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Abstract: South China with its rich fossil record is an important region for studying Early Cambrian metazoan diversity patterns. Compilation of a database of metazoan genera from the traditional Lower Cambrian (Cambrian Series 1–2) of South China allows a quantitative analysis of biodiversity changes. The dataset shows that about 876 genus names have been reported from the Lower Cambrian of South China. Of these only about 582 genera are now considered valid after taxonomic scrutiny. Plotting generic diversity shows a two-part, first-order trend: 1, diversity increase through the most of the Meishucunian and Qiongzhusian stages (South China usage), but punctuated by a modest decline during the upper Meishucunian; followed by 2, diversity decline in the Canglangpuan and Longwangmiaoan stages (South China usage). About 155 genera of small shelly fossils (SSFs) occur in Meishucunian rocks. The Meishucunian fauna is dominated by helcionellids, orthothecimorph hyoliths and other enigmatic fossils and is very consistent with the pretrilobitic ‘Tommotian Fauna’. Diversity increases in the Qiongzhusian stage, reaching 304 genera, the peak value known for the Cambrian. The Qiongzhusian fauna is characterized mainly by the occurrence of dominant pan-arthropods and crown-group brachiopods, representing an initial phase of the Cambrian Evolutionary Fauna. A substantial decline in generic diversity through the Canglangpuan (ca 153 genera) and Longwangmiaoan (ca 69 genera) stages is due partly to a eustatic regression, and it approximates the global pattern, reflecting an extinction event at the end of the Lower Cambrian. The observed regional biodiversity through this interval has both evolutionary and taphonomic influences: the decline in SSF diversity beginning in the upper Meishucunian was not only related to the decimation of SSFs, but also to a reduction in phosphogenesis. An apparent peak in diversity during the Qiongzhusian was somewhat exaggerated by the exceptional Chengjiang Biota. [Copyright &y& Elsevier]

Abstract: Small shelly fossils (SSFs) occur widely on the Yangtze Platform and have great potential for biostratigraphic subdivision of pre-trilobitic Early Cambrian strata. Based on the SSF record of the shallow water realm, five biozones can be recognized for the Meishucunian Stage. In ascending order the biozones are: Anabarites trisulcatus–Protohertzina anabarica Assemblage Zone; Paragloborilus subglobosus–Purella squamulosa Assemblage Zone; Watsonella crosbyi Assemblage Zone (formerly Heraultipegma yunnanensis Zone); poorly fossiliferous interzone; Sinosachites flabelliformis–Tannuolina zhangwentangi Assemblage Zone. In addition one SSF biozone is recognized for the overlying Qiongzhusian Stage: Pelagiella subangulata Taxon-range Zone. The formerly used Siphogonuchites triangularis–Paragloborilus subglobosus Zone and Heraultipegma yunnanensis Zone are discussed and redefined. Approximately 80 species of SSFs were screened for their spatial and temporal distribution on the Yangtze Platform. Variations in lithofacies and biofacies can be recognized throughout the Yangtze Platform, extending over an area of 2000×900 km. In a deeper water shelf setting the first zone is represented by the Protohertzina anabarica–Kaiyangites novilis Assemblage Zone, while younger zones are not recognized in the southern region. At the northern platform margin the Qiongzhusian is represented by the Ninella tarimensis–Cambroclavus fangxianensis Assemblage Zone and the Rhombocorniculum cancellatum Taxon-range Zone. The southeastern Shaanxi–western Hubei region followed a slightly different lithological and faunal development than the rest of Yangtze Platform, indicating a stronger similarity with parts of East Gondwana. Some taxa such as W. crosbyi, P. subangulata, R. cancellatum, Microdictyon effusum, A. trisulcatus, Protohertzina unguliformis, and P. anabarica occur nearly worldwide and support an international correlation of Early Cambrian sequences between the Yangtze Platform and smaller West Gondwanan blocks, Siberia, Newfoundland, and Australia. The six investigated zones of the Yangtze Platform comprise an interval spanning the early Nemakit–Daldynian to the late Atdabanian/early Botoman Stage on the Siberian Platform. Palaeobiogeographic analysis revealed a strong taxic similarity between the Yangtze Platform and the Tarim Platform. A smaller number of species are shared with other West Gondwanan platform fragments such as India and Iran. Palaeobiogeographic results do not support the previously reported position of the South China Block between Australia and Siberia during the Early Cambrian. [Copyright &y& Elsevier]

Abstract: Eggs, embryos, and hatchlings of olivooids are common and distinctive components of the earliest Cambrian small shelly fossil assemblage (Anabarites trisulcatus–Protohertzina anabarica Zone) at the northern and northwestern edge of the Yangtze Platform. Embryos are preserved in 3D as phosphatized permineralizations with superficial cellular structures. Nanofocus X-ray computed tomography reveals that diagenetic structures characterize the interior of most phosphatized eggs and embryos. The Olivooidae fam. nov. is herein distinguished from the possibly related hexaconulariids. Statistical size investigations and morphological studies indicate that at least three, but possibly five or more species of olivooids occur in the early Cambrian of South China. Fossilized smooth egg stages reflect a high biodiversity, encompassing 8–10 or more biological species. Eggs and embryos of Olivooides multisulcatus, Olivooides mirabilis and Quadrapyrgites quadratacris are represented by distinct and characteristic size clusters. The size ranges of embryonic parts of hatchlings roughly correspond to the size clusters of the embryos. From reconstruction of the developmental series of Olivooides multisulcatus it appears that the cleavage mode of olivooids was transitional between superficial and total cleavage. Late embryos and hatchlings of Olivooides multisulcatus and Olivooides mirabilis are characterized by pentaradial symmetry, whereas tubes of Quadrapyrgites quadratacris exhibit tetraradial symmetry. Tubular hatchlings of olivooids may represent larval or adult stages, or perhaps, nonfeeding larvae. Although the biological affinity of olivooids is weakely constrained, we hypothesize that they belong within an extinct clade of the Cycloneuralia. Their assignment to the Cnidaria is considered unlikely. [Copyright &y& Elsevier]

Abstract: The study discusses the detailed composition of Terreneuvian small shelly fossil (SSF) assemblages of eastern Yunnan Province (Meishucun, Xianfeng and Huize regions). The described sections represent shallow water deposits of the Yangtze Platform that include a number of stratigraphical hiati. The reported SSF assemblages can be assigned to the regional SSF biozones I–III (Anabarites trisulcatus–Protohertzina anabarica Assemblage Zone; Paragloborilus subglobosus–Purella squamulosa Assemblage Zone; Watsonella crosbyi Assemblage Zone). Zones II and III are slightly redefined based on new data. Two subzones of Zone II are proposed here, namely, Annelitellus yangtzensis–Obtusoconus honorabilis Subzone and Maikhanella cambriana–Oelandiella korobkovi Subzone. Claviconchella qianyii gen. et sp. nov. and Bubiites simplex gen. et sp. nov. are reported. Here we also provide a critical revision of the SSF distribution for the well-known Meishucun section. The correlation scheme for East Yunnan documents the extent and duration of stratigraphical hiati. The distribution of SSFs and the lithological characterization indicate three major events of exposure of the carbonate platform resulting in sedimentary breaks and possible karstification. This prompts new consideration of the previously described stable isotope stratigraphy of shallow platform strata. Considering the complexity of the Terreneuvian Earth history as well as the uniqueness of SSF, an integration of sedimentology, geochemistry and paleontology is needed for future SSF study. [Copyright &y& Elsevier]

The operculum of the problematic tubular fossil Cupitheca holocyclata Bengtson in Bengtson et al., (1990) is described for the first time based on collections from South Australia and North China. The phosphatized sub-circular operculum exhibits well defined cardinal processes and a narrow cardinal shield unequivocally demonstrating that Cupitheca is a hyolith, probably an orthothecid. C . holocyclata has an almost global distribution in Cambrian Stages 3–4. The apical structure of the operculum is an elevated, disc-shaped platform with a concave base and a marginal rim that could represent the scar of a specialized larval attachment structure, perhaps anchoring the larval hyolith to a sediment grain, algae or other benthic substrate. Cupitheca probably had a pelagic larval stage and settled on the seafloor by attachment of the apical disc to suitable substrates before developing a free-living benthic adult lifestyle. This contrasting mode of life compared to other hyolith genera suggests that the group had already evolved a range of distinct lifestyles in the Cambrian, providing significant clues into their ecology and distribution. [ABSTRACT FROM AUTHOR]

Abstract: Brachiopods are usually thought to contribute little to the tiering complexity from the Paleozoic to the Recent, mainly due to the fact that their recent representatives live primarily in lower tiers directly above or below the water–sediment interface. Here we present the first and oldest record of varied levels of secondary tiering in minute brachiopods attached by exceptionally preserved thread-like pedicles around the branched fronds of the algae-like Malongitubus kuangshanensis Hu, 2005. The specimens illustrated herein were recovered from the Chengjiang fauna (Series 2, Stage 3) in the Lower Cambrian Heilinpu Formation at the Kuangshan section in Malong County, Yunnan Province, southern China. The micro-morphology and oval outline of the attached brachiopods demonstrate that they can be assigned to acrotretoid brachiopods (Linguliformea, Lingulata, Acrotretoidea), described here as Kuangshanotreta malungensis gen. et sp. nov. This is the first report on the occurrence of acrotretoid brachiopods in the Lower Cambrian muddy deposits from southern China. The posterior margins of the Kuangshanotreta shells are invariably either in direct contact with, or directed towards, and then in situ attached to the algal frond of M. kuangshanensis, indicating a secondary tiering in the ecological structure of Early Cambrian brachiopods. The acrotretoid Kuangshanotreta/algae malongitubus association represents both the first and oldest evidence into the enigmatic paleoecology of the diverse acrotretoid linguliformean stock that comprises an important component of the Cambrian evolutionary fauna, and sheds light on medium-high levels of secondary tiering (+5 to +10cm) Cambrian soft substrate suspension-feeding communities. When compared to other Chengjiang brachiopods, the miniature morphology and concomitant weight reduction of the shell of K. malungensis may be a good adaption to a suspended epifaunal tiering life style. We infer that the diversification of micromorphic and miniaturized acrotretoids that occurred from the Middle Cambrian to Early Ordovician may be the result of the increased availability of habitable surfaces provided by high levels of tiering in new types of ecological spaces. [Copyright &y& Elsevier]

Helens, the curved lateral spines inserted between the conch and operculum of some hyoliths, are a unique morphological adaptation characterizing the order Hyolithida. These structures are paired, movable and had a mechanical function, probably related to orienting the hyolith conch and lifting its aperture above the sea floor. We show that helens are intimately associated with the hyolith opercula and are structurally comparable to the rod like units that constitute the clavicles, internal wall-like structures of the hyolithid operculum that probably evolved to secure the operculum from lateral displacement in the conch aperture. In some early Cambrian hyolith taxa that lack helens, such as Paramicrocornus , new clavicle rods are added in the gap separating the clavicles from the cardinal processes, the same position where helens are inserted in later hyolithids. We also show that the size of incipient helens at the earliest ontogenetic stage matches the size of the clavicles in associated opercula. We propose that helens are modified clavicle rods that were detached from the operculum and developed into lateral spines through allometric growth during early ontogeny. Further, we suggest a four-step model for the evolution of hyolithid hyoliths from orthothecid ancestors: 1, Externally fitting operculum; 2, Stabilizing, radially arranged structures on the inside of the operculum; 3, Ligula and folded operculum; 4, Detachment of clavicle rods and origin of helens. • Hyolithids are extinct benthic invertebrates with a conch and an external operculum. • Helens, lateral processes of Hyolithids, helped stabilize, re-orient and lift the conch. • Helens share a fibrous ultrastructure with the clavicles of hyolith opercula. • Helens evolved from detached clavicle rods. • Through allometric growth, clavicle rods developed into functional clavicles. [ABSTRACT FROM AUTHOR]

The lower Cambrian (Stage 3) Yu'anshan Formation in Yunnan Province, South China hosts the Chengjiang Lagerstätte , an exceptional window on the Cambrian Explosion. However, skeletal microfossils from this formation have long been neglected owing to the dominance of siliciclastics over carbonates. Here we characterize a new assemblage of small shelly fossils from the Maotianshan Shale Member of the Yu'anshan Formation. The assemblage consists of univalved mollusc, hyolith and brachiopod shells, trilobite and bradoriid carapaces, chancelloriid sclerites, sponge spicules, and echinoderm ossicles. Combining this information with taxonomic data previously tabulated for fossil assemblages from three other facies, we performed quantitative comparisons of the genus-level taxonomic diversity and modes of life and nutrition of four mutually equivalent, early Cambrian (Age 3) paleocommunities: (1) skeletal faunas of the inner shelf; (2) predominantly soft-bodied faunas of the intra-shelf basin; (3) archaeocyathan faunas of the carbonate shelf; and (4) sponge-dominated faunas of the slope to basin areas. The taxonomic and paleoecological distinctness of these four paleocommunities indicate possible low levels of competition within the communities and rapid expansion of habitat (ecospace) ranges, though with similarities in the taxonomic composition of poriferans and arthropods reflecting the rapid migration of these two groups into new habitats. The present, integrative study provides a more comprehensive portrait of metazoan diversification and paleocommunity evolution during the peak phase of the Cambrian Explosion than do investigations based on single communities. • New assemblage of skeletal fossils from the Cambrian Stage 3, South China. • The fauna is dominated by hyoliths, chancelloriids, bradoriids and molluscs. • The different paleocommunity compositions and structures are documented. • The paleocommunitites are highly facies-controlled. • The highest biodiversity of the Cambrian Explosion in South China [ABSTRACT FROM AUTHOR]