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Benthic-fixed metazoans and macroalgal holdfasts from the upper Doushantuo Formation (Ediacaran) of the Yangtze Block, South China. A, B, discoidal holdfast; C, spherical holdfast;. D, asteroidal holdfast; E, F, cone-shaped holdfast; G, I, K, Cucullus fraudulentus (Steiner, 1994); H, magnified view of G, showing the complex nonmineralized spongin fiber networks; J, magnified view of I, showing the organic wall; L, magnified view of K, showing the top pores and the side openings; M, two specimens of Cucullus; N, P, Protoconites minor (Chen et al., 1994b); O, magnified view of N, showing the transverse veins and the longitudinal ridge with tumors; Q, distribution of macroscopic algae. 

Benthic-fixed metazoans and macroalgal holdfasts from the upper Doushantuo Formation (Ediacaran) of the Yangtze Block, South China. A, B, discoidal holdfast; C, spherical holdfast;. D, asteroidal holdfast; E, F, cone-shaped holdfast; G, I, K, Cucullus fraudulentus (Steiner, 1994); H, magnified view of G, showing the complex nonmineralized spongin fiber networks; J, magnified view of I, showing the organic wall; L, magnified view of K, showing the top pores and the side openings; M, two specimens of Cucullus; N, P, Protoconites minor (Chen et al., 1994b); O, magnified view of N, showing the transverse veins and the longitudinal ridge with tumors; Q, distribution of macroscopic algae. 

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The evolution of paleoecosystems was often accompanied by the expansion of ecological niches; organismal habitats extended from the sediment surface to the water column, and then to the interior part of the sediment. A major step in ecosystem innovation was recorded in the macrobiota of the upper Doushantuo Formation during the middle-late Ediacara...

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... branching forms were Jiangk- ouphyton guizhouensis ( Wang et al., 2007a), which had main and side branchlets, and gemmiform protuberances on its topmost branchlets ( Figure 4L, M), Konglingiphy- ton erecta (Chen and Xiao, 1992), which had dichoto- mous branches and an increase in the width of the branch segments (see Chen and Xiao, 1992, pl. 1, fig. 5; Xiao et al., 2002, figs. 5.13-5.17), Longifuniculum dissolutum (Steiner et al., 1992), which possessed a twisted bundle of many dichotomously branching filaments ( Figure 4O), Miaohephyton bifurcatum Chen ( Chen and Xiao, 1991), which had Y-shaped, dichotomous branches ( Figure 4S), Sectoralga wenghuiensis ( Wang et al., 2007a) and S. ...
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... dissolutum (Steiner et al., 1992), which possessed a twisted bundle of many dichotomously branching filaments ( Figure 4O), Miaohephyton bifurcatum Chen ( Chen and Xiao, 1991), which had Y-shaped, dichotomous branches ( Figure 4S), Sectoralga wenghuiensis ( Wang et al., 2007a) and S. typica Hu ( Ding et al., 1996), which had a fan-shaped thallus that was bundled by dichotomous filaments ( Figure 4Q, R, U), Wenghuiphyton erecta ( Wang et al., 2007a), which had dichotomously branching foliation and a complex rhizoid system of the columnar and fila- mentous rhizoids ( Figure 3T), and Zhongbaodaophyton crassa (Chen et al., 1994a) and Z. robustus ( Wang et al., 2007a), which had dichotomous branching that was two branchlets in width ( Figure 4W, V). In addition, abundant macroalgal holdfasts of unknown affinity have been found, including discoidal ( Figure 5A, B), spherical (Fig- ure 5C), asteroidal ( Figure 5D), and cone-shaped hold- fasts ( Figure 5E, F), in both the Miaohe and Wenghui biota. A general feature of the middle-late Ediacaran macroalgae is a large branching or unbranching thallus with a developed holdfast. ...
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... dissolutum (Steiner et al., 1992), which possessed a twisted bundle of many dichotomously branching filaments ( Figure 4O), Miaohephyton bifurcatum Chen ( Chen and Xiao, 1991), which had Y-shaped, dichotomous branches ( Figure 4S), Sectoralga wenghuiensis ( Wang et al., 2007a) and S. typica Hu ( Ding et al., 1996), which had a fan-shaped thallus that was bundled by dichotomous filaments ( Figure 4Q, R, U), Wenghuiphyton erecta ( Wang et al., 2007a), which had dichotomously branching foliation and a complex rhizoid system of the columnar and fila- mentous rhizoids ( Figure 3T), and Zhongbaodaophyton crassa (Chen et al., 1994a) and Z. robustus ( Wang et al., 2007a), which had dichotomous branching that was two branchlets in width ( Figure 4W, V). In addition, abundant macroalgal holdfasts of unknown affinity have been found, including discoidal ( Figure 5A, B), spherical (Fig- ure 5C), asteroidal ( Figure 5D), and cone-shaped hold- fasts ( Figure 5E, F), in both the Miaohe and Wenghui biota. A general feature of the middle-late Ediacaran macroalgae is a large branching or unbranching thallus with a developed holdfast. ...
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... dissolutum (Steiner et al., 1992), which possessed a twisted bundle of many dichotomously branching filaments ( Figure 4O), Miaohephyton bifurcatum Chen ( Chen and Xiao, 1991), which had Y-shaped, dichotomous branches ( Figure 4S), Sectoralga wenghuiensis ( Wang et al., 2007a) and S. typica Hu ( Ding et al., 1996), which had a fan-shaped thallus that was bundled by dichotomous filaments ( Figure 4Q, R, U), Wenghuiphyton erecta ( Wang et al., 2007a), which had dichotomously branching foliation and a complex rhizoid system of the columnar and fila- mentous rhizoids ( Figure 3T), and Zhongbaodaophyton crassa (Chen et al., 1994a) and Z. robustus ( Wang et al., 2007a), which had dichotomous branching that was two branchlets in width ( Figure 4W, V). In addition, abundant macroalgal holdfasts of unknown affinity have been found, including discoidal ( Figure 5A, B), spherical (Fig- ure 5C), asteroidal ( Figure 5D), and cone-shaped hold- fasts ( Figure 5E, F), in both the Miaohe and Wenghui biota. A general feature of the middle-late Ediacaran macroalgae is a large branching or unbranching thallus with a developed holdfast. ...
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... dissolutum (Steiner et al., 1992), which possessed a twisted bundle of many dichotomously branching filaments ( Figure 4O), Miaohephyton bifurcatum Chen ( Chen and Xiao, 1991), which had Y-shaped, dichotomous branches ( Figure 4S), Sectoralga wenghuiensis ( Wang et al., 2007a) and S. typica Hu ( Ding et al., 1996), which had a fan-shaped thallus that was bundled by dichotomous filaments ( Figure 4Q, R, U), Wenghuiphyton erecta ( Wang et al., 2007a), which had dichotomously branching foliation and a complex rhizoid system of the columnar and fila- mentous rhizoids ( Figure 3T), and Zhongbaodaophyton crassa (Chen et al., 1994a) and Z. robustus ( Wang et al., 2007a), which had dichotomous branching that was two branchlets in width ( Figure 4W, V). In addition, abundant macroalgal holdfasts of unknown affinity have been found, including discoidal ( Figure 5A, B), spherical (Fig- ure 5C), asteroidal ( Figure 5D), and cone-shaped hold- fasts ( Figure 5E, F), in both the Miaohe and Wenghui biota. A general feature of the middle-late Ediacaran macroalgae is a large branching or unbranching thallus with a developed holdfast. ...
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... addition, numerous metazoans have been found in this ecological layer. Both elongated, sac-shaped Cucullus (Steiner, 1994) (Figure 5G-M) and Sinospongia Chen (see Chen and Xiao, 1992, pl. 6, Figs. ...
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... considered as metazo- ans (Chen et al., 1994b;Ding et al., 1996;Xiao et al., 2002;Finks and Rigby, 2004;Wang et al., 2007aWang et al., , 2011Wang and Wang, 2011). They had an organic wall (Fig- ure 5I, J), complex non-mineralized spongin fiber net- works ( Figure 5G, H), side openings that served as incurrent canals ( Figure 5G-K), and top pores that served as excurrent canals (Figure 4K, L). Wang and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. ...
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... considered as metazo- ans (Chen et al., 1994b;Ding et al., 1996;Xiao et al., 2002;Finks and Rigby, 2004;Wang et al., 2007aWang et al., , 2011Wang and Wang, 2011). They had an organic wall (Fig- ure 5I, J), complex non-mineralized spongin fiber net- works ( Figure 5G, H), side openings that served as incurrent canals ( Figure 5G-K), and top pores that served as excurrent canals (Figure 4K, L). Wang and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. ...
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... considered as metazo- ans (Chen et al., 1994b;Ding et al., 1996;Xiao et al., 2002;Finks and Rigby, 2004;Wang et al., 2007aWang et al., , 2011Wang and Wang, 2011). They had an organic wall (Fig- ure 5I, J), complex non-mineralized spongin fiber net- works ( Figure 5G, H), side openings that served as incurrent canals ( Figure 5G-K), and top pores that served as excurrent canals (Figure 4K, L). Wang and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. ...
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... had an organic wall (Fig- ure 5I, J), complex non-mineralized spongin fiber net- works ( Figure 5G, H), side openings that served as incurrent canals ( Figure 5G-K), and top pores that served as excurrent canals (Figure 4K, L). Wang and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...
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... had an organic wall (Fig- ure 5I, J), complex non-mineralized spongin fiber net- works ( Figure 5G, H), side openings that served as incurrent canals ( Figure 5G-K), and top pores that served as excurrent canals (Figure 4K, L). Wang and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...
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... and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...
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... and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...
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... and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...
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... and Wang (2011) and Wang et al. (2011) suggested that Cucullus fraudulentus ( Figure 5G-M) was a primitive member of the Demospongiae, and that the base of Cucullus nestled into the muddy substrate ( Figure 5G) while the sacciform body floated in the water. Similarly, the conical Protoconites minor (Chen et al., 1994b) (Fig- ure 5N-P), which had a thin wall (see Xiao et al., 2002, Figure 7.12, 7.13; Wang et al., 2007b, pl. 1, Figure 4), an outer sinus opening ( Figure 5N), transverse veins ( Figure 5N), and a longitudinal ridge with tumors ( Figure 5N, O), is considered as a bilateral animal, and that the sharp base was used to insert it into the sediment. ...

Citations

... 9,23,24 Among putative Ediacaran animals are a group of macrofossils with a distinctive triradial, discoidal bodyplan, known from the White Sea assemblage of Australia and Russia [25][26][27][28][29][30][31] and potentially from late Ediacaran sections of the Doushantuo Formation from Guizhou, China. 32 This group comprises the genera Tribrachidium, Anfesta, Albumares, Hallidaya, Skinnera, and Rugoconites, 25,[33][34][35][36][37] and might also include several other problematica in need of restudy (see 16 ). ...
Article
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The late Ediacaran Jiangchuan biota, from the Dengying Formation in eastern Yunnan, is well-known for its diverse macroalgal fossils, opening a window onto eukaryotic-dominated ecosystems from the late Neoproterozoic of South China. Although multiple lines of evidence suggest that metazoans had already evolved by the late Ediacaran, animal fossils have not yet been formally described from this locality. Here, we report a putative disc-shaped macrofossil from the Jiangchuan biota, Lobodiscus tribrachialis gen. et sp. nov. This specimen shows the triradial symmetry characteristic of trilobozoans, a group of Ediacaran macrofossils previously documented in Australia and Russia. Lobodiscus could record the youngest known occurrence of trilobozoans, strengthening taxonomic and ecological continuities between the Ediacaran “White Sea” and “Nama” assemblages. Our findings may expand the known paleogeographical distribution of trilobozoans and provide data for Ediacaran biostratigraphic correlations across the Yangtze block and globally, helping to track the diversification of early metazoan-grade organisms.
... It has also been proposed the denomination of vendobionts for these organisms which could constitute a separate phylum, today without descendants (Seilacher, 1989(Seilacher, , 1992Buss et Seilacher, 1994;Xiao et Laflamme, 2009;Dunn et al., 2018). In the end, the Precambrian fauna was represented as a phylogenetically diverse assemblage of organisms (Narbonne, 2005;Narbonne et al., 2014;Singer et al., 2012), totally extinct at the end of the Neoproterozoic, affected by the first mass extinction (Amthor et al., 2003;Laflamme et al., 2013;Buatois et al., 2014;Mángano and Buatois, 2014;Yue et al., 2015;Budd and Jensen, 2017;Darroch et al., 2015Darroch et al., , 2018Tarhan et al., 2018;Cribb et al., 2019;Evans et al., 2022). ...
... Transport of the fossils by gravity flows aided in their preservation (Maloney et al., 2021) but also contributed to fragmentation of the fossilized specimens, possibly masking more-complex structures and branching. These morphological advancements could indicate ecosystem-level change driven by growing competition for resources and/or space between species (Wang et al., 2015). ...
Article
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Proterozoic eukaryotic macroalgae are difficult to interpret because morphological details required for proper phylogenetic studies are rarely preserved. This is especially true of morphologically simple organisms consisting of tubes, ribbons, or spheres that are commonly found in a wide array of bacteria, plants, and even animals. Previous reports of exceptionally preserved Tonian (ca. 950−900 Ma) fossils from the Dolores Creek Formation of Northwestern Canada feature enough morphological evidence to support a green macroalgal affinity. However, the affinities of two additional forms identified on the basis of the size distribution of available specimens remain undetermined, while the presence of three unique algal forms supports other reports of increasing algal diversity in the early Neoproterozoic. Archaeochaeta guncho new genus new species is described as a green macroalga on the basis of its well-preserved morphology consisting of an unbranching, uniseriate thallus with uniform width throughout and possessing an elliptical to globose anchoring holdfast. A larger size class of ribbon-like forms is interpreted as Vendotaenia sp. A third size class is significantly smaller than Archaeochaeta n. gen. and Vendotaenia, but in the absence of clear morphological characters, it remains difficult to assign. As Archaeochaeta n. gen. and Vendotaenia represent photoautotrophic taxa, these findings support the hypothesis of increasing morphological complexity and phyletic diversification of macroalgae during the Tonian, leading to dramatic changes within benthic marine ecosystems before the evolution of animals.
... Macroscopic algae have been reported in the Paleoproterozoic (e.g., Hofmann and Chen, 1981;Han and Runnegar, 1992;Yan, 1995;Zhu and Chen, 1995;Yan and Liu, 1997;Zhu et al., 2000;Sun et al., 2004), Mesoproterozoic (e.g., Walter et al., 1976;Du et al., 1986;Kumar, 1995Kumar, , 2001Dutta et al., 2006;Sun et al., 2006;Sharma and Shukla, 2009a, b;Babu and Singh, 2011;Zhu et al., 2016), and early-middle Neoproterozoic (e.g., Hofmann and Aitken, 1979;Du, 1982;Du andTian, 1982, 1985a, b;Duan, 1982;Duan et al., 1985;Hofmann, 1985Hofmann, , 1992Walter et al., 1990;Ye et al., 2015). However, abundant and diverse macroalgae are found from the late Neoproterozoic Ediacaran strata (e.g., Gnilovskaya, 1971Gnilovskaya, , 1990Zhu and Chen, 1984;Xiao, 1991, 1992;Ding et al., 1992Ding et al., , 1996Steiner et al., 1992;Chen et al., 1994;Steiner, 1994;Yuan et al., 1995Yuan et al., , 1999Yuan et al., , 2011Yuan et al., , 2016Xiao et al., 2002Xiao et al., , 2013Grazhdankin et al., 2007;Wang et al., 2007Wang et al., , 2014Wang et al., , 2015aWang et al., , 2016aWang et al., , b, 2017Tang et al., 2008aTang et al., , 2009Singh et al., 2009;Marusin et al., 2011;Pandey and Sharma, 2017;Ye et al., 2019). Macroalgae emerged in the Paleoproterozoic and prosperously developed in the Ediacaran, which is important for understanding the biotic and environmental evolution at the dawn of animal life. ...
... Metaphytes and eukaryotic algae were considered to have tissue and/or organ differentiation, among which the macroalgal holdfast generally was used in previous publications as one of the distinguishing features of organ differentiation (e.g., Du and Tian, 1985a, b;Liu and Du, 1991;Yuan et al., 1995Yuan et al., , 2011Zhu and Chen, 1995;Ding et al., 1996;Chen et al., 2000;Xiao et al., 2002Xiao et al., , 2013Wang and Wang, 2006;Xiao, 2013;Wang et al., 2015bWang et al., , 2016aWang et al., , 2017Wang et al., , 2020a. The Precambrian macroalgal holdfast commonly consists of a rhizome that developed from the base of the thallus or stipe and a rhizoid growing on the rhizome (Wang et al., 2015a(Wang et al., , 2017(Wang et al., , 2020a. With a holdfast (a simple rhizome without a rhizoid) extended by its thallus (Walter et al., 1990;Wang et al., 2016bWang et al., , 2020a, the eukaryotic macroalga Grypanis spiralis (Walcott, 1899) Walter et al., 1976(e.g., Walter et al., 1976, 1990Runnegar, 1991;Han and Runnegar, 1992;Hofmann, 1992;Sharma and Shukla, 2009a;Wang et al., 2016aWang et al., , b, 2020a, which has been reported from the Paleoproterozoic Negaunee Iron Formation (dated to ca. 1870 Ma by Schneider et al., 2002) in the USA (Han and Runnegar, 1992), was regarded as the earliest fossil record of eukaryotic algae (Runnegar, 1991;Han and Runnegar, 1992;Wang et al., 2016bWang et al., , 2020a. ...
... 1700 Ma) in North China (see Hofmann and Chen, 1981;Yan, 1995;Zhu and Chen, 1995;Yan and Liu, 1997;Wang et al., 2020a). With a holdfast composed of a tapering or globular rhizome and filamentous or disk-like rhizoid, abundant and diverse eukaryotic macroalgae were found in the Ediacaran from Australia , India (Singh et al., 2009;Sharma et al., 2016;Pandey and Sharma, 2017), Namibia (Leonov et al., 2009), Russia (Gnilovshaya, 1971, 1990Gnilovshaya et al., 1988;Grazhdankin et al., 2007;Marusin et al., 2011), and South China Xiao, 1991, 1992;Ding et al., 1992Ding et al., , 1996Steiner et al., 1992;Chen et al., 1994Chen et al., , 2000Steiner, 1994;Yuan et al., 1995Yuan et al., , 1999Yuan et al., , 2011Yuan et al., , 2016Xiao et al., 1998, *Corresponding authors 2002Wang et al., 2007Wang et al., , 2014Wang et al., , 2015aWang et al., , 2016aWang et al., , 2017Tang et al., 2008aTang et al., , 2009Ye et al., 2019). However, the emergence of pithy macroalgae has been debated over whether a tubular or cylindrical structure within macroalgal stipes in the middlelate Ediacaran are natural biotic structures or diagenetic abiotic structures (e.g., Ding et al., 1996;Xiao et al., 2002;Wang et al., 2007Wang et al., , 2015bWang et al., , 2020aYe et al., 2019). ...
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With differentiated tissues and organs, a high-level eukaryotic macroalga Lanceaphyton xiaojiangensis n. gen. n. sp. lived on the middle–late Ediacaran (ca. 560–551 Ma) seafloor in South China. Its body had a pith (perhaps mechanical tissue) and outer tissue (perhaps epidermis and/or cortex). The lance-like macroalga consists of an unbranching thallus that grew over the sediment surface for sunlight and a holdfast grown into sediments to keep the thallus fixed on the seafloor. The pithy stipe (lower thallus) might have served to support the upper pithless thallus for photosynthesis. The holdfast is composed of a tapering pithy rhizome growing down into the sediments, with many filamentous pithless rhizoids dispersedly growing within the sediments. With the differentiated tissues and organs, especially the pith accounting for about half of the width of the rhizome and stipe, Lanceaphyton n. gen. was a high-level eukaryotic macroalga, similar to phaeophytes in morphological features, but further research is needed on its microstructural details. The pithy macroalga shows that the macroalgal pith had emerged in the Ediacaran. UUID: http://zoobank.org/bc924c5c-84e4-4170-9ca1-caee0d56c6d5 .
... Sinosabellidites and pararenicolids from the Liulaobei Formation in North China (Dong et al., 2008) also exhibits a cylindrical thallus ornamented with transverse annulations and bear a distinct holdfast structures. These morphological innovations highlight the growing competition for resources and/or space between species (Wang et al., 2015). (Jefferson and Parrish, 1989;Thorkelson, 2000;Macdonald et al., 2010;Van Acken et al., 2013;Strauss et al., 2014;Rooney et al., 2015;Baldwin et al., 2016;Milton et al., 2017;Gibson et al., 2018) this paper. ...
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Molecular phylogenetic data suggest that photosynthetic eukaryotes first evolved in freshwater environments in the early Proterozoic and diversified into marine environments by the Tonian Period, but early algal evolution is poorly reflected in the fossil record. Here, we report newly discovered, millimeter- to centimeter-scale macrofossils from outershelf marine facies of the ca. 950–900 Ma (Re-Os minimum age constraint = 898 ± 68 Ma) Dolores Creek Formation in the Wernecke Mountains, northwestern Canada. These fossils, variably preserved by iron oxides and clay minerals, represent two size classes. The larger forms feature unbranching thalli with uniform cells, differentiated cell walls, longitudinal striations, and probable holdfasts, whereas the smaller specimens display branching but no other diagnostic features. While the smaller population remains unresolved phylogenetically and may represent cyanobacteria, we interpret the larger fossils as multicellular eukaryotic macroalgae with a plausible green algal affinity based on their large size and presence of rib-like wall ornamentation. Considered as such, the latter are among the few green algae and some of the largest macroscopic eukaryotes yet recognized in the early Neoproterozoic. Together with other Tonian fossils, the Dolores Creek fossils indicate that eukaryotic algae, including green algae, colonized marine environments by the early Neoproterozoic Era.
... Abundant and diverse carbonaceous compressions (the Wenghui biota), including macroscopic macroalgae, metazoans, and ichnofossils, are found in the Ediacaran black shales of the upper Doushantuo Formation in northeast Guizhou, South China (Wang et al., , 2007(Wang et al., , 2010(Wang et al., , 2012(Wang et al., , 2015a(Wang et al., , 2016aTang et al., 2008aTang et al., , 2008bTang et al., , 2009Wang andWang, 2008, 2011;Zhu et al., 2008). A disc-like compression, with an unbranching carbonaceous stipe growing out from its center, is one of the abundant taxa in the Wenghui biota. ...
... The uppermost Member IV consists of fossiliferous black shales, underlying bedded cherts of the Liuchapo Formation (Fig. 1). Apart from the disc-like compression, abundant and diverse macroscopic fossils (i.e., the Wenghui biota) have been collected in black shales of the Member IV (e.g., Wang et al., 2005Wang et al., , 2007Wang et al., , 2009Wang et al., , 2010Wang et al., , 2014Wang et al., , 2015aWang et al., , 2016aWang and Wang, 2006Tang et al., 2008aTang et al., , 2008bTang et al., , 2009Zhu et al., 2008;Cheng et al., 2013). ...
... In the Wenghui biota, not only macroscopic fossils but also filamentous rhizoids of macroalgae are well preserved, therefore previous researchers generally considered that this biota lived in a relatively low-energy environment and was preserved in situ or nearby their growth position (Wang et al., , 2007(Wang et al., , 2015a(Wang et al., , 2016aWang andWang, 2006, 2011;Cheng et al., 2013). ...
Article
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The fixing organ of the Precambrian macroalga was briefly described by most researchers as a holdfast or rhizoid, suggesting a fixation structure and/or tissue differentiation. An Ediacaran macroscopic alga, Discusphyton whenghuiensis n. gen. n. sp., with a complex disc-like holdfast and an unbranching thallus, has been collected, together with abundant and diverse macrofossils (i.e., the Wenghui biota) in black shales of the upper Doushantuo Formation (~560–551 Ma) in northeastern Guizhou, South China. The Wenghui biota lived in a relatively low-energy marine environment and was preserved in situ or nearby their growth position. Morphologically, the macroalgal thallus, including the compressed lamina and cylindrical stipe, might have been suspended in the water column for photosynthesis. Its holdfast, a rare fixing form, is complex in structure and construction, consisting of a globular rhizome and a discoidal rhizoid. The large-sized discoidal rhizoid is regarded as a flat-bottomed and dome-shaped organ to attach the macroalga on the water-rich muddy seafloor. The globular rhizome, expanded by a thallus on the substrate, was originally harder and spherical nature within the dome-shaped rhizoid. It may have been an important organ as a steering knuckle to connect between the stipe and the rhizoid. The macroscopic metaphyte D . whenghuiensis n. gen. n. sp. shows the appearance of complex holdfast in morphology and bio-functions. However, not enough is known, in the absence of more information, to decipher the phylogenetic affinity of D . whenghuiensis n. gen. n. sp. and the origin of a discoidal rhizoid.
... Grypania differs from Vendotaenia Gnilovskaya, 1971 in that the ribbon-like body of the latter has longitudinal veins and is irregularly folded or curved, whereas Grypania is usually coiled, curled, or regularly curved. Grypania is also distinct from Linbotulichnus Li & Ding in Ding et al., 1996, a taenioid compression meandering on bedding planes hosting the Miaohe biota (Ding et al. 1996 and the Wenghui biota , 2009, 2015b, Tang et al. 2008, because the latter consists of bead-shaped segments with crescentic spaces (Ding et al. 1996, 2015b. Grypania spiralis is similar to Jiuqunaoells simplicis Chen in Chen & Xiao, 1991, a carbonaceous compression from the Miaohe biota, in representing a curved ribbon-like compression, but the latter is irregularly curved and bears discontinuous and close transverse wrinkles on a ribbon of variable width (Chen & Xiao 1991. ...
... Grypania differs from Vendotaenia Gnilovskaya, 1971 in that the ribbon-like body of the latter has longitudinal veins and is irregularly folded or curved, whereas Grypania is usually coiled, curled, or regularly curved. Grypania is also distinct from Linbotulichnus Li & Ding in Ding et al., 1996, a taenioid compression meandering on bedding planes hosting the Miaohe biota (Ding et al. 1996 and the Wenghui biota , 2009, 2015b, Tang et al. 2008, because the latter consists of bead-shaped segments with crescentic spaces (Ding et al. 1996, 2015b. Grypania spiralis is similar to Jiuqunaoells simplicis Chen in Chen & Xiao, 1991, a carbonaceous compression from the Miaohe biota, in representing a curved ribbon-like compression, but the latter is irregularly curved and bears discontinuous and close transverse wrinkles on a ribbon of variable width (Chen & Xiao 1991. ...
... Advancements in analytical facilities and their access have helped fathom the hitherto unknown world of the biosphere (Schopf et al., 2016; Porter et al., 2016; Cole et al., 2016). These studies have shown newer ecological niches adopted by the biotic communities with sudden or gradual changes in the hydrosphere/ atmosphere or lithosphere (Ye et al., 2015; Wang et al., 2015b; Yuan et al., 2011 Yuan et al., , 2013 Zhu et al., 2016). ...
... Wang et al. (2015a) described a body fossil of Ediacara carbonaceous compression fossil Zhongaodaophyton Chen et al. from the black shale of the Upper Doushantuo Formation, south China as a macroalgal form comparable to the members of phaeophyta. On the basis of Upper Doushantuo biotic assemblage, Wang et al. (2015b) suggested the existence of three tiered palaeoecosystem of macroalgal forms and established a multilayered ecological pyramid in the biosphere with increasing demand of oxygen which pinnacled at Pc-C boundary. Ye et al. (2015) reported carbonaceous compression fossils (benthic macroalgae ) from the Marinoan-age Nantuo Formation in South China. ...
... Advancements in analytical facilities and their access have helped fathom the hitherto unknown world of the biosphere (Schopf et al., 2016; Porter et al., 2016; Cole et al., 2016). These studies have shown newer ecological niches adopted by the biotic communities with sudden or gradual changes in the hydrosphere/ atmosphere or lithosphere (Ye et al., 2015; Wang et al., 2015b; Yuan et al., 2011 Yuan et al., , 2013 Zhu et al., 2016). ...
... Wang et al. (2015a) described a body fossil of Ediacara carbonaceous compression fossil Zhongaodaophyton Chen et al. from the black shale of the Upper Doushantuo Formation, south China as a macroalgal form comparable to the members of phaeophyta. On the basis of Upper Doushantuo biotic assemblage, Wang et al. (2015b) suggested the existence of three tiered palaeoecosystem of macroalgal forms and established a multilayered ecological pyramid in the biosphere with increasing demand of oxygen which pinnacled at Pc-C boundary. Ye et al. (2015) reported carbonaceous compression fossils (benthic macroalgae ) from the Marinoan-age Nantuo Formation in South China. ...
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Many facets of the early biosphere preserved in the Proterozoic and Early Cambrian successions, recorded during 2011-2015, are reviewed. Recent advancements made in the Precambrian palaeobiology and newer steps recognized in theorganismal evolution in the global perspective vis-a-vissignificance of Indian records are discussed. The palaeobiologicalevidence reported from India are chronicled and grouped under seven categories: MISS (Microbially Induced SedimentaryStructure) & stromatolites, acritarchs, OWM (OrganicWalled Microfossils), carbonaceous remains, trace-fossils & Ediacaranfossils, stable isotope studies and organic geochemistry. Present article is a continuum of Sharma et al. (2012), provides thestatus of Precambrian palaeobiological studies in the country, enumerates the importance of Indian records, and enlistunsolved problems and future research directions.
... Grypania differs from Vendotaenia Gnilovskaya, 1971 in that the ribbon-like body of the latter has longitudinal veins and is irregularly folded or curved, whereas Grypania is usually coiled, curled, or regularly curved. Grypania is also distinct from Linbotulichnus Li & Ding in Ding et al., 1996, a taenioid compression meandering on bedding planes hosting the Miaohe biota (Ding et al. 1996 and the Wenghui biota , 2009, 2015b, Tang et al. 2008, because the latter consists of bead-shaped segments with crescentic spaces (Ding et al. 1996, 2015b. Grypania spiralis is similar to Jiuqunaoells simplicis Chen in Chen & Xiao, 1991, a carbonaceous compression from the Miaohe biota, in representing a curved ribbon-like compression, but the latter is irregularly curved and bears discontinuous and close transverse wrinkles on a ribbon of variable width (Chen & Xiao 1991. ...
... Grypania differs from Vendotaenia Gnilovskaya, 1971 in that the ribbon-like body of the latter has longitudinal veins and is irregularly folded or curved, whereas Grypania is usually coiled, curled, or regularly curved. Grypania is also distinct from Linbotulichnus Li & Ding in Ding et al., 1996, a taenioid compression meandering on bedding planes hosting the Miaohe biota (Ding et al. 1996 and the Wenghui biota , 2009, 2015b, Tang et al. 2008, because the latter consists of bead-shaped segments with crescentic spaces (Ding et al. 1996, 2015b. Grypania spiralis is similar to Jiuqunaoells simplicis Chen in Chen & Xiao, 1991, a carbonaceous compression from the Miaohe biota, in representing a curved ribbon-like compression, but the latter is irregularly curved and bears discontinuous and close transverse wrinkles on a ribbon of variable width (Chen & Xiao 1991. ...
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Full-text available
Wang, Y., Wang, Y. & Du, W., February 2016. The long-ranging macroalga Grypania spiralis from the Ediacaran Doushantuo Formation, Guizhou, South China. Alcheringa 40, xxx–xxx. ISSN 0311-5518 Grypania spiralis (Walcott) Walter et al., a macroalga previously reported in pre-Ediacaran successions, has been collected, together with abundant macrofossils (i.e., the Wenghui biota), from black shales of the upper Doushantuo Formation (ca 593 to 551 Ma) in northeastern Guizhou, South China. Morphologically, G. spiralis represents a carbonaceous ribbon with a continuum of forms from coiled to nearly straight. Its helicoid main body might have been suspended in the water column for photosynthesis with one end anchored or nestled into soft sediments. Grypania possessed morphological stability, and its habit endowed great competitiveness for sunlight. Remarkably, it did not change significantly in size or morphology over more than 1200 Myrs. Ye Wang [[email protected]], School of Earth Sciences and Resources, PR China University of Geosciences, Beijing 100083, PR China; Yue Wang [[email protected]] (corresponding author), School of Resources and Environments, Guizhou University, Guiyang 550003, PR China; Wei Du [[email protected]], Department of Earth Science and Astronomy, The University of Tokyo, Tokyo 153-8902, Japan.