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Sponge Spicules from the Lower Cambrian in the Yanjiahe Formation, South China: the Earliest Biomineralizing Sponge Record

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Sponge Spicules from the Lower Cambrian in the Yanjiahe Formation, South China: the Earliest Biomineralizing Sponge Record PALAEO-07880; No of Pages 9 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx–xxx Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Sponge spicules from the lower Cambrian in the Yanjiahe Formation, South China: The earliest biomineralizing sponge record Shan Chang a,b,QinglaiFenga,b,⁎, Sébastien Clausen c, Lei Zhang b a State Key Laboratory of Geological Processes and Mineral Resources, Faculty of Earth Sciences, China University of Geosciences, Wuhan 430074, China b School of Earth Science, China University of Geoscience, Wuhan 430074, China c Univ. Lille, CNRS, UMR 8198 - Evo-Eco-Paleo, F-59000 Lille, France article info abstract Article history: Sponges are the earliest metazoans, and according to molecular-clocks, have a deep history back into the Received 17 February 2016 Cryogenian. However, the record of their biomineralized elements below the Ediacaran-Cambrian Boundary Received in revised form 10 June 2016 has been controversial. Here we present the first and earliest indisputable record of hexactinellid spicules in Accepted 21 June 2016 the lowest Cambrian, Terreneuvian, below the small shelly fossil Anabarites trisulcatus–Protohertzina anabarica Available online xxxx assemblage zone (Zone I). Spicules recovered from cherts of the lowest Yanjiahe Formation in the Yichang, Hubei province, South China, include monaxons, diaxons, triaxons, and that are all siliceous in composition. Keywords: fi Terreneuvian This earliest record of silica biomineralization by lter feeding metazoan suggests sponges have contributed to Hexactinellid spicules the cycling of silica in the oceanic system as early as the beginning of the Phanerozoic. These results, along Diversification with a review of previously reported sponge occurrences, suggest a two-step evolution on sponges of the early Biomineralization Cambrian. Evolution © 2016 Published by Elsevier B.V. 1. Introduction a matter of current controversy (Borchiellini et al., 2001; Medina et al., 2001; Muscente et al., 2015). During the last decades, increasing attention has been paid on the The ongoing work for a high-resolution chronostratigraphic and cor- origin and early biomineralization of sponges. The phylum Porifera relative scheme of the Cambrian System (e.g., Clausen et al., 2015 and (sponges) is widely acknowledged to be the most ancient and basal the references therein), is of great significance to decipher the exact se- group of multicellular animals, or Metazoa (Halanych, 2004; Taylor quence of appearance of fossils and the detailed bio-radiation process et al., 2007; Srivastava et al., 2010). Recent molecular clocks indicate a during the Neoproterozoic–Cambrian interval. It is undeniable that deep Precambrian origin of the Metazoa, with a divergence of sponge sponges are one of the main sessile suspension feeders and were quite classes at about 750 million years ago (Sperling et al., 2007, 2010; common and diversified in the Cambrian metazoan community from Erwin et al., 2011). Moreover, sponges are also one of the earliest ani- the Stage 3 upward, as argued by both complete exceptionally pre- mals to secrete a three dimensional, mineral skeletal-framework, al- served specimens and abundant isolated spicules. Numerous diversified though the timing of the earliest biomineralized sponges still remains and well-preserved sponges occur in the exceptional faunas from Cam- controversial (Muscente et al., 2015). brian Series 2 of the Yangtze Platform, China, such as the Chengjiang, the Even if biomarkers, paleontological and molecular clocks all con- Niutitang, the Sancha, the Zunyi, and the Hetang faunas (e.g., Zhang and verge to a probable Cryogenian origin of sponges, their fossil record re- Pratt, 1994; Yuan et al., 2002; Xiao et al., 2005; Wu et al., 2014). Along mains controversial until the Cambrian. Consequently, it is not clearly with comparable coeval sponge faunas from South Australia and Siberia established yet if there is a bias in the fossil record of sponge biominer- (e.g., Shabanov et al., 1987; Bengtson, 1990; Rozanov and Zhuravlev, alization or if they evolved biomineralization long after their divergence 1992), they suggest that sponge groups diversified quickly during the (Muscente et al., 2015). Moreover, whether the molecular data support Cambrian Series 2. On the opposite, Terreneuvian sponge fossils are a demosponge + hexactinellid grouping to form a monophyletic silica is conspicuously rare and poorly-substantiated, and even appear to be rare to absent in the earliest Cambrian (Terreneuvian) as well as the late Neoproterozoic (Debrenne and Reitner, 2001; Xiao et al., 2005; Antcliffe et al., 2014). To date, the oldest known reliable sponge spicules ⁎ Corresponding author at: State Key Laboratory of Geological Processes and Mineral came from the Fortunian small shelly fossil Protohertzina anabarica Zone Resources, Faculty of Earth Sciences, China University of Geosciences, Wuhan 430074, China. of the Soltanieh Formation, Iran (Antcliffe et al., 2014). In South China, E-mail address: [email protected] (Q. Feng). the oldest convincing sponge spicules were reported from the lower http://dx.doi.org/10.1016/j.palaeo.2016.06.032 0031-0182/© 2016 Published by Elsevier B.V. Please cite this article as: Chang, S., et al., Sponge spicules from the lower Cambrian in the Yanjiahe Formation, South China: The earliest biomineralizing sponge record, Palaeogeogr. Palaeoclimatol. Palaeoecol. (2016), http://dx.doi.org/10.1016/j.palaeo.2016.06.032 2 S. Chang et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx–xxx Cambrian Yangjiaping Formation in Hunan (Ding and Qian, 1988)and basal dolomitic conglomerates, mostly consists of dolostone to sandy the Gezhongwu Formation in Guizhou (Yang et al., 2004). The paucity dolostone and banded black cherts. Bed 2 is dominated by siliceous of biomineralizing sponge fossils in the Precambrian and their abun- phosphatic dolostone with flat pebble conglomerates. It is followed by dance in the early Cambrian have been considered to reflect an impor- a thick succession of alternation of shale, in which siliceous- tant evolutionary process that closely linked to the major geologic, phosphatic nodules are common, and limestone (Bed 3). Bed 4 com- geomorphic and geochemical changes in the oceanic realm of the prises carbonaceous limestones. The uppermost Bed 5 is characterized Precambrian-Phanerozoic transition (Brasier, 1992; Bengtson, 1994). by cherts and siliceous phosphatic dolostone with flat pebble However, new data is required to assess this assertion and to fill the conglomerates. gap between fossil record and biomarkers or molecular data, to reveal The sponge spicules described here were obtained from the upper the early biomineralization history of sponges and its possible environ- dark cherts of Bed 1 (sample: ljc-1-2; Fig. 2). Occurrence of the acritarch mental controls. Micrhystridium regulare has been reported from the black cherts of Bed 1 In this paper we report the oldest sponge spicules from cherts of the (Ding et al., 1992). Other SSFs, including Anabarites trisulcatus, first basal Yanjiahe Formation, in Luojiacun section, Yichang, Hubei Province, occur in Bed 2 (sample ljc-2-2), suggesting that Bed 1 is earlier than South China. They occur below the small shelly fossil Anabarites SSFs Zone 1. The SSFs zone 1 has been provisionally dated from 537 to trisulcatus–Protohertzina anabarica assemblage zone (Zone I, early 532 Ma (Fortunian; Peng et al., 2012). Furthermore, a negative excur- Fortunian). The paleontological data on the Precambrian-Cambrian ap- sion of δ13Ccarb was detected from the base of the Yanjiahe Formation, pearance of sponges are summarized and discussed in the light of this which reaches its nadir (−7‰) below the SSF Zone I (Ishikawa et al., new discovery in order to understand their diversification and biomin- 2008). This excursion can be correlated with the Basal Cambrian Carbon eralization during the Cambrian explosion. Isotope Excursion (‘BACE’)(Peng et al., 2012). The Bed 1 sponge spicules therefore are confidently referred to as Fortunian in age. 2. Geological setting and stratigraphy Fossil assemblage from Bed 2 includes tubular microfossils Megathrix longus and poorly preserved acritarchs (observed in thin sec- The studied section crops out in a quarry (30°47′40.95″N, 110°54′ tion of phosphatic nodules). Radiolarians have been recovered from the 49.81″E) closed to the Luojiacun village, western of Yichang, Hubei limestone nodules of Bed 3. Megascopic algae are very abundant in Bed Province, China (Fig. 1). The Ediacaran to lower Cambrian succession 4. Bed 5 contains a new SSF assemblage, which is currently under study. is well exposed. It includes, in ascending order, the Dengying, the Yanjiahe, the Shuijingtuo, and the Shipai formations. The basal Cam- brian Yanjiahe Formation, disconformably overlies the Baimatuo Mem- 3. Material and methods ber of the Ediacaran Dengying Formation. There is also a remarkable sedimentary hiatus between the Yanjiahe and the Shuijingtuo forma- About 10 Kg of dark cherts were collected from the basal Yanjiahe tions (Fig. 2). Formation. Samples were processed following the long known method The current definition of the Precambrian–Cambrian boundary, for extracting radiolaria from cherts (Pessagno and Newport, 1972). based on the first appearance datum (FAD) of ichnospecies Trichophycus They were crushed into 1 to 2 cm pieces and then etched in 5%
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