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Phylogenetic position of every species between Naraoia and Misszhouia. Numbers refer to derived characteristics as follows: 1-genal spines; 2-subcircular cephalic shield, 3-circular cephalic shield, 4-round genal angle, 5-with marginal spines of the posterior shield; 6-elongate posterior shield, 7-long elliptic posterior shield, 8-wide elliptic posterior shield, 9-single spine of the posterior shield rear margin; 10-one pair spines of posterior shield rear margin; 11-wide doublure (based on Caron et al. 2004, fig. 5, modified).

Phylogenetic position of every species between Naraoia and Misszhouia. Numbers refer to derived characteristics as follows: 1-genal spines; 2-subcircular cephalic shield, 3-circular cephalic shield, 4-round genal angle, 5-with marginal spines of the posterior shield; 6-elongate posterior shield, 7-long elliptic posterior shield, 8-wide elliptic posterior shield, 9-single spine of the posterior shield rear margin; 10-one pair spines of posterior shield rear margin; 11-wide doublure (based on Caron et al. 2004, fig. 5, modified).

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Naraoia is a taxon represented by commonly non-mineralized fossil found in lower middle Cambrian strata and widely distributed in China, North America, and is an important component of Burgess Shale-type biotas. Well-preserved representatives of Naraoia are known from the Chengjiang and Kaili biotas of South China. Here, we report a new species of...

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... The formation is a heterogeneous unit but is dominated by greenish-grey shale and calcareous mudstone shale with intercalations of thin-bedded argillaceous carbonates. Abundant fossil assemblages are excavated from the middle to upper part of the Balang Formation at different localities (Peng et al., 2005;Peng et al., 2010;Peng, Zhao & Sun, 2012;Peng et al., 2016;Sun et al., 2014;Shen et al., 2016;Wen et al., 2019;Liu et al., 2018). The biostratigraphic framework of the entire Balang Formation belongs to Arthricocephalus chauveaui trilobite Zone dated to the global Cambrian Series 2, Stage 4 (Yan et al., 2014). ...
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Moulting behaviors in trilobites are a crucial strategy during development. Previous studies have demonstrated inter-and intraspecific variability of moulting behavior in trilobites. Currently, ecdysial motifs for trilobites are considered not stable even within species and fewer detailed studies dealt with moulting behaviors in a single species of trilobite during development. Here a large sample of meraspid to holaspid exuviae of Arthricocephalites xinzhaiheensis (131 specimens) from the Cambrian Balang Formation of South China has allowed description of the reasonably complete ontogenic moulting sequence. Both ontogenetic stage and body size reveal gradual transition of configuration from Somersault configuration to Henningsmoen’s configuration during development. Somersault configuration is exclusive till meraspid degree five and exists in subsequent growth stages. This suggests that opening of the facial and rostral sutures allowing the emergence forward of the post-ecdysial trilobite was prevalent in early growth stages. In later development, Henningsmoen’s configuration (showing disarticulation of the cranidium) became more dominant. This study indicates that gradual transition of ontogenetic moulting behavior occurred in oryctocephalid trilobites in the early Cambrian.
... The Balang Formation is dominated by greenish-grey shale and calcareous mudstone shale with intercalations of thin-bedded argillaceous carbonates (Fig. 1C). Abundant fossil assemblages have been excavated from the middle to upper part of the Balang Formation at different localities (Peng et al. 2005(Peng et al. , 2010(Peng et al. , 2012Sun et al. 2014;Shen et al. 2016;Wen et al. 2017;Liu et al. 2018a,b). The biostratigraphical framework of the entire Balang Formation belongs to Arthricocephalus chauveaui trilobite Zone of the global Cambrian Series 2, Stage 4 (Yan et al. 2014). ...
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It is important to understand moulting behaviours in trilobites through different growth stages. Most studies have focused on patterns of moulting in trilobites based on a few exuviae. The present study is based on 177 disarticulated specimens of Arthricocephalites xinzhaiheensis from the Balang Formation (Cambrian Series 2, Stage 4, Guizhou Province, South China). The abundance of disarticulated assemblages allows the distinction between carcasses and exuviae to be made with confidence and enables a detailed exploration of moulting behaviours. About 96.6% of exuviae of both meraspides and holaspides of Ar. xinzhaiheensis show a disarticulated cranidium characterizing Henningsmoen's configuration. The trilobites commonly used the cephalic sutures and cephalothoracic joint for moulting, involving lateral rotation or inversion of the cranidium to create a wide exuvial gape. Only one specimen displays the Somersault configuration characterized by an overturned lower cephalic unit and the data also show that trilobites employing the cephalic sutures to create an anterior exuvial gape are rare in holaspides. Though some specimens are similar to Salter's configuration with an inverted cephalon with respect to the thoracopygon, the specimens show closed cephalic sutures and thus dissociated or inverted cephala are likely to represent disturbed carcasses. Several other species of oryctocephalid trilobites display Henningsmoen's configuration indicating that exuviation by disarticulated cranidium to enlarge the exuvial gape was prevalent in the family from Cambrian Series 2 Stage 4 to the succeeding Wuliuan Stage of the Miaolingian Series.
... Many fossil assemblages have been discovered from the middle to upper parts of the Balang Fm. at different localities (Peng et al., 2005(Peng et al., , 2012a(Peng et al., , 2012bPeng, 2009;Ma et al., 2011;Liu, 2018;Liu et al., 2018a); the Balang Fauna comprises at least 80 species of 63 genera (Liu, 2018;Liu et al., 2018b), including chancelloriids, cnidarians, sponges, worms, brachiopods, echinoderms, molluscs, hyoliths, arthropods, algae and many trace fossils (Peng et al., 2006(Peng et al., , 2010a(Peng et al., , 2010b(Peng et al., , 2012cFu et al., 2010;Qin et al., 2010;Liu, 2013aLiu, , 2013bLiu, , 2018Zhao et al., 2015;Sun et al., 2015Sun et al., , 2016Sun et al., , 2017Shen et al., 2016;Liang et al., 2017;Wen et al., 2017, in press;Liu et al., 2018a;Wang et al., 2018). There are currently 163 sponge specimens (including the 36 specimens collected in 2018) collected from the highly fossiliferous Lazizhai section at Lazizhai Village, Jianhe County, Kaili City, Guizhou, China ( Fig. 1(A)). ...
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... The Balang Formation is exposed in eastern Guizhou and western Hunan, China (Figure 1 Many fossil assemblages are discovered from the middle to upper part of the Balang Formation at different localities (Peng et al. 2005(Peng et al. , 2012a(Peng et al. , 2012bPeng 2009;Ma et al. 2011;Zhao et al. 2015;Liu et al. 2018a) and the Balang Fauna comprises at least 80 species of 63 genera (Liu 2018;Liu et al. 2018b), including algae, chancelloriids, cnidarians, sponges, worms, brachiopods, echinoderms, molluscs, hyoliths and arthropods, as well as many trace fossils (Peng et al. 2006(Peng et al. , 2007(Peng et al. , 2010a(Peng et al. , 2010bPeng et al. 2012cPeng et al. , 2016Fu et al. 2010;Liu 2013aLiu , 2013bSun et al. 2015Sun et al. , 2016Sun et al. , 2017Zhao et al. 2015;Liang et al. 2017;Wen et al. 2017;Liu et al. 2018a). Since there is no isotopic dating available for the Balang Formation, its age is regarded as Stage 4 based on a series of trilobitebased correlations of Cambrian strata between South China and other continents Peng and Babcock 2001;Yuan et al. 2001Yuan et al. , 2006Yuan et al. , 2011Peng 2009;Qin et al. 2010;Yan et al. 2014;Yuan and Ng 2014;Shen et al. 2016). ...
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The genus Naraoia Walcott, 1912, a Burgess Shale-type fossil known from the lower and middle Cambrian of British Columbia (Canada), Idaho and Utah (USA), as well as from Yunnan and Guizhou provinces (China), is now reported from the middle Cambrian of Murero (Zaragoza, Spain), which is the first record in the Acadobaltic province. The only fragmented specimen found is determined as Naraoia sp., its age being Pardailhania multispinosa Zone (Drumian Stage). This new datum reinforces the hypothesis of the existence of a cosmopolitan faunal substrate in early Cambrian times, which is to some extent refl ected in the mid Cambrian by faunal groups of low evolutionary potential as the family Naraoiidae and other soft-bodied fossil taxa.
... Naraoia taijiangensis Peng, Zhao, et al. 2012, holotype GUT GY-172-56. This fossil was found in the Geyi section of the Balang Formation, near Geyi Town, Taijiang, Guizhou, south China (Peng, Zhao, et al. 2012). ...
... Naraoia taijiangensis Peng, Zhao, et al. 2012, holotype GUT GY-172-56. This fossil was found in the Geyi section of the Balang Formation, near Geyi Town, Taijiang, Guizhou, south China (Peng, Zhao, et al. 2012). The Geyi section corresponds to the Arthricocephalus chauveaui-Changaspis elongata trilobite Biozone (Peng 2005;Peng, Zhao, et al. 2012). ...
... This fossil was found in the Geyi section of the Balang Formation, near Geyi Town, Taijiang, Guizhou, south China (Peng, Zhao, et al. 2012). The Geyi section corresponds to the Arthricocephalus chauveaui-Changaspis elongata trilobite Biozone (Peng 2005;Peng, Zhao, et al. 2012). The trilobite Arthricocephalus chauveaui indicates the Balang Formation belongs to the early Cambrian Series 2, Stage 4 , specifically the Duyunian regional Stage. ...
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Wolfe 2017. Metamorphosis Is Ancestral for Crown Euarthropods, and Evolved in the Cambrian or Earlier. Supplementary Data 1. List of node calibrations, list of tip calibrations, specimen information, age details, and supplementary references.
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... The Balang Fauna, a Burgess Shale-type biota from the lower Cambrian of China (Peng et al. 2005), also yields representatives of this group (Peng et al. 2010;Peng, Zhao & Sun, 2012). These fossils are primarily known from the very fossiliferous Geyi Section of the Balang Formation in Geyi Town, Taijiang County, where the section contains the best examples of the Balang Fauna (Peng, Zhao & Sun, 2012). These fossils display a cylindrical body with clear annulations densely separated by intersegment furrows and possessing a cuticle consisting of numerous, tiny, individual button-shaped plates on each annulation. ...
... The underlying Bianmachong Formation is mainly composed of grey-black mudstone, but it changes to yellowgreyish shale southwards (Yin, 1987). The previous biostratigraphic framework was described as a single genus-level assemblage zone (Zhou & Yuang, 1980;Yin, 1987), considered first to represent up to four species-level zones (Yuan, Zhao & Li, 2001;Yuan, Zhao & Yang, 2006), but subsequent work recognized only one species-level assemblage zone (Peng, 2009;Peng et al. 2010;Peng, Zhao & Sun, 2012). Most recently, a single species-level zone, that is the Arthricocephalus chauveaui Zone, has been proposed (Yan et al. 2014). ...
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Wronascolex is a taxon of palaeoscolecids. It is commonly represented by isolated button-like microfossils or compressed individuals, which are found worldwide in strata ranging from the Cambrian Series 2 to Series 3. The earliest representative of Wronascolex is known from the Sinsk Formation of the lower Cambrian of the Siberian Platform. Other species occur in Burgess Shale-type biotas of Cambrian age from Australia, Spain and North America. New palaeoscolecid material from the Balang Fauna of the Cambrian Series 2 of eastern Guizhou represents a new species of Wronascolex , W. geyiensis sp. nov., and extends the geographic and stratigraphic distribution of Wronascolex , as well as providing additional evidence for understanding its morphology based upon correlation of body configuration and cuticular ornaments.
... After years of quarrying, the unit has yielded a deposit of diverse taxa known as the Balang Biota, which contains thousands of articulated specimens of Guizhoueocrinus yui Zhao, Parsley and Peng, 2007a. With additional discoveries of soft-bodied arthropods, such as Naraoia (Yang et al., 2011;Peng et al., 2012a) and vermiform taxa (Peng et al., 2012b), the Balang Biota is undoubtedly a classic Burgess Shale-type deposit. ...
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A new eocrinoid locality of the Balang Formation (Cambrian Series 2) near Kaili City is reported. The fauna is associated with index trilobites, such as Redlichia (Pteroredlichia) murakamii Resser and Endo in Kobayashi, 1935 and Arthricocephalus chauveaui Bergeron, 1899, that are common in the Balang Biota (Cambrian Series 2) but absent in the younger Kaili Biota (Cambrian Series 3). This new locality contains a new eocrinoid fauna (n = 22) that is different from Guizhoueocrinus yui Zhao, Parsley and Peng, 2007a in bearing a smaller theca, a shorter stalk, and a robust attachment disk; thus, a taxon Globoeocrinus zhaoyuanlongensis n. sp. is proposed.
... The Balang Formation (Cambrian Stage 4) of eastern Guizhou, China, is an important Burgess Shale-type deposit that yields a diverse assemblage of biomineralized and nonbiomineralized fossils (Peng et al., 2005(Peng et al., , 2012. Most previous work has focused on the systematics and evolutionary history of the Balang Biota. ...
... 1), and its total thickness is the greatest near the center of the outcrop area (in the slope belt). It comprises a shallowing-upward sequence toward its contact with the overlying Tsinghsutung Formation (Peng et al., 2012). Zhou et al. (1979) first provided a biostratigraphic framework for the Balang Formation, and that work has been revised several times. ...
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The Balang Formation (Cambrian Stage 4) of eastern Guizhou, China, yields slender, sinuous, and branching ichnofossils associated with shells of hyolithids. The trace-makers, of speculative identity, are inferred to have been feeding on decaying soft parts of hyolithids and on the microbial halos formed around decaying bodily remains. The comprehensive observation of hyolithid shells and their associated traces suggests that the hyolithid shells may have been moved some distance after scavenging began. This preservational pattern can address some issues relating to hyolith preservation of the Balang Formation.
... In some respects, a comparable morphology of the digestive system, consisting of a midgut connected with a large, ramifying anterior pair of digestive diverticula u n c o r r e c t e d p r o o f s in the cephalon, followed by three pairs of much smaller cephalic diverticulae and at least ten pairs of trunk diverticulae has been described in Cambrian naraoids (e.g., whittington 1993; Chen et al. 1997;vannier & Chen 2002;lin 2006;BergströM et al. 2007;Zhang et al. 2007;Peng et al. 2012;ortega-hernándeZ & Brean 2012). Morphologically comparable cephalic diverticulae were described also in another nonbiomineralizing arthropod Burgessia (hughes 1975;ButterField et al. 2007;lin 2009). ...
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Remains of the digestive system are described in the holotype of the rare harpidid trilobite Harpides grimmi Barrande, 1852 collected from the Lower Ordovician Mílina Formation of the Prague Basin. The intestine (post-stomach alimentary canal) starts just behind the glabellar posterior margin and extends through the narrow axial region in all nineteen thoracic segments of this exceptionally preserved specimen. The anterior-most part of the digestive system is masked by the in situ hypostome preserved under the missing glabella. Similarly, also the posterior-most part of the post-stomach alimentary canal is absent, as the pygidium is not preserved. This specimen constitutes the first example of preserved digestive structures in the family Harpididae. Earlier finds of digestive system in Ordovician trilobites are briefly assessed.