai Group. Teichichnus is most abundant in the St. Non's Sandstone, where the proximal reaches of the delta experienced reduced salinity and possibly higher oxygen levels. Burrow diameter progressively decreases upsection, and bioturbation becomes less pervasive in subsequent units of the Caerfai Group. This indicates that by the Cambrian Stage 2, significant advances into reduced salinity environments had occurred, along with the dispersal of the Teichichnus producer throughout the Avalonia-Baltica seaway. Correlation within Avalonia indicates that a transgression in the Cambrian Stage 2 initiated the deposition of the Caerfai Group marking the onset of the Welsh Basin.