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Field aspect of Upper Hunghuayuan Formation at Zhangzhai. A, three reefs (outlined by dashed lines) surrounded by well-bedded limestone. Cystostroma has been found at AFO562 (see sample number in Fig. 1D). B, vertical surface of the lithistid sponge-Calathium reef at AFO562. Ruler = 5 cm. Ca, Calathium; Sp, lithistid sponge.
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Early Ordovician (early Floian) reefs of South China include lithistid sponge-Calathium reefs with a three-dimensional skeletal framework. These structures are among the first post-Cambrian skeletal-dominated reef structures and provides an opportunity to test how the novel metazoan builders changed the environments and increased topographic comple...
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... exposed at Zhangzhai, which can be subsumed under two broader categories ( Li et al. 2014): microbial-dominated (stromatolite and lithistid sponge-Calathium-calcimicrobial) reefs and metazoan-dominated (lithistid sponge-Calathium) reefs (Figs 1D, 2A). The lithistid sponge-Calathium reefs exhibit a high density of lithistid sponges and Calathium (Fig. 2B). Calathium fossils account for over 40% of the reef creating a striking frame-like cluster structure (Fig. 3A, B). Most Calathium indi- viduals are upright or slightly curved with a vase- shaped structure in longitudinal view (Fig. 3A). Lithistid sponges are also abundant reaching up to 20% in volume, but complete specimens are ...
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... This points to a shift of increased skeletal carbonate production within reefs by a diversity of organisms. By later Ordovician time, the sheet-like metazoan consortium was well established (Kroger et al., 2017), and corals were contributors to most reef ecosystems along with stromatoporoids (Cecile, 1988;Cooper et al., 2001;Li et al., 2017). There is also evidence that microbial fabrics become less abundant from Middle to later Ordovician time, with an overall increase in the skeletal framework builders in reefs (Webby, 2002). ...
... The records of early stromatoporoids clearly demonstrate that North China was the potential diversification centre during the Middle Ordovician (Jeon et al., 2017). After the earliest known appearance during the Early Ordovician in South China (Jeon et al., 2019;Q. J. Li et al., 2017), stromatoporoids subsequently diversified to 24 labechiid species in 10 genera in the Middle Ordovician of North China (Dong, 1982;Jeon et al., 2017Jeon et al., , 2019Jeon, Li, Chen et al., 2022;Ozaki, 1938;Sugiyama, 1941;Yabe & Sugiyama, 1930a, b). It is remarkable that the North Chinese stromatoporoids are much more diverse than their ...
... geographical change is also confirmed by the Late Ordovician Beiguoshan stromatoporoid assemblage. Such a pattern in stromatoporoids is possibly due to the different locations and times of origination of each stromatoporoid group. The earliest known labechiid stromatoporoids were reported from the Early Ordovician of South China (Jeon et al., 2019;Q. J. Li et al., 2017), but initial diversification occurred in North China in the late Middle Ordovician. Those Middle Ordovician labechiids from North China subsequently spread to other peri-Gondwanan terranes, resulting in a relatively low level of labechiid endemism during the Late Ordovician (Nestor & Webby, 2013;Stock et al., 2015). This dispersal patte ...
... The Late Ordovician stromatoporoids of North China are represented by some non-labechiid stromatoporoid groups, including clathrodictyids and stromatoporellids that appeared later (ca. 20 million years after the earliest record of labechiid stromatoporoids, Cystostroma; see Q. J. Li et al., 2017). Amongst those laminar-form stromatoporoids, i.e. clathrodictyids and stromatoporellids, only two clathrodictyid species (Clathrodictyon sp. ...
... The dominant cement composition is different from that of micrite-rich Lower Ordovician reefs, and the cement-rich reefs are indicative of global cooling and the glaciation event in the Late Ordovician (e.g., Lee and Lee, 2021;Li et al., 2023). However, ecosystem engineering, specifically the novel niches and biological interaction provided by early established skeletal-dominated reefs, may also have promoted the diversification of skeletal-reef builders (Adachi et al., 2011a;Li et al., 2016;Kröger et al., 2017). ...
... Apart from the environmental changes, a large increase in cyanobacterial calcification (Liu et al., 2020) and the rise of metazoan reef builders have been widely reported in the Ordovician (Kröger et al., 2017;Lee and Riding, 2018). Ecosystem engineering by those rising framework builders (Kröger et al., 2017;Li et al., 2017) would have played an important role in the formation of primary constructional vugs (Murray, 1960). The increased interframework pockets may be filled with sediment or later with cement. ...
... Together with pre-vious findings (Li et al., 2015), sponge-calcimicrobe-cement reefs on the Zhe-Gan platform are constructed by triple hybrid combinations of different carbonate-precipitating mechanisms, including biocontrolled (skeletal), bioinduced (mainly microbial), and abiotic carbonates (Riding and Virgone, 2020). Although the relative contributions of inorganic and organic carbonates are profoundly different, our Late Ordovician triple hybrid carbonates share a few features with their counterparts of late Palaeozoic reefs (Wood et al., 1996): The biotic association of sphinctozoan-calcimicrobes and rich synsedimentary cementations might probably indicates global cooling and glaciation events (Lee and Lee, 2021) as well as the increased ecosystem engineering of reefbuilders (Wright and Cherns, 2015;Li et al., 2017). These results suggest changes in early diagenesis and carbonate precipitation on the seafloor (Wright and Cherns, 2016), which could have played a crucial role in the consolidation of Late Ordovician reefs during the icehouse conditions before the Hirnantian glacial maximum (Vandenbroucke et al., 2010). ...
With a significant rise of metazoan reef-builders in shallow marine environments, the Late Ordovician saw a substantial increase in precipitation of crystalline crusts in reef systems, which may reflect a dramatic environmental change in the ocean. Isopachous crystalline crusts occur abundantly in boundstone of Katian sphinctozoan-bearing microbial reefs that of the Zhe-Gan Platform, South China. These deposits are characterized by in-place carbonate textures that may be interpreted as associations of abiotic, bioinduced and biocontrolled fabrics, considered here to be hybrid carbonates. Little previous work has been done to investigate how these hybrid carbon-ates are related to the transition in biotic composition during the Ordovician. These fabrics occur in well-preserved sphinctozoan-bearing reefs in the lower member of the Sanqushan Formation at the Jitoushan Section in Yushan County, Jiangxi Province. The meter-scale reefs are dominated by Corymbospongia, Amsassia and microbial components, and are cemented by fibrous sparry crusts. The intertwined Corymbospongia and Amsassia build the primary skeletal framework. Microbes and subsequent cementation enhance the stability of the framework by covering most of the macrofossil skeletons. The studied skeletal-microbial-cement reefs share a lot similarities with contemporaneous reefs of North China and Central Nepal, but differ from Early Ordovician reefs, which lack sparry crusts. Our results provide evidences for extensive synsedimentary cementation on the seafloor, which could have played a crucial role in the consolidation of Late Ordovician reefs during the icehouse conditions before the Hirnantian glacial maximum.
... STROMATOPOROIDS with skeletal structures consisting of cyst plates, denticles and pillars are traditionally classified within the Order Labechiida, and have the longest stratigraphical range amongst Palaeozoic stromatoporoid groups , Li et al. 2017, Jeon et al. 2019, Kershaw & Sendino 2020. The labechiid group has been regarded as the earliest stromatoporoid-type sponge based on its occurrence in the Floian Stage of the Lower Ordovician in South China (Li et al. 2017, Jeon et al. 2019. ...
... STROMATOPOROIDS with skeletal structures consisting of cyst plates, denticles and pillars are traditionally classified within the Order Labechiida, and have the longest stratigraphical range amongst Palaeozoic stromatoporoid groups , Li et al. 2017, Jeon et al. 2019, Kershaw & Sendino 2020. The labechiid group has been regarded as the earliest stromatoporoid-type sponge based on its occurrence in the Floian Stage of the Lower Ordovician in South China (Li et al. 2017, Jeon et al. 2019. This group diversified substantially in the late Darriwilian of the Middle Ordovician, reaching a total of 12 genera that contributed to the Great Ordovician Biodiversification Event , 2015a, 2015b, Stearn 2015. ...
... These stromatoporoids are all labechiidtype structures characterized by cyst plates with or without denticles and pillars (Webby 2015b). The earliest known stromatoporoid, Cystostroma primordia Jeon et al., 2019, occurs in the Lower Ordovician (lower Floian) strata of South China, and possesses the simplest skeletal structures amongst labechiid stromatoporoids (Li et al. 2017, Jeon et al. 2019). There are no reports of Ordovician stromatoporoids in the Dapingian to the middle Darriwilian interval, and an $18 million-year (Ma) gap seems to separate the early Floian records of stromatoporoids from the late Darriwilian ones on a global scale. ...
The Darriwilian stromatoporoid assemblage of North China represents a critical stage in early stromatoporoid development across the peri-Gondwanan regions, reaching the highest generic diversity than elsewhere as part of the Great Ordovician Biodiversification Event. We describe four labechiid stromatoporoid species, Labechia variabilis Yabe & Sugiyama, 1930, Labechiella mingshankouensis
(Ozaki, 1938), Thamnobeatricea suxianensis (Dong, 1982), and Sinodictyon columnare Yabe & Sugiyama, 1930 from the Middle Ordovician Machiakou Formation in the Xiaonanhai section near Anyang City in northern Henan Province of China. The conodont Plectodina onychodonta occurs in association with these labechiids, implying a middle to late Darriwilian age. This assemblage extends the stratigraphic range of early stromatoporoids in North China to as early as the middle Darriwilian, thus slightly older than the previously known late Darriwilian earliest records.
Mutual encrustations of Labechia variabilis and Labechiella mingshankouensis in the present assemblage constructed a microbioherm, together with unidentified tubular organisms, Ortonella-like calcimicrobes, a tubeworm-like organism, and borings. The assemblages of the Darriwilian labechiid stromatoporoids vary from place to place palaeogeographically in the shallow epicontinental marine environment of North China, i.e., in a patchy manner. The significant diversification of Darriwilian labechiid stromatoporoids in North China was likely promoted by a combination of
environmental and ecological factors, which include: (1) the occurrence of a very shallow epicontinental marine environment, (2) a possible increase in the carbonate saturation state in seawater, (3) favourable seawater temperature for stromatoporoids during the Middle Ordovician and (4) the scarcity of bryozoans and corals as ecological competitors in North China.
... In Paleozoic, there were many other organisms besides hyolithids that served as the hosts for epibionts, including moulting animals (like trilobites and mollusc Wiwaxia) (Brandt, 1996;Key et al., 2010;Topper et al., 2014), sessile animals (like brachiopodas, corals, echinoderms and sponges) (Carrera, 2000;Baumiller and Gahn, 2002;Głuchowski, 2005;Sutton et al., 2005;Zhang et al., 2010Zhang et al., , 2020Mistiaen et al., 2012;Li et al., 2017;Chen et al., 2022;Zatoń et al., 2022) and many other organisms (Cong et al., 2017;Zicha et al., 2020;Li et al., 2020;Wang et al., 2021). Even so, hyolithids were one of the most preferred hosts from the Cambrian to the Devonian in terms of the number and species of epibionts (Marek and Galle, 1976;Galle and Parsley, 2005;Sun et al., 2016Sun et al., , 2021Zicha et al., 2020), and based on this, we believe the following factors contribute to making hyolithids the preferred hosts for other epibionts: 1) The slow mobile hyolithids were unable to escape when the epibionts attached on them; 2) Hyolithids provided a fixed habitat for epibionts to inhabit for long, differing from the moulting hosts whose frequent moulting would obviously had a negative impact on the epibionts (Topper et al., 2014); 3) The mobile hyolithids could keep the epibionts in an optimum position for food, oxygen, and waste removal by facing into the current (Galle and Parsley, 2005;Horný, 2006), while the sessile animals offered less food for epibionts and likely led to burial; 4) the ornaments (growth lines and rugae) on hyolithid conchs may be helpful for attachment, fixation and stability for epibionts. ...
Reliable fossil record regarding symbiotic relationship is a key to understanding the ecology and environment in Cambrian. Herein, we exhibited an exceptionally well preserved hyolithid with intact stalked eocrinoids attaching on the operculum and conch venter, indicating the symbiotic relationship between hyolithids and eocrinoids. By that we carried out a reconstruction for their epibiotic relationship on the Cambrian sea floor. The ontogenetic features of two eocrinoids on the operculum indicated they were immature (in the early stage of ontogeny). And considering the correlation between the the orientation of eocrinoids and the mud flow in event-induced rapid burial, we proposed the orientation of eocrinoids implies the direction of mud flow. Additionally, based on specimens with signs of transport, we found that taphonomical factors besides ecological factors could also lead eocrinoids to preferentially attach on the dorsal or dorso-lateral surface of the conch, which was usually interpreted as a mutualism in previous reports though.
... The Ordovician is a crucial period for the early evolution of stromatoporoids, manifested by their first-known appearance (Li et al., 2017;Jeon et al., 2019) and early diversification . Stromatoporoids achieved one of their highestdiversity and widest-circumequatorial distributions throughout the late Middle to Late Ordovician times, as many as 26 genera on a global scale (Webby, 1979c(Webby, , 1980(Webby, , 1994(Webby, , 2015aStock et al., 2015). ...
... After the appearance of the pioneering genus Cystostroma in the early Floian in South China (Li et al., 2017;Jeon et al., 2019), 12 labechiid genera demonstrate stromatoporoid diversification in the late Darriwilian from North China, Sibumasu, Siberia, Tasmania, and Laurentia (Stock et al., 2015), in conjunction with significant global-scale development of reefbuilding organisms, including bryozoans, sponges, and corals (Carrera and Rigby, 2004;Webby, , 2015aErnst, 2018;Servais and Harper, 2018). During this period, stromatoporoid species in different terranes show a high level of endemism (Nestor and Webby, 2013;Stock et al., 2015). ...
A diverse labechiid stromatoporoid assemblage that includes 16 species in 8 genera was found in the Upper Ordovician Xiazhen Formation (mid–late Katian) at Zhuzhai, Jiangxi Province of South China. The assemblage is characterized by a combination of (1) North China provincial species succeeding from their origination in the Darriwilian, including Pseudostylodictyon poshanense Ozaki, 1938, Labechia shanhsiensis Yabe and Sugiyama, 1930, Labechia variabilis Yabe and Sugiyama, 1930, and Labechiella regularis (Yabe and Sugiyama, 1930) and (2) South China endemic species, including three new species ( Labechia zhuzhainus Jeon n. sp., Labechiella beluatus Jeon n. sp., Sinabeatricea luteolus Jeon n. gen. n. sp.), and four species in open nomenclature ( Rosenella sp., Cystostroma sp., Pseudostylodictyon sp., and Labechia sp.). The finding of Labechiella gondwanense Jeon n. sp., Stylostroma bubsense Webby, 1991, Stylostroma ugbrookense Webby, 1991, and Thamnobeatricea gouldi Webby, 1991 in the formation indicates that Tasmania was closely related to South China and had a closer paleobiogeographical relation with peri-Gondwanan terranes than with Laurentia. In addition, the occurrences of Labechia altunensis Dong and Wang, 1984 and Stylostroma species support a close biogeographic link between Tarim and South China through the Middle to Late Ordovician interval, corresponding with the results from other fossil groups such as brachiopods, conodonts and chitinozoans. The diverse labechiids from the Xiazhen Formation improve our understanding of the diversity of Ordovician stromatoporoids in peri-Gondwanan terranes and the biogeographic affinities among Australia (especially Tasmania), Tarim, and South China.
UUID: http://zoobank.org/4f46c91b-fa4c-4fe5-bea9-e409f1785677
... During the calculation of biodiversity, many problematic and phylogenetically unresolved fossil groups were wrongly categorized, especially some calcimicrobes, calcareous algae and some sponges. For example, the macrofossil calathid sponge by most authors is considered a coralline sponge which in gross morphology resembles archaeocyathids Liu et al., 1996Li et al., , 2017a. However, in biodiversity curves calathid sponges were classified as green algae together with other receptaculids (Nitecki et al., 2004). ...
... These mounds occur in tropical to subtropical carbonate platform and upper carbonate ramp depositional environments of the Paleotethyan and Iapetian realms (Appendix 6). Calathid sponges (receptaculids, Soanitidae), though phylogenetically unresolved, by most authors are considered coralline sponges which in gross morphology resemble archaeocyathids Liu et al., 1996Li et al., , 2017a. The primary mineralogy of their calcareous skeleton was aragonite (Nitecki et al., 1999;Church, 2009). ...
... Calathid-demosponge carbonate mounds contain a significant amount of non-skeletal microcrystalline Ca-carbonate (Toomey, 1970;Alberstadt et al, 1974;Church, 1974Toomey and Nitecki, 1979;Pratt and James, 1982;Johns, 1995;Kwon et al., 2003;Li et al., , 2017a for other reports see Appendix 6). A number of studies generalized the microcrystalline Ca-carbonate as lime mud or carbonate mudstone, some authors even used the term mud-mound (Krause, 2001;Pearce, 2012). ...
Bulk-rock based carbon-oxygen chemostratigraphy should be combined with a detailed understanding of depositional facies (mineralogy, porosity), its 2D-chronostratigraphic architecture, and diagenesis. The Ordovician of the western Tarim Basin recorded a peculiar litho-biostratigraphic succession. The Darriwilian Yijianfang Formation formed part of a carbonate ramp dominated by filter feeders. Toward its top, there is condensation succeeded by a multi-Myr hiatus. The hiatus correlative succession is a black-shale (Darriwilian to early Sandbian Saergan Formation) preserved in slope-to-basin settings. A marine red-bed interval (Sandbian Tumuxiuke Formation) diachronously succeeded toward a more basin-wide record. Finally, the late Sandbian to Katian Lianglitag Formation re-established a shallow-water carbonate factory (ramp-to-platform), but this time being highly productive and hosting a diversifying assemblage of benthic primary producers. By exploring diagenesis associated with a first component-specific data-set of δ13C - δ18O values and by integrating and filtering respective literature bulk-rock data, a synoptic chemo-chronostratigraphic sequence is presented. It displays segments lasting from tens of Myrs to several 100 kyrs. There is a long-term trend of increasing δ13C values culminating in the early Katian at 3.2 ‰ followed by a steady decrease. This tipping point is associated with a precursory baseline shift that interferes with the short-term Guttenberg carbon-isotope excursion presumably associated with a positive shift of δ18O values. The baseline shift was driven by regional effects of photosynthesis and a boosting (dasycladacean-related) production of aragonite along the Sandbian-Katian boundary interval. There is a medium-term (Darriwilian to earliest Sandbian) negative δ13C excursion coinciding with both a basal positive δ18O excursion and the demise of the Darriwilian carbonate ramp (Suecicus-Event, new term). It might represent the effects of volcanism/SO2-outgassing during the switch from a passive to an active continental arc. This event masquerades the elsewhere recorded middle Darriwilian carbon-isotope excursion. Caution is needed to consider the Tarim realm for global Ordovician chemostratigraphy.
... Stromatoporoids with clathrodictyid skeletal structure are the second-emerged group of stromatoporoids after labechiids, and first appeared during Late Ordovician (Webby, 2015a;, ca. 20 million years after the earliest record of stromatoporoids (labechiids) in the Floian stage of Early Ordovician (Li et al., 2017;Jeon et al., 2019). The clathrodictyid group is characterized by distinctive skeletal elements of planar, crumpled and wrinkled laminae, which are remarkably different from those of labechiids (Nestor, 2015). ...
This study reports four species belonging to two genera of clathrodictyid stromatoporoids from the Sanqushan Formation (Katian, Upper Ordovician), Changshan County, Zhejiang Province, East China: Clathrodictyon cf. mammillatum (Schmidt), C. plicatum Webby and Banks, C. cf. microundulatum Nestor and Labyrinthodictyon cascum (Webby and Morris). Of these, three species (C. cf. mammillatum, C. plicatum and L. cascum) show a wide range of skeletal variation, representing the variation from the normal skeletal phase to abnormal Ecclimadictyon-like crumpled skeletal phase, especially in the axial part of the columnar growth forms. It is uncertain whether the skeletal variation in early clathrodictyids was the result of environmental stress (such as abrupt sediment influxes that often cause growth interruptions of the stromatoporoids) or genetic inheritance. These characteristics suggest that this group had greater environmental tolerance than the earlier labechiids and increased ecological complexity during Late Ordovician by providing a stable, solid and elevated substrate for the other benthic organisms. This may imply that the labechiids and clathrodictyids are quite different groups, probably independently derived during the early evolution of stromatoporoids.
... In Paleozoic, there were many other organisms besides hyolithids that served as the hosts for epibionts, including moulting animals (like trilobites and mollusc Wiwaxia) (Brandt, 1996;Key et al., 2010;Topper et al., 2014), sessile animals (like brachiopodas, corals, echinoderms and sponges) (Carrera, 2000;Baumiller and Gahn, 2002;Głuchowski, 2005;Sutton et al., 2005;Zhang et al., 2010Zhang et al., , 2020Mistiaen et al., 2012;Li et al., 2017;Chen et al., 2022;Zatoń et al., 2022) and many other organisms (Cong et al., 2017;Zicha et al., 2020;Li et al., 2020;Wang et al., 2021). Even so, hyolithids were one of the most preferred hosts from the Cambrian to the Devonian in terms of the number and species of epibionts (Marek and Galle, 1976;Galle and Parsley, 2005;Sun et al., 2016Sun et al., , 2021Zicha et al., 2020), and based on this, we believe the following factors contribute to making hyolithids the preferred hosts for other epibionts: 1) The slow mobile hyolithids were unable to escape when the epibionts attached on them; 2) Hyolithids provided a fixed habitat for epibionts to inhabit for long, differing from the moulting hosts whose frequent moulting would obviously had a negative impact on the epibionts (Topper et al., 2014); 3) The mobile hyolithids could keep the epibionts in an optimum position for food, oxygen, and waste removal by facing into the current (Galle and Parsley, 2005;Horný, 2006), while the sessile animals offered less food for epibionts and likely led to burial; 4) the ornaments (growth lines and rugae) on hyolithid conchs may be helpful for attachment, fixation and stability for epibionts. ...
