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.     

A crown‐group demosponge from the early Cambrian Sirius Passet Biota,North Greenland

Calibration of the divergence times of sponge lineages and understanding of their phylogenetic history are hampered by the difficulty in recognizing crown versus stem groups in the fossil record. A new specimen from the lower Cambrian (Series 2, Stage 3; approximately 515 Ma) Sirius Passet Biota of North Greenland has yielded a diagnostic spicule assemblage of the extant demosponge lineages Haploscleromorpha and/or Heteroscleromorpha. The specimen has disarticulated approximately in situ, but represents an individual sponge that possessed monaxon spicules combined with a range of slightly smaller sigma, toxa and unique spiral morphologies. The combination of spicule forms, together with their relatively large size, suggests that the sponge represents the stem lineage of Haploscleromorpha + Heteroscleromorpha. This is the first crown‐group demosponge described from the early Cambrian and provides the most reliable calibration point currently available for phylogenetic studies.     

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.     

葫芦虫化石在寒武纪关山生物群中的发现及其意义

报道产自昆明高楼房的寒武纪早期关山生物群中的葫芦虫化石,这是该类动物在全球的第三例化石记录。葫芦虫化石在关山生物群的发现对了解该类动物以及寒武纪动物群生态和演化等均有着重要意义。     

Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China

Botting, J.P., Muir, L.A., Xiao, S., Li, X. & Lin, J.‐P. 2012: Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China. Lethaia, Vol. 45, pp. 463–475. The relationships of the extant sponge classes, and the nature of the last common ancestor of all sponges, are currently unclear. Early sponges preserved in the fossil record differ greatly from extant taxa, and therefore information from the fossil record is critical for testing hypotheses of sponge phylogenetic relationships that are based on modern taxa. New specimens of the enigmatic sponge Lenica sp., from the Early Cambrian Hetang Biota of South China, exhibit an unusual spicule structure. Each spicule consists of a siliceous core with an axial canal, an organic outer layer and a middle layer interpreted to have been originally calcium carbonate. This finding confirms previous work suggesting the existence of biminerallic spicules in early sponges. Combined with data from other early sponges, the new findings imply that the two fundamental spicule structures of modern sponges were derived from a compound, biminerallic precursor. Spicules are therefore homologous structures in Calcarea and Silicea, and if sponges are paraphyletic with respect to Eumetazoa, then spicules may also have been a primitive feature of Metazoa. □Calcarea, Early Cambrian, Hetang Biota, phylogeny, Silicea, taphonomy.     

陕西镇巴下寒武统水井沱组小壳化石新属种

本文描述了陕西镇巴下寒武统水井沱组中的小壳化石３新属５新种。样品采自镇巴小洋坝曾家坡,共生化石有三叶虫、古介形虫、海绵骨针和大量软舌螺等。化石组合面貌表明其地质时代属筇竹寺期。另外,在峡东水井沱组中发现了微网虫（Ｍｉｃｒｏｄｉｃｔｙｏｎｓｐ．）化石骨片（图版Ｉ,图１８,１９）,可与云南筇竹寺组及陕南水井沱组对比,说明Ｍｉｃｒｏｄｉｃｔｙｏｎ广泛分布于扬子地台区。     

Additional trilobite fauna from the basal part of the Cambrian Series 3, Stage 5, Kunzam La (=Parahio) Formation,Parahio Valley,Spiti (Northwest Himalaya), India and its biostratigraphic significance

Additional Cambrian Series 3, Stage 5, trilobite fauna comprising Peronopsis sp., Eosoptychoparia cf. Spinosa Gaotanaspis cf. pingzhaiensis and Gaotanaspis cf. transversa have been recorded just immediately above the known Oryctocephalus indicus biozone (Cambrian Series 3, Stage 5) in the Parahio valley section (Spiti region). The FAD of Peronopsis and LAD of Gaotanaspis are used to establish a Peronopsis–Gaotanaspis concurrent biozone immediately above the Oryctocephalus indicus biozone. The first records of Gaotanaspis cf. pingzhaiensis and G. transversa from the Cambrian of Spiti region and the other faunal elements are correlated with the Peronopsis taijiangensis biozone of the Kaili Formation (South China). The stratigraphic thickness from the base of the O. indicus biozone to the top of the Peronopsis–Gaotanaspis concurrent biozone in the Kunzam La (=Parahio) Formation and its comparison to the Kaili Formation (South China) indicate a possible stratigraphic condensation in the basal part of the Cambrian Series 3, Stage 5 of the Parahio valley (Spiti). Sedimentological and sequence stratigraphic analysis supports this contention.     

皖南早寒武世荷塘组海绵骨针化石

本文报道皖南休宁县早寒武世荷塘组黑色页岩中产出的海绵骨针化石组合,这些海绵骨针化石具有较高的丰度和分异度,它们以二轴四射针、T型针、三轴六射针和三轴五射针为主。骨针形态完整,并保存了内部轴丝、轴管以及同心圈层等微细构造。黄铁矿化在化石的保存中起了重要的作用,化石产出的时代可能为梅树村阶至筇竹寺阶（Tommotian-Atdabanian),这个化石组合证实了海绵动物在早寒武世已开始迅速分异。     

Early sponge evolution: A review and phylogenetic framework

Sponges are one of the critical groups in understanding the early evolution of animals. Traditional views of these relationships are currently being challenged by molecular data, but the debate has so far made little use of recent palaeontological advances that provide an independent perspective on deep sponge evolution. This review summarises the available information, particularly where the fossil record reveals extinct character combinations that directly impinge on our understanding of high-level relationships and evolutionary origins. An evolutionary outline is proposed that includes the major early fossil groups, combining the fossil record with molecular phylogenetics. The key points are as follows. (1) Crown-group sponge classes are difficult to recognise in the fossil record, with the exception of demosponges, the origins of which are now becoming clear. (2) Hexactine spicules were present in the stem lineages of Hexactinellida, Demospongiae, Silicea and probably also Calcarea and Porifera; this spicule type is not diagnostic of hexactinellids in the fossil record. (3) Reticulosans form the stem lineage of Silicea, and probably also Porifera. (4) At least some early-branching groups possessed biminerallic spicules of silica (with axial filament) combined with an outer layer of calcite secreted within an organic sheath. (5) Spicules are homologous within Silicea, but also between Silicea and Calcarea, and perhaps with Homoscleromorpha. (6) The last common ancestor of extant sponges was probably a thin-walled, hexactine-bearing sponge with biminerallic spicules. (7) The stem group of sponges included tetraradially-symmetric taxa that grade morphologically into Cambrian fossils described as ctenophores. (8) The protomonaxonid sponges are an early-branching group, probably derived from the poriferan stem lineage, and include the problematic chancelloriids as derived members of the piraniid lineage. (9) There are no definite records of Precambrian sponges: isolated hexactine-like spicules may instead be derived from radiolarians. Early sponges had mineralised skeletons and thus should have a good preservation potential: the lack of sponge fossils in Precambrian strata may be due to genuine absence of sponges. (10) In contrast to molecular clock and biomarker evidence, the fossil record indicates a basal Cambrian diversification of the main sponge lineages, and a clear relationship to ctenophore-like ancestors. Overall, the early sponge fossil record reveals a diverse suite of extinct and surprising character combinations that illustrate the origins of the major lineages; however, there are still unanswered questions that require further detailed studies of the morphology, mineralogy and structure of early sponges.     

Benthic primary producers in exceptionally preserved Cambrian biotas of North China

Strata situated within the South China Craton host several key Cambrian macroalgal assemblages, including those of the Chengjiang, Kaili, and Tsinghsutung Lagerstätte. In this report, Cambrian macroalgae are described for the first time from the North China Craton, an area that, during the early Paleozoic, was not contiguous with the South China Craton. The material collected from the Mantou Formation and overlying Zhangxia Formation (Miaolingian Series) in Shandong Province is preserved as carbonaceous compressions and includes abundant simple tubular forms. These fall into two groupings on the basis of size and have characteristics broadly comparable to Fuxianospira gyrata and Sinocylindra yunnanensis, both of which are well-known from the Chengjiang biota and may represent simple siphonous green algae. Additional material includes irregular circular to oval films referable to Morania fragmenta and a tuft-like form superficially similar to Marpolia spissa, taxa with possible cyanophyte affinities described initially on the basis of material from the Burgess Shale but also known from the Cambrian of South China. The material is typical of the “Cambrian Flora” and provides further support for the cosmopolitan distribution of this noncalcified macroalgal flora at low latitude settings during the Cambrian.     

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.     

A new eocrinoid from the Guanshan Biota (Cambrian Series 2, Stage 4), with implication of the development of different attachment modes in early Cambrian

Eocrinoids are scarce in the Guanshan Biota (Cambrian Series 2, Stage 4), Yunnan Province, southwest China. Here, we introduce a new gogiid: Kunmingeocrinus cupuliformis n. gen. n. sp. which is characterized by a short stalk and a well-developed attachment disc. Preservation may indicate a weakly biomineralized body for the new taxon. Morphological similarities between the new taxon and other eocrinoids with attachment discs from Cambrian Lagerstätten of Guizhou Province (Series 2, Stage 4 and Miaolingian, Wuliuan) might suggest a similar mode of life. The eocrinoids from the Guanshan Biota possibly utilized different attachment modes.     

全尾宽背虫Bathynotus holopygus (Hall, 1859)在 贵州剑河八郎寒武系凯里组的首次发现及意义

宽背虫属Bathynotus (Hall, 1859)广泛分布于劳伦古大陆、中国华南地区、澳大利亚和西伯利亚, 为寒武纪第二世末期最重要的标志性三叶虫之一,已成为寒武系划分的重要指示化石。其模式种B. holopygus以其狭长的颊刺、胸轴宽, 第11节肋刺巨大, 第12、13节肋刺不发育为主要特征;具有很强的地域性, 仅分布于华南和北美, 在华南主要见于寒武系扬子区与江南区之间过渡区的黔东地区。B. holopygus产出地层延限较窄, 仅见于寒武系第二统第四阶顶部。本文所描述的部分标本采自贵州剑河八郎苗板坡剖面苗岭统及乌溜阶界线之下2 m左右, 是在该地的首次报道, 也是全球仅有的第4个产地。笔者通过数理统计值分析进一步描述B. holopygus的特征和发育趋势, 为进一步属种研究提供新材料; 根据新发现的唇瓣标本进一步推测其唇瓣与腹边缘为愈合状态。     

贵州松桃寒武系第二统清虚洞组腕足动物 Eohadrotreta zhenbaensis的发现

微型腕足动物化石镇巴始壮贝Eohadrotreta zhenbaensis常见于陕南、峡东水井沱组与黔东麻江九门冲组等地寒武纪地层中,在贵州松桃盘信寒武系第二统清虚洞组薄层灰岩中为首次发现。通过与华南其他地区E. zhenbaensis的背壳长宽比及中隔板等结构特征比较,发现松桃盘信清虚洞组E. zhenbaensis与浅水扬子地台的陕南、峡东地区的E. zhenbaensis壳体内部特征较为相似,包括肉茎孔形成阶段、肉茎孔封闭阶段及交互沟伸长阶段3个生长发育阶段标本,与较深水过渡区麻江的E. zhenbaensis略有差异。贵州松桃盘信寒武系第二统清虚洞组E. zhenbaensis的发现,扩展了该属化石在华南寒武纪第二世的古地理分布,丰富了该组腕足动物组成面貌,为研究E. zhenbaensis形态结构特征提供了新材料。     

First report of Schizopholis (Lingulata,Brachiopoda) from the Tsinghsutung Formation (Cambrian Series 2, Stage 4) in Guizhou,China

Schizopholis Waagen, 1885 is a genus of linguliform brachiopod, which is known from Cambrian Stage 4 to the Wuliuan Stage of Australia, Antarctica, Pakistan and China. Recently, new material of Schizopholis was discovered from the upper part of the Tsinghsutung Formation (Cambrian Series 2, Stage 4) near Balang village, Jianhe County, Guizhou Province, China. These specimens display the oval pedicle opening in the ventral valve, a median tongue and a pair of tubercles in the dorsal valve that are characteristic features of Schizopholis napuru (Kruse, 1990). This is the first time that this species has been described from Guizhou Province, China. Previously, fossils of this species are usually preserved in carbonate deposits, but the new material documented herein preserved in mudstone of the Tsinghsutung Formation, providing new information regarding both the paleogeographic distribution and paleoecology of this species.     

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.     

Carbonaceous preservation of Cambrian hexactinellid sponge spicules

Early fossil sponges offer a direct window onto the evolutionary emergence of animals, but insights are limited by the paucity of characters preserved in the conventional fossil record. Here, a new preservational mode for sponge spicules is reported from the lower Cambrian Forteau Formation (Newfoundland, Canada), prompting a re-examination of proposed homologies and sponge inter-relationships. The spicules occur as wholly carbonaceous films, and are interpreted as the remains of robust organic spicule sheaths. Comparable sheaths are restricted among living taxa to calcarean sponges, although the symmetries of the fossil spicules are characteristic of hexactinellid sponges. A similar extinct character combination has been documented in the Burgess Shale fossil Eiffelia. Interpreting the shared characters as homologous implies complex patterns of spicule evolution, but an alternative interpretation as convergent autapomorphies is more parsimonious. In light of the mutually exclusive distributions of these same characters among the crown groups, this result suggests that sponges exhibited an early episode of disparity expansion followed by comparatively constrained evolution, a pattern shared with many other metazoans but obscured by the conventional fossil record of sponges.     

‘Cambrian’ demosponges in the Ordovician of Morocco: Insights into the early evolutionary history of sponges

Cambrian spicular sponge faunas are dominated by a distinctive assemblage of demosponges and hexactinellids that are known from Burgess shale-type faunas worldwide. Most of these are previously unknown outside the Lower-Middle Cambrian (and perhaps Tremadoc) and have no obvious close relatives in later sequences. This paper describes examples of Choia sp., Pirania auraeum sp. nov. and Hamptonia christi sp. nov. from the Arenig of Morocco, associated with isolated hexactinellid spicules. A summary of the stratigraphic ranges of the major Cambrian sponge lineages is provided. These indicate an environmental contrast in the Lower Palaeozoic evolution of hexactinellids and non-lithistid demosponges, with demosponges probably undergoing cryptic diversification in nearshore environments during the Upper Ordovician.     

Early Cambrian organophosphatic brachiopods from the Xinji Formation,at Shuiyu section,Shanxi Province,North China

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.