Fossil preservation in the Neoproterozoic Doushantuo phosphorite Lagerstätte, South China

Phosphorites of the late Neoproterozoic Doushantuo Formation exposed in the vicinity of Weng'an, Guizhou Province, and Chadian, Shaanxi Province, South China, contain exceptionally well-preserved algal thalli, acritarchs, and globular microfossils interpreted as animal embryos. Combined optical microscopic and SEM observations provide insights into the taphonomy of phosphatized fossils. Algal cells and tissues are variably resistant to decay, and within preserved populations permineralization began at varying stages of degradation. In consequence, there is a spectrum of quality in cellular preservation. Algal cell walls, acritarch vesicles, and embryo envelopes are commonly encrusted by an isopachous rim of apatite, with cell interiors filled by collophane and later diagenetic dolomite. In contrast, blastomere surfaces of animal embryos are encrusted primarily by minute phosphatic spherules and filaments, possibly reflecting an immediately postmortem infestation of bacteria that provided nucleation sites for phosphate crystal growth. Thus, the same processes that gave rise to Phanerozoic phosphatized Lagerstätten - phosphatic encrustation, and impregnation, probably mediated by microbial activity - effected soft-tissue preservation in the Doushantuo Lagerstätte. It remains unclear how phosphatic ions and organic macro-molecules interact at the molecular level and to what extent specific microbial metabolisms or microenvironmental conditions control the phosphatization of soft tissues. New observations of phosphatized Doushantuo fossils include: a second locality (Chadian) for Wengania globosa , interpreted as an algal thallus and previously known only from Weng'an; microtunnels in Weng'an phosphorites interpreted as pyrite trails; and new taxa described from Weng'an: Meghystrichosphaeridium reticulatum (acritarch), Sarcinophycus radiatus (algal thallus), and one unnamed problematic form.     

湖北保康陡山沱组磷酸盐化微体化石组合

中国南方震旦系陡山沱组磷块岩、燧石和页岩中陆续发现的精美保存的化石生物群,为我们提供了一个栩栩如生的新元古代晚期生物演化的图景,表明多细胞真核生物在新元古代冰期之后发生了非常显著的分异。其中磷块岩化石库以生物细胞结构的完好保存而著称。继在贵州瓮安、陕西勉县茶店和江西上饶朝阳之后,湖北保康白竹磷矿陡山沱组磷块岩中也发现了类似的磷酸盐化化石生物群。保康化石组合中包括多细胞藻类Wengania,Paramecia和原地保存的Thallophyca;大型复杂疑源类Meghystrichosphaeridium,Echinosphaeridium和目前仅在保康发现的Bacatisphaera;球状蓝菌化石Archaeophycus,丝状蓝菌化石Siphonophycus和在陡山沱组磷块岩中首次发现的Obruchevella;同时,内部保存不同数量小球体的球状化石在保康也大量产出。古地理分析表明中国南方扬子地台陡山沱组磷块岩中保存的磷酸盐化化石库大都形成于浅水台地边缘地带。     

疑源类Tianzhushania在贵州瓮安震旦纪陡山沱组磷块岩中的发现

首次报道贵州瓮安地区震旦系陡山沱组磷块岩中的大型具刺凝源类Tianzhushania spinosa及其新种Tianzhushania tuberifera sp.nov.前者形态特征及内部结构与湖北峡尔地区震旦系陡山沱组隧石结核中发现的典型Tianzhushania spinosa 标本（硅化）完全相同,但多数未遭受重力挤压呈球形保存。新种特征明显,也呈球形保存,但内层表面具有明显瘤突,瘤突顶端向外延伸出单一或多条管状制,另外还探讨大型具刺疑源类Tianzhushania属的特征,已定种的地理分布,地层意义和亲缘关系。     

Precambrian animal life: Taphonomy of phosphatized metazoan embryos from southwest China

Phosphatized fossils from the Neoproterozoic Doushantuo Formation have provided valuable insight into the early evolution of metazoans, but the preservation of these spectacular fossils is not yet fully understood. This research begins to address this issue by performing a detailed specimen-based taphonomic analysis of the Doushantuo Formation phosphatized metazoan embryos. A total of 206 embryos in 65 thin sections from the Weng'an Phosphorite Member of the Doushantuo Formation were examined and their levels of pre-phosphatization decay estimated. The data produced from this examination reveal a strong taphonomic bias toward earlier (2-cell and 4-cell) cleavage stages, which tend to be well-preserved, and away from later (8-cell and 16-cell) cleavage stages, which tend to exhibit evidence for slight to intense levels of organic decay. In addition, the natural abundances of these embryos tend to decrease with advancement in cleavage stage, and no evidence of more advanced (beyond 16-cell) cleavage stages or eventual adult forms were found in this study. One possible explanation for this taphonomic bias toward early cleavage stages is that later cleavage stages and adult forms were more physically delicate, allowing them to be more easily damaged during burial and reworking, allowing for more rapid decay. The spectacular preservation of these embryos was probably aided by their likely internal enrichment in phosphate-rich yolk, which would have caused their internal dissolved phosphate levels to reach critical levels with only miniscule organic decay, thereby hastening phosphatization. If internal sources of phosphate did indeed play a role in the phosphatization of these embryos, it may explain their prolific abundance in these rocks compared to other phosphatized fossils as well as indicating that metazoans lacking such internal phosphate sources were likely much more difficult to preserve. The phosphatic fossils of the Doushantuo Formation, therefore, provide an indispensable, yet restricted, window into Neoproterozoic life and metazoan origins.     

Wengania globosa在陕西勉县茶店陡山沱组磷块岩中的发现

本文报道陕西勉县茶店陡山沱组磷块岩中的疑源多细胞生物化石,其中Ｗｅｎｇａｎｉａｇｌｏｂｏｓａ属于首次报道。勉县茶陡山组磷块岩中的化石组合与贵州瓮安陡山沱组与可以直接对比,进一步了此两地的磷块岩属同期沉积产物。这一发现表明新元古代磷块岩中可能保存了多细胞生物早期演化中最精采的章节,因而进一步深入研究陡山沱组磷块岩大有必要。本文还讨论了疑在Ｍｅｇｈｙｓｔｒｉｃｈｏｓｐｈａｅｒｉｄｉｕｍ ｃｈａｄｉａｎ     

扬子区晚震旦世动物化石新材料

扬子区晚震旦世地层中已有多种类型的动物及遗迹化石的报道。文中补充了不同地区和层位的骨骼及遗迹化石新材料,包括：江西上饶王家院陡山沱组顶部磷块岩中的磷酸盐化“十”字形骨针和云南晋宁王家湾灯影组上段隧石条带中的单轴双射硅质骨针,它们可能分属于钙质海绵和普通海绵类;贵州瓮安陡山沱组上段磷质白云岩中的可疑壳状化石;宜昌三斗坪金瓜墩陡山沱组下部（第二段）遗迹化石;瓮安陡山沱组上段及湖北南漳灯影组上段底部磷质硬底中的生物扰动构造。另外还报道了贵州清镇桃子冲剖面桃子冲组最底部的瓶状动物（Pro-tolagena limbata）化石,并认定桃子冲组与灯影组之间为假整合接触,其底界并无跨入震旦系的证据。     

贵州瓮安陡山沱组磷块岩中疑源类化石壳壁超微结构

未经化学处理的原始岩石样品的扫描电镜观察,确定了瓮安陡山沱组型刺饰疑源类Meghystri chospaeridium chadianensis的微结构特征。磷酸盐化疑源类化石由表面棘刺和球体组成,球体分为壳壁和腔体二部分。球体壳壁具黑色外层和白色内层的多层结构。棘刺具有与球体相似的多层状的壳壁和腔体。首次发现了发育于球体表面和体壳壁内部的网格超微结构。球体的腔体内存在的残余有机质。     

Preservational modes of some ichthyosaur soft tissues (Reptilia,Ichthyopterygia) from the Jurassic Posidonia Shale of Germany

Konservat-Lagerstätten, such as the Toarcian (Early Jurassic) Posidonia Shale of southwestern Germany, are renowned for their spectacular fossils. Ichthyosaur skeletons recovered from this formation are frequently associated with soft tissues; however, the preserved material ranges from three-dimensional, predominantly phosphatized structures to dark films of mainly organic matter. We examined soft-tissue residues obtained from two ichthyosaur specimens using an integrated ultrastructural and geochemical approach. Our analyses revealed that the superficially-looking ‘films’ in fact comprise sections of densely aggregated melanosome (pigment) organelles sandwiched between phosphatized layers containing fibrous microstructures. We interpret this distinct layering as representing condensed and incompletely degraded integument from both sides of the animal. When compared against previously documented ichthyosaur fossils, it becomes readily apparent that a range of preservational modes exists between presumed ‘phosphatic’ and ‘carbonized’ soft-tissue remains. Some specimens show high structural fidelity (e.g. distinct integumentary layering), while others, including the fossils examined in this study, retain few original anatomical details. This diversity of soft-tissue preservational modes among Posidonia Shale ichthyosaurs offers a unique opportunity to examine different biostratinomic, taphonomic and diagenetic variables that potentially could affect the process of fossilization. It is likely that soft-tissue preservation in the Posidonia Shale was regulated by a multitude of factors, including decay efficiency and speed of phosphatic mineral nucleation; these in turn were governed by a seafloor with sustained microbial mat activity fuelled by high organic matter input and seasonally fluctuating oxygen levels.     

现代丰年虾卵的腐解实验——对早期化石胚胎研究的启示

早寒武世和埃迪卡拉纪中的球状化石,一些已被归入可能的后生动物胚胎化石,由于具较为完好的三维保存方式以及近乎完美的胚胎发育序列,为早期后生动物的起源、分类、谱系演化及发育生物学提供了难得的实证材料。然而随着研究的深入,多数寒武纪胚胎的生物学分类位置未定;而数量异常巨大、又有独自的保存方式的晚元古代陡山沱组胚胎的真伪和生物学归属,更是争议未消。通过对现生生物胚胎的实验埋藏研究,可以揭示出各类生物胚胎在腐解、埋藏各阶段的保存潜力,而现代胚胎在各实验埋藏阶段形态、结构的变化,也能为化石胚胎的研究提供重要的实证材料。本文就是通过对虾卵胚胎各发育阶段腐解保存潜力的实验模拟研究,试图为球状化石的形成机制和化石归属提供一些实验室依据。     

寒武纪早期宽川铺生物群中圆管螺化石显微结构及显微谱学分析

陕南寒武系底部宽川铺组不仅保存了大量的小壳化石,还以磷酸盐化的方式立体保存了多种软躯体动物的胚胎和成体化石,为研究早期动物矿化模式、个体发育方式以及动物躯体构型的早期辐射过程提供了非常关键的信息。然而宽川铺生物群的研究大多聚焦于化石系统分类及亲缘关系探索,对生物群的沉积背景、埋藏过程和保存模式的分析甚少。本文以宽川铺生物群的核心产出层位——陕西省西乡县大河镇宽川铺组下部磷质碎屑灰岩为研究对象,对其中的一类管状化石圆管螺及其围岩进行了显微结构学和显微谱学综合分析,获得了化石显微结构和关键结构对应的元素、矿物成分等信息,讨论了管状化石圆管螺的埋藏过程和保存模式,并在此基础上对其生物学特征进行了初步探讨。该研究对理解宽川铺生物群的埋藏机制提供了新的线索。     

Experimental taphonomy of giant sulphur bacteria: implications for the interpretation of the embryo-like Ediacaran Doushantuo fossils

The Ediacaran Doushantuo biota has yielded fossils interpreted as eukaryotic organisms, either animal embryos or eukaryotes basal or distantly related to Metazoa. However, the fossils have been interpreted alternatively as giant sulphur bacteria similar to the extant Thiomargarita. To test this hypothesis, living and decayed Thiomargarita were compared with Doushantuo fossils and experimental taphonomic pathways were compared with modern embryos. In the fossils, as in eukaryotic cells, subcellular structures are distributed throughout cell volume; in Thiomargarita, a central vacuole encompasses approximately 98 per cent cell volume. Key features of the fossils, including putative lipid vesicles and nuclei, complex envelope ornament, and ornate outer vesicles are incompatible with living and decay morphologies observed in Thiomargarita. Microbial taphonomy of Thiomargarita also differed from that of embryos. Embryo tissues can be consumed and replaced by bacteria, forming a replica composed of a three-dimensional biofilm, a stable fabric for potential fossilization. Vacuolated Thiomargarita cells collapse easily and do not provide an internal substrate for bacteria. The findings do not support the hypothesis that giant sulphur bacteria are an appropriate interpretative model for the embryo-like Doushantuo fossils. However, sulphur bacteria may have mediated fossil mineralization and may provide a potential bacterial analogue for other macroscopic Precambrian remains.     

Three-Dimensional Phosphatic Preservation Of Giant Acritarchs From The Terminal Proterozoic Doushantuo Formation In Guizhou And Hubei Provinces, South China

During the last two decades, a diverse assemblage of multicellular algae, spherical microfossils, bacteria and cyanobacteria (the Weng'an Biota) has been reported from the Terminal Proterozoic Doushantuo phosphorite in south China, which provides an important window for evolutionary studies of multicellular life just before the Ediacaran animal radiation. In this paper we report on three-dimensionally preserved acritarchs from the Doushantuo phosphorites at Weng'an in Guizhou Province and, for the first time, from Baokang in Hubei Province. Our assemblage contains species of the genus Meghystrichosphaeridium Zhang, Yin, Xiao and Knoll, 1998 non Chen and Liu, 1986 including M. chadianensis (Chen and Liu) Zhang, Yin, Xiao and Knoll, emend. 1998, and M. reticulatum Xiao and Knoll, 1999; the genus Echinosphaeridium Knoll, 1992 including the species E. maximum (Yin) Knoll, 1992; the new genus Bacatisphaera including the new species B. baokangensis and the new genus Castaneasphaera including the new species C. speciosa . The latter shows a marked resemblance to similarly preserved phosphatized Palaeozoic acritarchs called 'mazuelloids', consistent with a phytoplanktonic mode of life for the global Doushantuo/Pertatataka microflora (DPM).     

A new look at saponite formation and its implications for early animal records in the Ediacaran of South China

Acanthomorphic acritarch fossils, including some interpreted to be the fossils of the earliest animal embryos, first appear in the lower Doushantuo Formation of the Yangtze Gorges area (YGA). Further, the complete paleontological and geochemical record for the YGA has played a central role in defining the global biological and geochemical backdrop that presaged and witnessed the dawn of diverse animal life. Despite the importance of the YGA in our understanding of Neoproterozoic Earth history, basic aspects about its depositional history remain debated. Foremost among the controversies, extensively studied sections in the YGA were recently tied to deposition in an alkaline lake, casting new but contentious light on the environments of early animal evolution and the broader significance of geochemical records from the YGA. Arguments for a lacustrine setting hinged on the presence of trioctahedral clays (saponite–corrensite). However, this clay type commonly forms in other environments, including the weathering profiles of mafic and ultramafic volcanics. Using a coupled geochemical and sedimentological approach, we argue that the trioctahedral clays in the lower Doushantuo of the YGA are better explained as weathering products from a regional mafic‐to‐ultramafic hinterland delivered by rivers to a shelf or lagoon in the Yangtze Gorges Basin. These novel provenance relationships for YGA sediments and associated clays are consistent with a marine setting for the early animal records and must factor in our current understanding of the broader geochemical fabric of the Doushantuo Formation.     

贵州瓮安新元古代陡山沱组磷酸盐化管柱状微体化石新材料

在用醋酸浸泡处理采自贵州瓮安新元古代陡山沱组磷酸质岩石样品时,获得大量微体动物骨骼化石。文中重点报道部分管柱状微体化石,即：Sinoquadraticus poratus gen.et sp.nov和Sinoquadraticus wenganensis gen. et sp.nov.。并对Sinocyclocylicus的部分特征进行补充。     

Phosphatized multicellular algae in the Neoproterozoic Doushantuo Formation, China, and the early evolution of florideophyte red algae

Phosphatic sediments of the Late Neoproterozoic (ca. 600 million years old [Myr]) Doushantuo Formation at Weng'an, South China, contain fossils of multicellular algae preserved in anatomical detail. As revealed by light microscopy and scanning electron microscopy, these fossils include both simple pseudoparenchymatous thalli with apical growth but no cortex-medulla differentiation and more complex thalli characterized by cortex-medulla differentiation and structures interpretable as carposporophytes, suggesting a multiphasic life cycle. Simple pseudoparenchymatous thalli, represented by Wengania, Gremiphyca, and Thallophycoides, are interpreted as stem group florideophytes. In contrast, complex pseudoparenchymatous thalli, such as Thallophyca and Paramecia, compare more closely to fossil and living corallinaleans than to other florideophyte orders, although they also differ in some important aspects (e.g., lack of biocalcification). These more complex thalli are interpreted as early stem group corallinaleans that diverged before Paleozoic stem groups such as Arenigiphyllum, Petrophyton, Graticula, and Archaeolithophyllum. This phylogenetic interpretation implies that (1) the phylogenetic divergence between the Florideophyceae and its sister group, the Bangiales, must have taken place before Doushantuo time-an inference supported by the occurrence of bangialean fossils in Mesoproterozoic rocks; (2) the initial diversification of the florideophytes occurred no later than the Doushantuo time; and (3) the corallinalean clade had a "soft" (uncalcified) evolutionary history in the Neoproterozoic before evolving biocalcification in the Paleozoic and undergoing crown group diversification in the Mesozoic.     

陕西西乡寒武纪幸运期宽川铺生物群网格状微体化石埋藏学与生物属性研究*

磷酸盐化保存是软躯体化石特异埋藏的一个重要途径, 而微生物在软躯体磷酸盐化过程中可能发挥了重要作用。前人通过埋藏学实验发现, 微生物会在动物胚胎等软躯体组织内部快速滋生, 充填生物体内部空间, 以微生物假形的方式复制了生物体的原始形态。但化石的磷酸盐化过程是否与埋藏学实验模拟的过程一致, 目前仍有争议。本次研究在寒武纪早期宽川铺生物群中发现了一类网格状微体化石。此类化石的保存状态可以根据其中丝状微生物滋生的程度分为三种类型, 它们展示了生物从死亡到微生物侵入、滋生, 最后被磷酸盐化的全过程。这些标本显示, 微生物假形在生物软组织磷酸盐化过程中扮演了重要角色, 但并不是化石磷酸盐化的必由之路, 尤其是当生物体具有矿化硬骨骼或者几丁质软骨骼等抗腐性较强的结构时。此类标本多以不完整保存的残片为主, 正反两面结构一致, 具有典型列状排列的近圆形与哑铃形网孔。由于化石结构简单, 生物学性状较少, 因此它们的亲缘关系尚不明确, 是一类需要继续研究的疑难化石。     

Distinguishing geology from biology in the Ediacaran Doushantuo biota relaxes constraints on the timing of the origin of bilaterians

The Ediacaran Doushantuo biota has yielded fossils that include the oldest widely accepted record of the animal evolutionary lineage, as well as specimens with alleged bilaterian affinity. However, these systematic interpretations are contingent on the presence of key biological structures that have been reinterpreted by some workers as artefacts of diagenetic mineralization. On the basis of chemistry and crystallographic fabric, we characterize and discriminate phases of mineralization that reflect: (i) replication of original biological structure, and (ii) void-filling diagenetic mineralization. The results indicate that all fossils from the Doushantuo assemblage preserve a complex mélange of mineral phases, even where subcellular anatomy appears to be preserved. The findings allow these phases to be distinguished in more controversial fossils, facilitating a critical re-evaluation of the Doushantuo fossil assemblage and its implications as an archive of Ediacaran animal diversity. We find that putative subcellular structures exhibit fabrics consistent with preservation of original morphology. Cells in later developmental stages are not in original configuration and are therefore uninformative concerning gastrulation. Key structures used to identify Doushantuo bilaterians can be dismissed as late diagenetic artefacts. Therefore, when diagenetic mineralization is considered, there is no convincing evidence for bilaterians in the Doushantuo assemblage.     

陕南寒武系下部宽川铺组帽球状化石*

陕南寒武纪早期宽川铺组微体球状化石类型多样,如何正确辨别这些球状化石的生物属性是当前早期生命演化古生物学研究中的一个难点。我们在宽川铺组中发现了一类为数众多,具有一个或者多个不规则的帽状隆起结构以及翻边帽沿的帽状化石。研究表明这些帽状化石均为不完整的个体,其完整形态呈不规则的凹球形。根据帽球状化石帽身的数量和相对位置,这类化石可划分为"单帽型"、"双帽型"和"复帽型"三种类型。这些凹球状化石呈双层壳壁,内壁光滑,外壁粗糙。因为凹球状化石形态及其表面小孔与微体藻类、后生动物的胚胎以及壳体化石差异都非常显著,所以推测这类化石可能与带壳原生动物亲缘关系最为紧密。     

陕南早寒武世宽川铺生物群沉积环境及埋藏机制研究

陕南早寒武世宽川铺生物群三维立体保存了大量结构精美的磷酸盐化胚胎及小壳化石,为后生动物演化及化石胚胎学的研究提供了珍贵的实证材料。目前的研究主要集中在化石分类学及演化生物学上,而对沉积环境及埋藏机制的研究却很少涉及。文中通过对大量野外岩石照片、岩石薄片、小壳化石SEM照片的综合研究后认为,宽川铺生物群不是原地埋藏的产物,而是事件沉积的结果。该研究对于探讨宽川铺生物群化石的埋藏环境及保存机制具有一定的意义。     

陕西宁强早寒武世磷酸盐化具星状表饰胚胎化石研究

后生动物胚胎化石研究对了解寒武纪大爆发中生命的个体及系统发育具有十分重要的意义。陕南早寒武世灯影组宽川铺段发现大量磷酸盐化的球状化石标本,其中的一些为保存有十分精细的星状表饰的胚胎化石,目前发现至少3种具星状纹饰的胚胎化石:圆球状胚胎、具中央收缩带的球状胚胎和表面具“拉伸构造”的花生状胚胎,其中后者为新类型。研究表明,此新类型在演化序列上处于圆球状胚胎、具中央收缩带的球状胚胎发育阶段之后,是胚胎向成体演化的一个重要中间过渡环节,这一发现为研究早寒武世早期后生动物胚胎及个体发育提供了新线索。