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The Cambrian cnidarian Cambroctoconus orientalis occurs in clusters exhibiting pendent growth in crypts within thrombolite frameworks. Here we successfully reconstruct its three‐dimensional (3D) in situ mode of life. The 3D reconstruction shows the initial individuals of colonies and attachment structures, budding sites and growth modes of coralla in response to the extent of sheltered spaces. The earliest individuals were attached to the framework by a suction‐cup‐like structure and were basically orthogonal to the walls, growing downward. Their calicular bases were pierced by a hole. Individuals were initially cylindrical and gradually formed an octagonal column during growth. Budding occurred everywhere, and toroidal budding traces are abundant upon the corallite surfaces. The attachment structures were secondarily reinforced by continuous secretion from the coenenchymal tissue. Offsets continued to grow only when sufficient growth space was available, while producing as many individuals as possible. Growth of individuals was commonly halted by the surrounding framework. Growth directions were in part modified by corallite curvature to avoid collisions between corallites and to make full use of the intricate void spaces. In deteriorating conditions, offset corallites appeared through regeneration rather than budding from the remaining soft tissues. Active exploitation of the sheltered spaces within thrombolites, which remained dominant during the Cambrian, was thus conducted by C. orientalis. Notably, the cryptobionts treated herein adopted subtle growth strategies that have not previously been fully understood and are not easily interpreted with reference to organisms living in open spaces. [ABSTRACT FROM AUTHOR]

The Cambrian coral Cambroctoconus occurs selectively in the crypts of calcimicrobe Epiphyton reefs of the Zhangxia Formation (Miaolingian) in Shandong Province, North China. These cryptobionts preferentially grow laterally and/or downward, in some cases showing pendent growth from the ceilings of the reef framework. The upside‐down growth is clearly shown by the downward‐oriented aperture and the presence of fork‐like holdfasts at its base. The coral aggregations reveal two cryptic growth strategies: one involves the prevalence of far smaller individuals in vertically narrow spaces; the other involves the predominance of modular individuals in large cavernous spaces. Offset individuals frequently appear projecting downward and are connected to each other in a chain‐like fashion. Individuals often undergo rejuvenescence laterally and downward, when the growth directions may change. Therefore, the size and modularity of individuals and the direction of budding depend largely upon the cryptic microenvironments available at the time. The cryptobionts make the best use of vacant spaces by modifying the location of budding and altering their growth directions through rejuvenescence. The cryptic growth strategies found in Cambroctoconus indicate a certain degree of morphological variability in the modules. More importantly, they indicate that the individuals flexibly and exquisitely utilized the microbial crypts that were still predominant even during the middle Cambrian. These modes of cryptic growth demonstrate the age‐specific exploitation of niches by sessile skeletal organisms that was facilitated by the development of a firm attachment structure. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Peloids are major contributors to modern and ancient limestones, although their origins are not yet comprehensively understood. Upper Permian to Lower Triassic carbonate successions are well preserved at the Bangeng area in Guizhou Province, South China, where microbialites (thrombolites), coccoidal microbes and peloids are preserved together. These peloids are classified into three types (Peloid-A, B, C) that typically occur together along with coccoidal microbes and restricted bioclasts in a variety of microenvironments. Peloid-A is most common and exhibits close relationship with coccoidal microbes. Peloid-A1 has spherical grain and a size range typical for coccoidal microbes (10–60 μm in diameter), whereas Peloid-A2 is marked by larger spherical and subrounded grains with diffuse margins, ranging from 70 to 200 μm in diameter. Peloid-A1 is in part produced by the complete micritic filling in coccoidal microbes by their metabolic activities, whereas Peloid-A2 is related to the calcification of a colony of coccoidal microbes by their metabolic activities, or simply to the aggregation of individual peloids and coccoidal microbes. Peloids of this type might also originate from other carbonate precipitation processes. Peloid-B is derived merely by micrite filling in ostracodes and gastropods skeletons, whereas Peloid-C is of micritized-bioclast origin. These peloidal origins do not explain those of all environments and ages. Peloids are polygenetic in origin, irrespective of the presence of coccoidal microbes. However, it is emphasized that the peloids formed after the end-Permian extinction are dominated by microbes related ones. Those kinds of peloids might have occurred repeatedly and predominantly, especially in severely deteriorated environments such as those after the end-Permian mass extinction. [Copyright &y& Elsevier]

Stromatolites just after the end-Permian extinction are exceptionally well developed in the Chongyang area of Hubei Province, South China. The stromatolites include endolithic coccoidal microbes and exhibit three growth forms: columnar-layered, pseudo-columnar, and columnar, each located in different stratigraphic horizons. The columnar type is the most conspicuous and provides key information on the mode of their construction. In this paper, we describe the mega- to microscopic features of these stromatolites together with interpretations of their construction. The columnar type typically comprises alternations of lighter and darker laminae with intercalations of dense micritic laminae. The lighter laminae are composed of fine to coarse calcite cement and fine dolomite, whereas the darker laminae are composed of micrite and microspar. Endolithic coccoids are sporadic in these laminae. In contrast, the dense micritic laminae consist of micrite with abundant endolithic coccoids, accentuating the laminae of stromatolites. The columnar stromatolites were formed under deteriorating conditions after the end-Permian mass extinction as follows: (1) the alternating development of lighter and darker laminae due to the colonization of different microbial communities in lower-energy, shallow- to deep-subtidal settings; (2) carbonate precipitation within each microbial community through microbial metabolic activity and degradation of extracellular polymeric substances (EPS), leading to the alternations of lighter and darker laminae; (3) episodic interruption of the growth of the lighter and darker laminae by endolithic coccoidal activity; and (4) high levels of endolithic activity, accompanied by the precipitation of carbonate within boreholes to produce the dense micritic laminae. In contrast, the columnar-layered and pseudo-columnar types comprise alternations of lighter and darker laminae, composed mainly of fine to coarse calcite cement, fine dolomite, micrite, peloids, and microspar with sporadic endolithic coccoids. The dense micritic laminae are absent or rare in these stromatolites. These types were essentially produced by the repetition of processes (1) and (2) documented in the columnar type. The finely alternating laminae of these stromatolites are therefore the product of the repeated initiation and cessation of growth of relevant microbial communities in response to changing environmental conditions. The stromatolites were formed immediately after the end-Permian extinction and persisted intermittently throughout the Early Triassic. The Chongyang stromatolites are undoubtedly some of the best-developed stromatolites near the Permian–Triassic boundary and retain a high-resolution record of microbial successions and palaeo-oceanic perturbations immediately following the end-Permian extinction. [ABSTRACT FROM AUTHOR]