The importance of Planolites in the Cambrian substrate revolution

Early Cambrian subtidal shelf substrates were characterized by low water content and steep chemical gradients, conditions likely facilitated by the presence of microbial mats as reflected by an abundance of microbially-mediated sedimentary structures in Lower Cambrian strata. Such substrate conditions would have been unfavourable for burrowing by benthic metazoans. A combination of environmental restrictions and a lack of adaptations to vertical burrowing likely prevented most benthic metazoans from burrowing infaunally in Early Cambrian subtidal shelf substrates. The eventual acquisition of burrowing adaptations by benthic metazoans later in the Cambrian promoted an increase in the depth and intensity of bioturbation and initiated a transition toward well-hydrated substrates in which extensive infaunal activity was possible.Siliciclastic units of the Lower Cambrian succession in the White–Inyo Mountains, eastern California, contain abundant horizontal bioturbation on bedding planes, as documented by bedding plane bioturbation indices, but little vertical bioturbation, as shown by ichnofabric indices and x-radiography. Planolites, a simple horizontal trace fossil, represents the dominant type of bioturbation in these units. Planolites is found in a range of diameters, indicating that more than one species of tracemaker likely produced this type of trace. Although these Planolites do not have a vertical component, their abundance on bedding planes indicates that the activities of Planolites tracemakers had a significant impact on subtidal shelf substrates, represented by Lower Cambrian units in the White–Inyo Mountains, early in the Cambrian substrate revolution.     

Paleoecology of a large Early Cambrian bioturbator

The Lower Cambrian Poleta Formation in the White-Inyo Mountains of eastern California contains well-preserved and laterally extensive exposures of the large looping and meandering trace fossil Taphrhelminthopsis nelsoni n.isp. Such traces are typical features on upper bed surfaces of Lower Cambrian shallow marine sandstones and occur with Ediacaran fossils at other localities. Morphologic, sedimentologic and goniogram analyses suggest that the inferred tracemaker was a large soft-bodied echinozoan- or mollusc-grade animal with a volume greater than 14 cm³ that actively grazed or ingested sediment at the sediment-water interface. Although portions of these traces appear to reflect relatively 'complex' behavior, looping patterns are not periodic as expected for a systematic foraging strategy. T. nelsoni traces are patchy in distribution and commonly associated with suspect-microbial features, suggesting that tracemakers may have been targeting microbial-based or related concentrations of food resources. Such behavioral patterns are typical of shallow late Neoproterozoic-early Cambrian settings, and like suspect-microbial structures are later restricted to deep marine or stressed settings.     

Facies control on lower Cambrian wrinkle structure development and paleoenvironmental distribution, southern Great Basin, United States

Assessing the role that physical processes play in restricting microbial mat distribution has been difficult due to the primary control of bioturbation in the modern ocean. To isolate and determine the physical controls on microbial mat distribution and preservation, a time in Earth’s history must be examined when bioturbation was not the primary control. This restricts the window of observation primarily to the Precambrian and Cambrian, which precede the development of typical Phanerozoic and modern levels of bioturbation. Lower Cambrian strata of the southern Great Basin, United States, record the widespread development of seafloor microbial mats in shallow shelf and nearshore settings. These microbial mats are recorded by wrinkle structures, which consist of millimeter-scale ridges and sinuous troughs that represent the former presence of a surface microbial mat. Wrinkle structures within these strata occur exclusively within heterolithic deposits of the offshore transition, i.e., between fair-weather wave base and storm wave base, and within heterolithic tidal-flat deposits. Wrinkle structures are not preserved in siltstone-dominated offshore deposits or amalgamated shoreface sandstones. The preservation of wrinkle structures within these environments is due to: (1) the development of microbial mats atop clean quartz-rich sands for growth and casting of the structures; and (2) the draping of the microbial mat by finer-grained sediment to inhibit erosion. The exclusion from offshore deposits may be due to a lack of sufficient sunlight, whereas the restriction from the shoreface is likely due to the amalgamation of proximal tempestites, resulting in the erosion of any incipient microbial mat development.     

Quantitative paleoecology of Cambrian (Series 2–Miaolingian) communities from central Sonora,Mexico

This work presents a paleoecological approach to study benthic communities preserved in Cambrian strata exposed at the Sahuaral locality in central Sonora, northwestern Mexico. Based on species richness and abundance, this work analyzed 627 individuals corresponding to 33 species from five distinctive Cambrian formations: the lowermost Proveedora Formation characterized by bioturbation horizons that record the occurrence of a benthic-infaunal biota from coastal shallow seas, followed by the Buelna Formation which contains reef deposits dominated by autotrophic epibenthic communities. The Cerro Prieto Formation was deposited in a high-energy shallow marine environment, dominated by primary consumers-epifaunal and suspensions feeders. The uppermost unit, the El Gavilán Formation, is an intercalation of shale and fossiliferous limestone characterized by the fossil remains of suspension feeders, detritus feeders and nektobenthic organisms of low energy deep water environments. The paleobiota consist of ichnofossils, microbial oncolites, chancellorids, brachiopods, hyolithids, and trilobites in a sequence of increasing diversity and complexity. Paleoecological indices (Simpson Dominance, Shannon-Weaver Index, Evenness Index, and Total Richness) were determined and compared in the study area. In addition, the effective number of species and the true diversities were obtained, which are similar between each station distributed in each formation. However, it is possible that the high degree of dominance is a factor to indicate an inequality between the true diversity and the species richness identified. These results provide new analytical and paleoecological criteria for better understanding depositional environments and fossil associations where the spatial distribution of benthic organisms, richness and abundance provide a valued window to Cambrian biotas. The structure of the marine community of the El Sahuaral area has a great affinity with other Cambrian communities, such as those from the Stephen Formation (Burgess Shale, British Columbia) and the Chengjiang biota of the Yangtze Shelf deposits in South China.     

Phosphatized rare star-like mouth disc of Punctatus and its functional morphology from the earliest Cambrian of the South Shaanxi China

The Meishucun stage is the prelude in decipher-ing the Cambrian Explosion. In this prominent stage, rapid radioactive evolution and body-plan innovation have taken place and different associations of organism have been shaped. In this paper we report several 3D-preserved rare star-like fossils with finely preserved soft tissues which were recovered from the Kuanchuanpu Member of the Dengying Formation in South Shaanxi, China in 2003. By studying on functional morphology and analogy with mouthpart of Punctatus, there are evidences that this star-like organism approaches the coelenterates in systematic classification and the centre of star-like organism is its mouth. The appearance of coelenterates marks the real beginning of metazoan evolution. Therefore, it has the prominent position in the origin and evolutionary history of organisms. Perhaps the star-like organism represents the early types of coelenterate with original tentacles. These new materials provide new evidence for the origin, evolution and the functional evolution of the metazoan during the early stage of the Cambrian Explosion.     

陕南早寒武世早期Quadrapyrgites再研究

处于"寒武纪生命大爆发"序幕阶段的梅树村期,生物类群大规模辐射,身体构型快速革新,与前寒武纪生物群面貌明显不同.最近在陕南宁强宽川铺地区梅树村期地层中发现了大量五辐射Punctatus及部分四辐射四方塔形壳属Quadrapyrgites,其中包括1个新种Quadrnpyrgites undulatuscostalis sp.nov..在此基础上对Quadrapyrgites进行了属征补充.双胚层腔肠动物的出现标志着地球生命史的真后生动物演化开端,在生物起源演化历程上占据着关键的位置.本文为研究真后生动物起源演化、生物辐射、体型构建提供了重要实证.     

Trace fossils in the Ediacaran-Cambrian transition: Behavioral diversification, ecological turnover and environmental shift

After taxonomic revision, trace fossils show a similarly explosive diversification in the Ediacaran-Cambrian transition as metazoan body fossils. In shallow-marine deposits of Ediacaran age, trace fossils are horizontal, simple and rare, and display feeding strategies related to exploitation of microbial matgrounds. Equally notable is the absence of arthropod tracks and sinusoidal nematode trails. This situation changed in the Early Cambrian, when a dramatic increase in the diversity of distinct ichnotaxa is associated was followed by the onset of vertical bioturbation and the disappearance of a matground-based ecology (‘‘agronomic revolution’’). On deep sea bottoms, animals have been present already in the Ediacaran, but ichnofaunas were poorly diverse and dominated by the horizontal burrows of undermat miners. As shown by the ichnogenus Oldhamia, this life style continued to be predominant into the Early, and to a lesser extent, Middle Cambrian. Nevertheless, there was an explosive radiation of behavioral programs during the Early Cambrian. When exactly the bioturbational revolution arrived in the deep sea is uncertain. In any case, the Nereites ichnofacies was firmly established in the Early Ordovician. The rich ichnofauna in the Early Cambrian Guachos Formation of northwest Argentina probably marks a first step in this ecological onshore-offshore shift.     

陕南早寒武世早期磷酸盐化Punctatus奇异星状口盘的发现及其形态功能分析

早寒武世最早期梅树村期处于“寒武纪大爆发”序幕阶段。在这一时期,生物类群发生了大规模辐射演化及其身体构型的快速革新,形成与前寒武纪生物群明显不同的生物组合面貌。最近在陕南宽川铺地区早寒武世早期灯影组宽川铺段地层中发现了数十枚呈三维立体精美保存的磷酸盐化奇异星状生物化石标本,通过形态功能分析及与其共生的Punctatus的口盘类比表明,该奇异星状生物很可能属于腔肠动物Punctatus的口盘,星状体的中心为该生物的口部。而腔肠动物的出现标志着真后生动物的开始,在生物起源演化历程上占据着极其关键的位置。本文报道的奇异星状生物可能代表了初具原始触手的腔肠动物早期演化类型,为研究真后生动物起源演化、功能进化提供了新的实证材料。     

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.     

Ichnology of the Winnipeg Formation,southeast Saskatchewan: a glimpse into the marine infaunal ecology of the Great Ordovician Biodiversification Event

The ichnology of the Middle Ordovician Winnipeg Formation has been analysed based on the study of cores from five wells drilled in southeast Saskatchewan (Canada). Six sedimentary facies, ranging from upper shoreface to lower offshore settings in a shallow‐marine environment, have been characterized. Ichnological attributes are consistent with those in currently proposed models for shallow‐marine wave‐dominated settings, but ichnodiversity is lower than in post‐Palaeozoic settings. Low ichnodiversity in the Winnipeg Formation most likely reflects evolutionary factors rather than environmental controls. Interestingly, low‐energy, distal deposits of the Winnipeg Formation display intense degree of bioturbation, reflecting a well‐developed mixed layer and underscoring the importance of the Great Ordovician Biodiversification Event in terms of sediment mixing.     

河南叶县下寒武统辛集组单板类和腹足类化石的研究

本文描述和讨论了河南叶县杨寺庄下寒武统辛集组单板类１３种,腹足类２种,未定类１种。其中包括２新属６新种：Ｒｅｐｅｎｏｃｏｎｕｓ ｘｉｎｊｉｅｎｓｉｓ ｇｅｎ．ｎｏｖ．,Ｓｃｅｎｅｌｌａ ｐｙｃｎａ ｓｐ．ｎｏｖ．,Ｓｅｃｕｒｉｃｏｎｕｓ ｖｕｌｇａｒｉｓ ｓｐ．ｎｏｖ．,Ｏｂｔｕｓｏｃｎｕｓ ｇｒｏｓｓｉｃｏｓｔｕｓ ｓｐ．ｎｏｖ．,Ｉｇｏｒｅｌｌｉｎａ ｐｒｏｂｏｓｃｉｓ ｓｐ．ｎｏｖ．,Ｇａｌ     

Trace fossil analysis of lacustrine facies and basins

Two ichnofacies are typical of lacustrine depositional systems. The Scoyenia ichnofacies characterizes transitional terrestrial/nonmarine aquatic substrates, periodically inundated or desiccated, and therefore is commonly present in lake margin facies. The Mermia ichnofacies is associated with well oxygenated, permanent subaqueous, fine-grained substrates of hydrologically open, perennial lakes. Bathymetric zonations within the Mermia ichnofacies are complicated by the wide variability of lacustrine systems. Detected proximal–distal trends are useful within particular lake basins, but commonly difficult to extrapolate to other lakes. Other potential ichnofacies include the typically marine Skolithos ichnofacies for high-energy zones of lakes and substrate-controlled, still unnamed ichnofacies, associated to lake margin deposits. Trace fossils are useful for sedimentologic analysis of event beds. Lacustrine turbidites are characterized by low-diversity suites, reflecting colonization by opportunistic organisms after the turbidite event. Underflow current beds record animal activity contemporaneous with nearly continuous sedimentation. Ichnologic studies may also help to distinguish between marine and lacustrine turbidites. Deep-marine turbidites host the Nereites ichnofacies that consists of high diversity of ornate grazing traces and graphoglyptids, recording highly specialized feeding strategies developed to solve the problem of the scarcity of food in the deep sea. Deep lacustrine environments contain the Mermia ichnofacies, which is dominated by unspecialized grazing and feeding traces probably related to the abundance and accessibility of food in lacustrine systems. The lower diversity of lacustrine ichnofaunas in comparison with deep-sea assemblages more likely reflects lower species diversity as a consequence of less stable conditions. Increase of depth and extent of bioturbation through geologic time produced a clear signature in the ichnofabric record of lacustrine facies. Paleozoic lacustrine ichnofaunas are typically dominated by surface trails with little associated bioturbation. During the Mesozoic, bioturbation depth was higher in lake margin facies than in fully lacustrine deposits. While significant degrees of bioturbation were attained in lake margin facies during the Triassic, major biogenic disruption of primary bedding in subaqueous lacustrine deposits did not occur until the Cretaceous.     

云南澄江早寒武世地层并论中国的Parabadiella,摩洛哥的Abadiella及澳大利亚的Abadiella huoi

根据近来的发现,区域地层的阐明及生物在寒武系界线上下的变化,澄江早寒武世地层划分作了修订。早寒武世岩石地层名称的更换,年代地层及生物地层单位的分带这里作了简述.目前的岩石地层单位从上到上是渔户村组（梅树阶）,黑林铺组（筇竹寺阶）,红井哨组、乌龙箐组（沧浪铺阶）及山邑村组（龙王庙阶）。此层序可分为2个亚带。所有的组合带在中国西南地台区都有存在,但两个亚带在澄江是区域性的。云南东部震旦系与寒武系的界线暂时置于渔户村且小歪头山段的底界,位于小壳化石最早出现的下部。中国的Parabadiella,摩洛哥的Abadiella及澳大利亚的Abadiella huoi这里重新作了研究。Parabadiella及Abadiella是两个不同的属。模式标本的图片在这里再次刊出。澳大利亚的Abadiella huoi一种应是Wutingaspis的一个新种（Wutingaspis jelli Chang sp.nov)。Opik(1975）的Dolerolenus(?)sp.nov.一种应归属Parabadiella huoi一种。Parabadiella是中国也是澳大利亚寒武系最古老的三叶虫。澳大利亚的Wutingaspis jelli的层位与摩洛哥Abadiella的层位相当,可与中国的Wutingaspis-Eoredichia带的上部对比。     

A scratch circle origin for the medusoid fossil Kullingia

Kullingia is considered a key taxon in demonstrating the presence of terminal Proterozoic-early Cambrian chondrophorine hydrozoans. However, Kullingia concentrica from the Lower Cambrian of northern Sweden possesses several features that show that it is not a body fossil but that it was formed by current or wave-induced rotation of an anchored tubular organism, possibly a sabelliditid. A scratch circle interpretation applies also to several other reports of Lower Cambrian Kullingia , including Kullingia delicata from the Chapel Island Formation of Newfoundland. Given the considerable number of problematic fossils that have been interpreted as chondrophorines, it is difficult to put an age on the oldest fossil chondrophorines, but it may be as late as the Devonian. Overall, scratch circles appear to be rarely preserved. The occurrence of these scratch circles in Lower Paleozoic storm deposits is probably related to low levels of bioturbation that enhanced both the likelihood of formation and preservation of these structures.     

Phosphatized rare star-like mouth disc of Punctatus and its functional morphology from the earliest Cambrian of the South Shaanxi China

The Meishucun stage is the prelude in deciphering the Cambrian Explosion. In this prominent stage, rapid radioactive evolution and body-plan innovation have taken place and different associations of organism have been shaped. In this paper we report several 3D-preserved rare star-like fossils with finely preserved soft tissues which were recovered from the Kuanchuanpu Member of the Dengying Formation in South Shaanxi, China in 2003. By studying on functional morphology and analogy with mouthpart of Punctatus, there are evidences that this star-like organism approaches the coelenterates in systematic classification and the centre of star-like organism is its mouth. The appearance of coelenterates marks the real beginning of metazoan evolution. Therefore, it has the prominent position in the origin and evolutionary history of organisms. Perhaps the star-like organism represents the early types of coelenterate with original tentacles. These new materials provide new evidence for the origin, evolution and the functional evolution of the metazoan during the early stage of the Cambrian Explosion. __________ Translated from Acta Micropalaeontologica Sinica, 2006, 23(1): 62–69 [译自: 微体古生物学报]     

Suspected microbial-induced sedimentary structures (MISS) in Furongian (Upper Cambrian; Jiangshanian,Sunwaptan) strata of the Upper Mississippi Valley

The Furongian (Upper Cambrian; Jiangshanian and Sunwaptan) Tunnel City Group (Lone Rock Formation and Mazomanie Formation), exposed in Wisconsin and Minnesota, represents a shallow-marine clastic environment during a time of exceptionally high sea level. Lithofacies from shoreface to transitional-offshore settings document deposition in a wave- and storm-dominated sea. Flooding of the cratonic interior was associated with formation of a condensed section and the extensive development of microbial mats. Biolamination, mat fragments, wrinkle structures, and syneresis cracks are preserved in various sandstone facies of the Lone Rock Formation, as is evidence for the cohesive behavior of sand. These microbial-induced sedimentary structures (MISS) provide unique signals of biological–physical processes that physical structures alone cannot mimic. The MISS are associated with a trilobite extinction event in the Steptoean–Sunwaptan boundary interval. This may support recent claims that Phanerozoic microbial mats were opportunistic disaster forms that flourished during periods of faunal turnover. Further investigation of stratigraphic, taphonomic, and other potential biases, however, is needed to fully test this hypothesis.     

Taphonomy of the middle Cambrian (Drumian) Ravens Throat River Lagerstätte,Rockslide Formation,Mackenzie Mountains,Northwest Territories,Canada

The Middle Cambrian (series 3, Drumian, Bolaspidella Biozone) Ravens Throat River Lagerstätte in the Rockslide Formation of the Mackenzie Mountains, northwestern Canada, contains a Burgess Shale‐type biota of similar age to the Wheeler and Marjum formations of Utah. The Rockslide Formation is a unit of deep‐water, mixed carbonate and siliciclastic facies deposited in a slope setting on the present‐day northwestern margin of Laurentia. At the fossil‐bearing locality, the unit is about 175 m thick and the lower part onlaps a fault scarp cutting lower Cambrian sandstones. It consists of a succession of shale, laminated to thin‐bedded lime mudstone, debris‐flow breccias, minor calcareous sandstone, greenish‐coloured calcareous mudstone and dolomitic siltstone, overlain by shallow‐water dolostones of the Broken Skull Formation, which indicates an overall progradational sequence. Two ~1‐m‐thick units of greenish calcareous mudstone in the upper part exhibit soft‐bodied preservation, yielding a biota dominated by bivalved arthropods and macrophytic algae, along with hyoliths and trilobites. It represents a low‐diversity in situ community. Most of the fossils occur in the lower unit, and only the more robust components are preserved. Branching burrows are present under the carapaces of some arthropods, and common millimetre‐sized disruptions of laminae are interpreted as bioturbation. The fossiliferous planar‐laminated calcareous mudstone consists of chlorite, illite, quartz silt, calcite and dolomite and is an anomalous facies in the succession. It was deposited via hemipelagic fallout of a mixture of platform‐derived and terrestrial mud. Geochemical analysis and trace‐element proxies indicate oxic bottom waters that only occasionally might have become dysoxic. Productivity in the water column was dominated by cyanobacteria. Fragments of microbial mats are common as carbonaceous seams. Complete decay of soft tissues was interrupted due to the specific sediment composition, providing support for the role of clay minerals, possibly chlorite, in the taphonomic process.     

Ecological drivers of the Ediacaran-Cambrian diversification of Metazoa

Organismal modifications to their physical and chemical environment play a significant role in structuring many modern ecosystems, and experimental evidence suggests that such behavior can increase diversity. Despite the important role such activities play in connecting ecology and evolution, less is known of the macroevolutionary impact of such influences, especially their role during major evolutionary transitions. The Ediacaran-Cambrian diversification of Metazoa encompassed the appearance and early diversification of virtually all major clades of marine animals and the establishment of metazoan-dominated ecosystems. Here we assess the role of positive ecological feedbacks using a new compilation of the first occurrences of all metazoan phyla, classes; orders and equivalent stem taxa, as well as data from a previously published compendium on fossils from the early to middle Cambrian of China. The results reveal relatively minor feedback during the Ediacaran, but a substantial increase during the Cambrian, principally through bioturbation and the appearance of a number of structural engineers, including sponges. Chemical modification of the environment through filtering and bioturbation seems to have had the largest impact. Data on taxic diversity is a poor proxy for abundance, or for the actual environmental impact of these activities, however. Future assessments of the influence of ecological feedbacks on this event will require standardized assessments of the abundance of taxa with different ecological roles.     

Secular distribution of Burgess-Shale-type preservation

Burgess-Shale-type preservation is defined as a taphonomic pathway involving the exceptional organic preservation of non-mineralizing organisms in fully marine siliciclastic sediments. In the Phanerozoic it occurs widely in Lower and Middle Cambrian sequences but subsequently disappears as a significant taphonomic mode. The hypothesis that this distribution derives solely from a secular increase in the depth of bioturbation is falsified: low bioturbation indices do not prevent the rapid enzymatic degradation of organic structure, nor do they account for the conspicuous absence of comparable preservation during the Vendian. An earlier, Late Riphean (ca. 750–850 Ma), interval of enhanced organic-walled fossil preservation suggests a long-term recurrence in Burgess-Shale-type taphonomy that is independent of metazoan activity. A model based on the potentially powerful anti-enzymatic and/or stabilizing effects of clay minerals on organic molecules is proposed to account for Burgess-Shale-type preservation. Long-term changes in average clay mineralogies and the ocean chemistry that determines their interaction with organic molecules are likely to have induced the pronounced secular distribution of these fossil biotas, while regional variations in tectonism, weathering, etc., explain their non-uniform geographic distribution; the close correlation between exceptional, organic-walled fossil preservation and volcano-genic sedimentation in Tertiary lake deposits provides a compelling analogue. Recognition of a temporal control on Burgess-Shale-type preservation constrains the evolutionary scenarios that can be drawn from such biotas; significantly, neither the initial rate of appearance, nor the ultimate fate of Burgess-Shale-type taxa can be directly assessed. □ Taphonomy, exceptional preservation, organic preservation, fossil Lagerstätten, Burgess Shale, clay mineralogy, clay-organic interactions, secular change, Cambrian, Proterozoic.     

河南登封寒武系第三统张夏组核形石与遗迹化石的耦合变化

河南登封关口剖面寒武系第三统张夏组下部碳酸盐岩中发育了大量的微生物成因的核形石和后生动物遗迹化石。在野外和显微镜下对核形石和遗迹化石进行观察,并统计它们在地层中所占比例,表明核形石和遗迹化石存在着耦合关系。下部地层以发育凝块石和形状不规则、纹层不连续、代表一种低能弱搅动水体的Ⅰ型核形石为主,不含遗迹化石,表明此时微生物在海洋生态系统中占主导地位。中部地层则以发育浑圆形、纹层连续的Ⅱ型核形石和遗迹化石Planolites为特征;而且随水体能量的增强,Planolites丰度逐渐升高,核形石丰度逐渐降低;二者的丰度变化说明后生动物的存在对核形石的数量有一定影响,但未破坏核形石的生长条件。上部地层发育大量Thalassinoides和生物扰动构造,缺乏核形石;后生动物对沉积基底进行反复扰动,彻底破坏了原始层理以及微生物造岩的环境,核形石消失。可见,在张夏组沉积时期,微生物与后生动物以及环境之间存在着特殊的相互作用关系。