Journal of Earth Science  2018, Vol. 29 Issue (4): 912-919   PDF    
New Siliceous Microfossils from the Terreneuvian Yanjiahe Formation, South China:The Possible Earliest Radiolarian Fossil Record
Shan Chang1,2, Qinglai Feng1,2, Lei Zhang1    
1. School of Earth Sciences, China University of Geosciences, Wuhan 430074, China;
2. State Key Laboratory of Geological Processes and Mineral Resources, China University of Geosciences, Wuhan 430074, China
ABSTRACT: Radiolarians form an important part of the planktonic realm in the ocean of Early Paleozoic, but their origin and evolutionary processes has long been enigmatic. The ancestral representatives of radiolarians have been considered to belong to the order Archaeospicularia, whose unquestionable fossil records were dated back to the Middle Cambrian. Here we report? Blastulospongia and unnamed spherical radiolarians in the Terreneuvian from the Yanjiahe Formation in Hubei Province, South China. Blastulospongia is an enigmatic siliceous microfossil genus, with affinities proposed amongst the radiolarian, sphinctozoan-grade sponges and uncertain protists. As for the newly discovered unnamed radiolarians, morphologically they possess latticed shell, spherical shape and are all small in size. Our discoveries support the idea that spherical radiolarians is an ancient representative, whose origin and diversification was probably much earlier than generally accepted. The hypothesis that the oldest radiolarians belong to the order Archaeospicularia needs to be re-examined.
KEY WORDS: Cambrian    radiolarian    ?Blastulospongia    silica-biomineralization    


Apart from exquisite skeletons and astonishing biodiversity, radiolarians are one of the key components in the marine system with silica-biomineralized skeletons. Their origin and early history has long been studied extensively, including molecular data and the fossil record (Decelle et al., 2012; Ishitani et al., 2011; Pouille et al., 2011; Danelian and Moreira, 2004; Nazarov, 1975, 1973). Identifying the earliest record of radiolarians is undoubtedly important in regard to the rise of siliceous biomineralization in eukaryotic lineages and its impact on the evolution of the silica cycle (Zhang et al., 2013 and references therein). Moreover, they provide information on the origin and initial establishment of "modern"-type marine ecosystems during the transitional interval between the Ediacaran and Cambrian, since they constitute the main representatives of the heterotrophic plankton (De Wever et al., 2001; Anderson, 1983). However, little is known concerning the morphology of the earliest radiolarians, since fossil records are scarce, especially in old sequences. Despite their origin can be dated back to the Neoproterozoic by molecular data (Decelle et al., 2012), the earliest convincing fossil record of radiolarians is from the Middle Cambrian (Won and Below, 1999). The ancestor of radiolarians has been considered to belong to the order Archaeospicularia, and the spicular or needle-like morphology has been postulated to represent an ancient characteristic (Maletz, 2011).

However, history of radiolarians in the Early Cambrian or the Neoproterozoic is still poorly known. So far, reports concerning Early Cambrian radiolarians are still highly incomplete, among them, few specimens have been described (e.g., Braun et al., 2007a; White, 1986), some of the specimens are misinterpreted as nonbiologic objects or skeletal plates of the lobopod animal Microdictyon (Zhang and Aldridge, 2007; Lipps, 1992; Shu and Chen, 1989; Peng, 1984), some specimens are poorly preserved (Cao et al., 2014, Fig. 4l therein; Braun et al., 2007a). Besides, ambiguous age of some sequences in the Lower Cambrian poses another challenge since there are neither proper fossil zones nor geochemical data.

Here we report new siliceous microfossils recovered from the Yanjiahe Formation in the Luojiacun Section, Zigui County, Hubei Province, South China. According to SSFs biostratigraphy, the Yanjiahe Formation belongs to Early to Middle Meishucunian in age, corresponding to the Terreneuvian internationally (Guo et al., 2014). The fossils, recovered from the limestone nodules in the middle of Yanjiahe Formation, include?Blastulospongia sp. and unnamed spherical radiolarians, they both represent the possible earliest radiolarian fossil record. These fossils are less likely to be contamination because Blastulospongia is a genus that only reported from the Cambrian; the unnamed radiolarians are very similar to those reported from the Kuanchuanpu Formation (Terreneuvian) and the Shuijingtuo Formation (Series 2) (Cao et al., 2014; Braun et al. 2007a). Our findings support the idea by Braun et al. (2007a) that spherical radiolarian is an original representative and has an older history than previous thought. Along with sponges (e.g., Chang et al., 2017; Braun et al., 2007b), it is likely that the Cambrian invasion of siliceous skeleton-bearing radiolaria in the oceanic realm altered profoundly oceanic silica cycle during the Ediacaran–Cambrian transition. Their early biomineralization and flourish might be related to the high concentration of silica in the seawater caused by the breakup of the Rodinia supercontinent during the Neoproterozoic– Cambrian transition.


The studied section crops out in a quarry (30°47'40.95"N, 110°54'49.81"E) closed to the Luojiacun Village, western Yichang, Hubei Province, China (Fig. 1). The Ediacaran to Lower Cambrian succession is well exposed. It includes, in ascending order, the Dengying, the Yanjiahe, the Shuijingtuo, and the Shipai formations. The basal Cambrian Yanjiahe Formation, disconformably overlies the Baimatuo Member of the Ediacaran Dengying Formation. There is a hiatus between the Yanjiahe and the Shuijingtuo formations (Fig. 2).

Figure 1. Geological map of Zigui County in South China and location of the studied area (schetch map of China modified after, No. GS(2016)1569).
Figure 2. Stratigraphic column of the Luojiaping Section and field photos. (1) Field photo showing the contact relationship of the Yanjiahe Fm. with the Shuijingtuo Fm.; (2) the magnified picture of the rectangle area in (1); (3) siliceous phosphatic-interclastic dolostone in Bed 5; (4) field photo showing the contact relationship of the Yanjiahe Fm. with the Dengying Fm.; (5) the weathering crust; (6) conglomerates (marked by the arrows) in the base of the Yanjiahe Fm. Fm. Formation; Mb. member.

In biostratigraphy, Chen (1984) identified two small shelly fossils (SSFs) assemblage zones within the Yanjiahe Formation: Circotheca-Anabarites-Protohertzina assemblage zone and Lophotheca-Aldanella-Maidipingoconus assemblage zone. Guo et al. (2014) revised the SSFs into three assemblage zones, namely: the Anabaritestrisulcatus-Protohertzinaanabarica assemblage zone (Zone Ⅰ), the Purellaantiqua assemblage zone (Zone Ⅱ) and the Aldanellayanjiaheensis assemblage zone (Zone Ⅲ) in ascending order (Fig. 2). Based on SSFs, the Yanjiahe Formation was deposited in the Early to Middle Meishucunian and corresponds to the Cambrian Terreneuvian internationally.

In the studied area, the Yanjiahe Formation can be subdivided into 5 parts. Bed 1, characterized by basal dolomitic conglomerates, mostly consists of dolostone to sandy dolostone and banded black cherts. Bed 2 is dominated by siliceous phosphatic dolostone with flat pebble conglomerates. It is followed by a thick succession of alternation of shale, in which carbonate and phosphatic nodules are common, and limestone (Bed 3). Bed 4 comprises carbonaceous limestones. The uppermost Bed 5 is characterized by cherts and siliceous phosphatic dolostone with flat pebble conglomerates. The radiolarian fossils were recovered from the limestone nodules in Bed 3, which belongs to the Purellaantiqua Zone (Zone Ⅱ) in the Yanjiahe Section. This bed can be further correlated with the Upper Zhongyicun Member in eastern Yunnan, the Upper Maidiping Formation in western Sichuan, the Upper Gezhongwu Member in northern Guizhou (Guo et al., 2014).


Recrystallization in cherts during diagenesis process can strongly influence the microstructures of radiolarian fossils. This study focused on limestone nodules to detach radiolarians. A large amount of limestone nodules were collected from the middle part of the Yanjiahe Formation (Bed 3) in the field work. Sample was then crushed into small pieces of about 1 cm3. The fragments were placed in plastic barrels with 10% acetic acid solution (nine parts water to one part acid) for a period of about one week, the upper acid is discarded and the sample is washed until enough residues were detached. Subsequently, the residue was sieved (diameter=0.038 and 0.083 mm) carefully, dried at room temperature, and then hand-picked under a binocular microscope. Isolated specimens were mounted on stubs with latex then examined on stereoscan electron microscope (SEM) and electronic differential system (EDS) analysis in the State Key Laboratory of Geological Process and Mineral Resources (Wuhan).

In our study, abundant microfossils have been recovered, including radiolarians, SSFs, sponge spicules, etc. Among them, six radiolarian fossils were described. According to small shelly fossil biostratigraphy, the radiolarian fossils, including?Blastulospongia sp. and unnamed spherical radiolarians, belong to Purellaantiqua assemblage zone of the Yanjiahe Formation (Guo et al., 2014), which can be correlated with the second SSF biozone, SiphogonuchitestriangularisPurellasquamulosa assemblage zone in eastern Yunnan (Steiner et al., 2007), corresponding to the Early Meishucunian Stage of Chinese usage, Upper Nemakit–Daldynian in Siberia and Terreneuvian, Fortunian Stage of international scheme (Peng et al., 2012; Khomentovsky and Karlova, 1993).


All of the specimens described here are deposited in the Geological Museum of China University of Geosciences (Wuhan), China.

Phylum uncertain

Class uncertain Family uncertain

Genus Blastulospongia Pickett and Jell, 1983

Diagnosis: Spherical to oval shell, perforated by numerous pores which are usually evenly distributed. Wall thin, interior of tests empty, the primary composition is uncertain.

?Blastulospongia sp.

Figure 3 (1–3)

Figure 3. Radiolarian fossils from the Yanjiahe Formation in the Loujiaping Section. (1, 3) Blastulospongia sp. (1) YJH03-11; (2) shows the magnified picture of the perforations; (3) YJH03-12. (4, 7) spherical radiolarian Ⅰ. (4) YJH03-21, with the pentagonal or hexagonal frames amplified in (5, 6); (7) YJH03-22. (8–11) Spherical radiolarian Ⅱ, poorly preserved. (8) YJH03-31; (9) YJH03-32; (10) YJH03-33; (11) YJH03-34. Scale bars length: 50 μm.

Material: Two specimens: YJH03-11, YJH03-12.

Occurrence: Early Cambrian–South China, Siberia; Middle Cambrian, West Utah; Late Cambrian, Queensland.

Description: Globular tests, single and small shell, about 70 μm in diameter, with rounded or angular perforations evenly distributed all over the specimen. Spineless, wall thin. Composed of microcrystalline silica. Most of the perforations are nearly circular in outline, fairly and evenly spaced, about 5–8 μm in diameter, some angular ones might partially be attributed to recrystallization.

Remarks: Blastulospongia is an enigmatic genus that only receives specimens from Cambrian strata, with siliceous wall and varied diameter. It was originally interpreted as sponge by Pickett and Jell (1983). The type species, B. monothalamos, was described to be extremely large (1–1.9 mm in diameter). Bengtson (1986) reported B. mindyallica from the Late Cambrian of Queensland and proposed that this genus would be a radiolarian, if the original composition could have been siliceous. White (1986) reported an unnamed spumellarian from the Middle Cambrian, which is recognised as Blastulospongia for its strong similarities with this genus. In South China, Conway Morris and Chen (1990) reported B. polytreca from the Shuijingtuo Formation (the Cambrian Series 2). The B. polytreca was characterized by thin wall, bearing dimples and folds of varying openness. However, possibility of post-mortem damage cannot be ruled out for these dimples and folds, since the shells of B. polytreca are rather thin and would be easily influenced by sedimentary compaction. Conway Morris and Chen (1990) further provided evidence that the Blastulospongia was unlikely to be sponge for its siliceous composition that probably being primary. Kouchinsky et al. (2017) reported specimens recovered from Cambrian Terreneuvian in Siberia, by experiments of energy-dispersive X-ray analyses and their co-occurrence with calcitic shelly fossils, the siliceous composition of Blastulospongia was further confirmed to be original, rather than reflecting selective replacement by silica during diagenesis.

New materials described here are siliceous (Fig. 4a), co-occurred with calcitic shelly fossils. However, our specimens have some differences with those mentioned above. The specimens reported from Medvezhya Formation (Terreneuvian, Stage 2) in Siberia includes 2 specimens, the smaller specimen is ca. 350 μm and the larger one ca. 800 μm in diameter (Kouchinsky et al., 2017). Our specimens are rather small in diameter, the perforations are evenly spaced, and shows no deformation in the wall. Compared to B. polytreca, the perforations of our specimens are more sparsely distributed; compared to B. mindyallica, the shell of our specimens are thicker. The diameter of specimens from the Shuijingtuo Formation reported by Conway Morris and Chen (1990) is between 350 to 370 μm; the specimen described by White (1986) from the Middle Cambrian was about 380 μm, but the size range of his materials was not provided; the diameters of specimens reported from the Late Cambrian were between 280–520 μm (Bengtson, 1986). The diameter of this genus varies from 280 to 1 900 μm, some of the specimens have rather thin wall and tend to be deformed with dimples and folds (e.g., B. polytreca and one specimen from Siberia) but others do not show deformation. Our specimens share the morphology of single chambered, asiphonate, porate, without internal skeletal structures with Blastulospongia, but is rather small in size, it's possible that these differences represent two different lineages, so they are tentatively assigned to Blastulospongia.

Figure 4. Electronic differential system (EDS) analysis for the siliceous microfossils from the Yanjiahe Foramtion. (a) EDS result of the specimen of Blastulospongiashown in Fig. 3-1; (b) EDS result of the specimen of the unnamed spherical radiolarianshown in Fig. 3-7.

As to this controversial question of whether Blastulospongia belongs to a sponge, radiolarian, or something else, observations of this study are in accordance with Conway Morris and Chen (1990) and Kouchinsky et al. (2017), which shows that Blastulospongia are less similar to sphinctozoan- grade. In morphology our specimens are most similar to that reported by White (1986) and they might belong to one species. In addition, if it is a radiolarian, to the familiar latticed architecture, radiolarians can also produce spherical tests of denser material with perforations, it is possible that radiolarian could have been diversified as a part of the 'Cambrian explosion'.

Blastulospongia is similar to a small shelly fossil, assigned as Aetholicopallaadnata (Conway Morris and Chen, 1990), in the round shape with perforations. Aetholicopallaadnata have been widely reported from the Lower Cambrian strata (Yang et al., 2014; Kouchinsky et al., 2013; Wrona, 2004; Elicki, 1998). But Aetholicopallaadnata is characteristic of double-walled shells connected by hollow pillars and preserved as phosphate (Wrona, 2004), which can be distinguished from Blastulospongia.

Suborder, family and genus unknown

Spherical radiolarian

Figure 3. 4–7

Material: Six specimens: YJH03-21, YJH03-22, YJH03-31, YJH03-32, YJH03-33, YJH03-34.

Occurrence: Early Cambrian, South China.

Description: Spherical, latticed shell, small in diameter, about 125 μm. The perforations are circular with pentagonal or hexagonal frames. Inner structure unclear. Siliceous in composition (Fig. 4b).

Remarks: These spherical radiolarians are very similar to those reported from the Kuanchuanpu Formation in Ningqiang, Shanxi Province, China (Braun et al., 2007a), they possessed latticed shell and spherical shape with small sizes, while the latter specimens possess short spines. In the Shuijingtuo Formation, Cao et al. (2014) reported similar spherical radiolarian, but its age is younger (Series 2, Stage 3) and its diameter is bigger (240 μm). The specimens described here are from the same region with those reported by Cao et al. (2014), not only validating the presence of radiolarians in the Early Cambrian sequences, but also providing opportunities to reveal characteristics of early radiolarian. The spherical morphology might represent an original characteristic of radiolarian, which was evolved much earlier than that has been generally accepted. The frame structure displayed similarities to some genera occurred in Ordovician, but due to recrystallization its inner structure was unclear. However, its systematic position and nomenclature requires additional material and more conspicuous characters showing the inner structure.


In the Yanjiahe Formation, all these siliceous fossils yielded are small in size, with the average diameter being approximately 100 μm. The specimens reported from the Kuanchuanpu Formation are also relatively small, about 160 μm in diameter (Braun et al., 2007a). The specimens reported by Obut and Iwata (2000) from the Botomian (Lower Cambrian) in the Altai Mountains (Siberia) are even smaller. Indeed, the shell diameter of species Archaeocenosphaeramuricata Obut and Iwata ranges between 80 and 100 μm (Obut and Iwata, 2000). The specimens of Spongomassanannosphaera Won, described from the Middle Cambrian of Australia, varies between 93 and 130 μm in diameter (Obut and Iwata, 2000). The relationship of body size with oxygen availability has been discussed widely, with lots of studies showing that the increase of oxygen level is a critical trigger for evolution of life (Li et al., 2017; Zhang and Cui, 2016; Zhang et al., 2014; Payne et al., 2009; Canfield et al., 2007; Cloud, 1968). Specifically increased oxygen availability leads to increased sizes, as demonstrated by fossil records of protists (primarily in aerobic heterotrophs) and animals (Payne et al., 2009) and experiments in laboratory settings (Klok et al., 2009). Studies from geochemistry show that oxygen availability was still very low in South China (e.g., Liu et al., 2017; Jin et al., 2016), lithologically the wide spread black shall and cherts composition also indicate a relatively poor redox condition in the lower sequences of South China (e.g., Zhou and Jiang, 2009; Li et al., 1999). The small size of the?Blastulospongia and the spherical radiolarian fossils in the Yanjiahe Formation might be an original character and the low oxygen availability might be a factor that limited their size, but considerable complexity remains.

The ancestral representatives of radiolarians have been considered to belong to the order Archaeospicularia, and the simple spicular or needle-like morphology has been postulated to represent an ancient characteristic (Maletz, 2011). However, no elements of Archaeospicularia has been found in the Lower Cambrian till now, moreover, the simple, 'original' morphology described by Maletz (2011) could be easily confusing with Hexactinellida sponge spicules or some abiogenetic minerals for their similar outline, size and siliceous composition.

The record of radiolarians from the Lower Cambrian or even the Late Neoproterozoic would certainly provide a great breakthrough as regards to the radiolarian research as well as the Cambrian research. Recently there have been multi-disciplinary works on the Edicaran–Cambrian transition from South China (e.g., Ding et al., 2017; Mason et al., 2017; Hu et al., 2016; Yin et al., 2016; Zhang et al., 2016), as the Proterozoic to Early Cambrian strata are well exposed there, many publications provided evidence of radiolarian fossils shown in thin sections (He et al., 2013; Zheng et al., 2012; Zhao, 1999; Yin et al., 1994), and all of them possess spherical shell. But due to recrystallization the inner structures are difficult to observe. Reports regarding old radiolarian fossils from South China have a potential for our knowledge on the early history of radiolarians. Morphologically they possess latticed shell and spherical shape that are quite different from the postulated ancestral radiolarians (Archaeospicularia). The original character of radiolarian is still open to question, but evidence from both thin sections and acid etching analysis shows that the morphology of radiolarian in South China is different from that of Australia specimens during Cambrian. The hypothesis that oldest radiolarians belong to the order Archaeospicularia needs to be re-examined, and a phylogenetic analysis concerning old radiolarians must await more investigations.


This research was supported by the NSFC (Nos. 41430101, 41502014). We express our sincere thanks to Yan Zhang, Fenggang Lan, Wei Guo and Ke Zhang for help in the fieldwork. Professor Jonathon Aichison and anonymous reviewers are greatly appreciated for improving the manuscript. The final publication is available at Springer via

Anderson, O. R., 1983. Radiolaria. Springer, New York. 271
Bengtson, S., 1986. Siliceous Microfossils from the Upper Cambrian of Queensland. Alcheringa:An Australasian Journal of Palaeontology, 10(3): 195-216. DOI:10.1080/03115518608619155
Braun, A., Chen, J. Y., Waloszek, D., et al., 2007a. First Early Cambrian Radiolaria. In: Vickers-Rich, P., Komarower, P., eds., The Rise and Fall of the Ediacaran Biota. Geological Society, London, Special Pub-lications, 286(1): 143-149.
Braun, A., Chen, J. Y., Waloszek, D., et al., 2007b. Siliceous Microfossils and Biosiliceous Sedimentation in the Lowermost Cambrian of China. In: Vickers-Rich, P., Komarower, P., eds., The Rise and Fall of the Ediacaran Biota. Geological Society, London, Special Publications, 286(1): 423-424.
Canfield, D. E., Poulton, S. W., Narbonne, G. M., 2007. Late-Neoproterozoic Deep-Ocean Oxygenation and the Rise of Animal Life. Science, 315(5808): 92-95. DOI:10.1126/science.1135013
Cao, W. C., Feng, Q. L., Feng, F. B., et al., 2014. Radiolarian Kalimnas-phaera from the Cambrian Shuijingtuo Formation in South China. Marine Micropaleontology, 110(2): 3-7. DOI:10.1016/j.marmicro.2013.06.005
Chang, S., Feng, Q. L., Clausen, S., et al., 2017. Sponge Spicules from the Lower Cambrian in the Yanjiahe Formation, South China:The Earliest Biomineralizing Sponge Record. Palaeogeography, Palaeoclimatology, Palaeoecology, 474: 36-44. DOI:10.1016/j.palaeo.2016.06.032
Chen, P., 1984. Discovery of Lower Cambrian Small Shelly Fossils from Jijiapo, Yichang, West Hubei and Its Significance. Professional Papers of Stratigraphy and Palaeontology, 2: 49-65.
Cloud, P. E., 1968. Pre-Metazoan Evolution and the Origins of the Metazoa. In: Drake, E. T., ed., Evolution and Environment. Yale University Press, New Haven. 72
Conway Morris, S., Chen, M. G., 1990. Blastulospongia Polytreta N. Sp., an Enigmatic Organism from the Lower Cambrian of Hubei, China. Journal of Paleontology, 64(1): 26-30. DOI:10.1017/s0022336000042207
Danelian, T., Moreira, D., 2004. Palaeontological and Molecular Arguments for the Origin of Silica-Secreting Marine Organisms. Comptes Rendus Palevol, 3(3): 229-236. DOI:10.1016/j.crpv.2004.01.005
De Wever, P., Dumitrica, P., Caulet, J. P., et al., 2001. Radiolarians in the Sedimentary Record. Gordon and Breach Science Publishers, London. 525
Decelle, J., Suzuki, N., Mahé, F., et al., 2012. Molecular Phylogeny and Morphological Evolution of the Acantharia (Radiolaria). Protist, 163(3): 435-450. DOI:10.1016/j.protis.2011.10.002
Ding, R. X., Zou, H. P., Min, K., et al., 2017. Detrital Zircon U-Pb Geochronology of Sinian-Cambrian Strata in the Eastern Guangxi Area, China. Journal of Earth Science, 28(2): 295-304. DOI:10.1007/s12583-017-0723-y
Elicki, O., 1998. First Report of Halkieria and Enigmatic Globular Fossils from the Central European Marianian (Lower Cambrian, Görlitz Syn-cline, Germany). Rev. Espa. Gonzalo Vidal, (1): 51-64.
Guo, J. F., Li, Y., Li, G. X., 2014. Small Shelly Fossils from the Early Cambrian Yanjiahe Formation, Yichang, Hubei, China. Gondwana Research, 25(3): 999-1007. DOI:10.1016/
He, T., Ling, H., Chen, Y., et al., 2013. Geochemical Character and For-mation of Cherts from the Ediacaran Piyuancun Formation of Lantian Section in Xiuning, Southern Anhui. Geological Journal of China Universities, 19(4): 620-633. DOI:10.16108/j.issn1006-7493.2013.04.016
Hu, R., Li, S., Wang, W., et al., 2016. Source Characteristics of Tillite the Nantuo Formation in Three Gorges, Northern Yangtze Block:Evidence from Zricon Ages and Geochemical Composition. Earth Science, 41(10): 1630-1654. DOI:10.3799/dqkx.2016.121
Ishitani, Y., Ishikawa, S. A., Inagaki, Y., et al., 2011. Multigene Phyloge-netic Analyses Including Diverse Radiolarian Species Support the "Retaria" Hypothesis-The Sister Relationship of Radiolaria and Foraminifera. Marine Micropaleontology, 81(1/2): 32-42. DOI:10.1016/j.marmicro.2011.06.007
Jin, C. S., Li, C., Algeo, T. J., et al., 2016. Evidence for Marine Redox Control on Spatial Colonization of Early Animals during Cambrian Age 3 (c. 521-514 Ma) in South China. Geological Magazine, 154(6): 1360-1370. DOI:10.1017/S0016756816001138
Khomentovsky, V. V., Karlova, G. A., 1993. Biostratigraphy of the Vendian-Cambrian Beds and the Lower Cambrian Boundary in Siberia. Geological Magazine, 130(1): 29-45. DOI:10.1017/S0016756800000960
Klok, C. J., Hubb, A. J., Harrison, J. F., 2009. Single and Multigenerational Responses of Body Mass to Atmospheric Oxygen Concentrations In-Drosophila Melanogaster:Evidence for Roles of Plasticity and Evolu-tion. Journal of Evolutionary Biology, 22(12): 2496-2504. DOI:10.1111/j.1420-9101.2009.01866.x
Kouchinsky, A., Bengtson, S., Clausen, S., et al., 2013. An Early Cambrian Fauna of Skeletal Fossils from the Emyaksin Formation, Northern Si-beria. Acta Palaeontologica Polonica, 60(2): 421-512. DOI:10.4202/app.2012
Kouchinsky, A., Bengtson, S., Landing, E., et al., 2017. Terreneuvian Stratigraphy and Faunas from the Anabar Uplift, Siberia. Acta Palaeontologica Polonica, 62(2): 311-440. DOI:10.4202/app.00289.2016
Li, C., Jin, C. S., Planavsky, N. J., et al., 2017. Coupled Oceanic Oxygenation and Metazoan Diversification during the Early-Middle Cambrian?. Geology, 45(8): 743-746. DOI:10.1130/G39208.1
Li, R. W., Lu, J. L., Zhang, S. K., et al., 1999. Organic Carbon Isotopes of the Sinian and Early Cambrian Black Shales on Yangtze Platform, China. Science in China Series D:Earth Sciences, 42(6): 595-603. DOI:10.1007/bf02877787
Lipps, J. H., 1992. Proterozoic and Cambrian Skeletonized Protists. In: Schopf, J. W., Klein, C., eds., The Proterozoic Biosphere: A Multidisciplinary Study. Cambridge University Press, Cambridge. 237-240
Liu, K., Feng, Q. L., Shen, J., et al., 2017. Increased Productivity as a Primary Driver of Marine Anoxia in the Lower Cambrian. Palaeogeography, Palaeoclimatology, Palaeoecology, 491: 1-9. DOI:10.1016/j.palaeo.2017.11.007
Maletz, J., 2011. Radiolarian Skeletal Structures and Biostratigraphy in the Early Palaeozoic (Cambrian-Ordovician). Palaeoworld, 20(2/3): 116-133. DOI:10.1016/j.palwor.2010.12.007
Mason, R., Li, Y. J., Cao, K. N., et al., 2017. Ediacaran Macrofossils in Shunyang Valley, Sixi, Three Gorges District, Hubei Province, China. Journal of Earth Science, 28(4): 614-621. DOI:10.1007/s12583-017-0773-1
Nazarov, B. B., 1973. Radiolarians from the Lowermost Horizons of the Batenev Mountain Ridge. In: Problems of Paleontology and Biostra-tigraphy of the Lower Cambrian of Siberia and the Far East. Novosibirsk, Nauka. 5-13 (in Russian)
Nazarov, B. B., 1975. Lower and Middle Paleozoic Radiolarians of Ka-zakhstan (Methods of Investigation, Systematics and Stratigraphic Significance). In: Raaben, M. E., ed., Trudy Akademiya Nauk SSSR, Geologicheskii Institut. Izdatelstvo Nauka, Moscow. 1-203 (in Russian)
Obut, O. T., Iwata, K., 2000. Lower Cambrian Radiolaria from the Gorny Altai (Southern West Siberia). News of Paleontology and Stratigraphy, 2/3: 33-38.
Payne, J. L., Boyer, A. G., Brown, J. H., et al., 2009. Two-Phase Increase in the Maximum Size of Life over 3.5 Billion Years Reflects Biological Innovation and Environmental Opportunity. Proceedings of the Na-tional Academy of Sciences, 106(1): 24-27. DOI:10.1073/pnas.0806314106
Peng, L., 1984. The Age and Tectonic Significance of Ophiolites of the Undorsum Group, Nei Mongol Autonomous Region. Science Bulletin, 29(7): 936-939.
Peng, S. C., Babcock, L. E., Cooper, R. A., 2012. The Cambrian Period. In: Gradstein, F. M., Ogg, J. G., Schmitz, M. D., et al., eds., The Geologic Time Scale 2012, Vol. 2. Elsevier BV, Amsterdam. 437-488.
Pickett, J. W., Jell, P. A., 1983. Middle Cambrian Sphinctozoa (Porifera) from New South Wales. Memoirs of the Association of Australasian Paleontologists, 1: 85-92.
Pouille, L., Obut, O., Danelian, T., et al., 2011. Lower Cambrian (Botomian) Polycystine Radiolaria from the Altai Mountains (Southern Siberia, Russia). Comptes Rendus Palevol, 10(8): 627-633. DOI:10.1016/j.crpv.2011.05.004
Shu, D., Chen, L., 1989. Discovery of Early Cambrian Radiolarian and Its Significance. Science in China, 32(8): 986-994.
Steiner, M., Li, G., Qian, Y., et al., 2007. Neoproterozoic to Early Cambrian Small Shelly Fossil Assemblages and a Revised Biostratigraphic Correlation of the Yangtze Platform (China). Palaeogeography, Pal-aeoclimatology, Palaeoecology, 254(2): 67-99. DOI:10.1016/j.palaeo.2007.03.046
White, R. D., 1986. Cambrian Radiolaria from Utah. Journal of Paleontology, 60(3): 778-780. DOI:10.1017/s0022336000022307
Won, M. Z., Below, R., 1999. Cambrian Radiolaria from the Georgina Basin, Queensland, Australia. Micropaleontology, 45(4): 325-363. DOI:10.2307/1486119
Wrona, R., 2004. Cambrian Microfossils from Glacial Erratics of King George Island, Antarctica. Acta Palaeontologica Polonica, 49(1): 13-56.
Yang, B., Steiner, M., Li, G., et al., 2014. Terreneuvian Small Shelly Faunas of East Yunnan (South China) and Their Biostratigraphic Implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 398: 28-58. DOI:10.1016/j.palaeo.2013.07.003
Yin, H., Zeng, Y., Xia, W., 1994. Chert on the Southeast Continental Margin of the Yangtze Platform. Acta Geologica Sinica, 68(2): 132-141.
Yin, L. M., Wang, C. J., Zhao, Y. L., et al., 2016. Early-Middle Cambrian Palynomorph Microfossils and Related Geochemical Events in South China. Journal of Earth Science, 27(2): 180-186. DOI:10.1007/s12583-016-0689-1
Zhang, L., Danelian, T., Feng, Q. L., et al., 2013. On the Lower Cambrian Biotic and Geochemical Record of the Hetang Formation (Yangtze Platform, South China):Evidence for Biogenic Silica and Possible Presence of Radiolaria. Journal of Micropalaeontology, 32(2): 207-217. DOI:10.1144/jmpaleo2013-003
Zhang, M. Z., Peng, S. B., Zhang, L., et al., 2016. New Recognition of Carbonate Nodules Genesis in Sinian Doushantuo Formation in Zigui Area and Its Geological Implication. Earth Science, 41(12): 1977-1994. DOI:10.3799/dqkx.2016.138
Zhang, X. G., Aldridge, R. J., 2007. Development and Diversification of Trunk Plates of the Lower Cambrian Lobopodians. Palaeontology, 50(2): 401-415. DOI:10.1111/j.1475-4983.2006.00634.x
Zhang, X. L., Cui, L. H., 2016. Oxygen Requirements for the Cambrian Explosion. Journal of Earth Science, 27(2): 187-195. DOI:10.1007/s12583-016-0690-8
Zhang, X. L., Shu, D. G., Han, J., et al., 2014. Triggers for the Cambrian Explosion:Hypotheses and Problems. Gondwana Research, 25(3): 896-909. DOI:10.1016/
Zhao, G., 1999. The Influence of Biogenic Procession on the Accumulation and Precipiation of Silica-An Example from South of Anhui and West of Zhejiang. Acta Sedimentologica Sinica, 17(1): 30-37. DOI:10.1427/j.cnki.cjxb.1999.01.005
Zheng, N., Song, T., Li, Y., et al., 2012. The Discovery of the Lower Cambrian and Middle Ordovician Radiolaria in the South China Orogenic Belt. Geology in China, 39(1): 260-265.
Zhou, C. M., Jiang, S. Y., 2009. Palaeoceanographic Redox Environments for the Lower Cambrian Hetang Formation in South China:Evidence from Pyrite Framboids, Redox Sensitive Trace Elements, and Sponge Biota Occurrence. Palaeogeography, Palaeoclimatology, Palaeoecology, 271(3/4): 279-286. DOI:10.1016/j.palaeo.2008.10.024