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Paterimitra Pyramidalis Laurie, 1986, the First Tommotiid Discovered From

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Paterimitra Pyramidalis Laurie, 1986, the First Tommotiid Discovered From 1 Paterimitra pyramidalis Laurie, 1986, the first tommotiid discovered from 2 the early Cambrian of North China 3 4 Bing Pana, b, Glenn A. Brockc, Christian B. Skovstedd, Marissa J. Bettse, Timothy P. Topperf, 5 Guo-Xiang Lia, * 6 7 a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and 8 Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China 9 b University of Science and Technology of China, Hefei 230026, China 10 c Department of Biological Sciences, Macquarie University, NSW 2109, Australia 11 d Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden. 12 e Palaeoscience Research Centre, School of Environmental and Rural Science, University of 13 New England, Armidale, NSW, Australia. 14 f Palaeoecosystems Group, Department of Earth Sciences, Durham University, Durham, UK. 15 * Corresponding author. 16 E-mail: [email protected] (B. Pan), [email protected] (G.A. Brock), 17 [email protected] (C.B. Skovsted), [email protected] (M.J. Betts), 18 [email protected] (T.P. Topper), [email protected] (G.X. Li) 19 20 ABSTRACT 21 The eccentrothecimorph tommotiid Paterimitra pyramidalis Laurie, 1986, was 22 previously only known from lower Cambrian rocks of the Northern Territory and South 23 Australia. Herein, we document the first occurrence of P. pyramidalis from the Xinji 24 Formation in the Shuiyu section at Ruicheng County, Shanxi Province, located at the 25 southwestern margin of the North China Platform. This represents the first report of a 1 26 tommotiid taxon from lower Cambrian strata of the North China Platform. All three sclerite 27 types that characterise the scleritome of P. pyramidalis have been recovered and are described, 28 permitting definitive identification to species level. The discovery of P. pyramidalis from the 29 North China Platform not only greatly extends the palaeogeographic range of this distinctive 30 tommotiid taxon, but also supports planktotrophic development of larvae in Paterimitra as a 31 stem group brachiopod. The discovery of P. pyramidalis supports a Cambrian, Epoch 2, late 32 Age 3 to early Age 4 age for the shelly fossil fauna from the Xinji Formation and indicates a 33 close palaeogeographic position between the North China Platform and Australian East 34 Gondwana during the early Cambrian. 35 36 Key words: Tommotiida, Paterimitra, North China, Cambrian Epoch 2, biostratigraphy, 37 palaeobiogeography 38 39 1. Introduction 40 The eccentrothecimorph tommotiids Paterimitra pyramidalis Laurie, 1986, 41 Eccentrotheca helenia Skovsted, Brock, Topper, Paterson and Holmer, 2011 and Kulparina 42 rostrata Conway Morris and Bengtson in Bengtson et al., 1990 are critical taxa for 43 interpreting the timing of character assembly and bivalved body plan evolution in 44 organophosphatic brachiopods (Skovsted et al., 2009, 2011, 2015; Larsson et al., 2014). 45 Identical mineralogical composition, external ornamentation and distinctive organic 46 polygonal ultrastructure shared between P. pyramidalis and early Cambrian paterinid 47 brachiopods such as Askepasma Laurie, 1986 (Balthasar et al., 2009; Topper et al., 2013) and 48 Salanygolina Ushatinskaya, 1987 (Holmer et al., 2009, 2011) led Larsson et al. (2014) to 49 suggest a homologous relationship between these taxa. The bivalved brachiopod shell evolved 50 through successive stages of sclerite reduction, specialisation and tube shortening from an 2 51 Eccentrotheca-like stem group ancestor through intermediary forms such as Paterimitra 52 pyramidalis and Micrina etheridgei Laurie, 1986 (Holmer et al., 2008; Skovsted et al., 2008, 53 2009, 2011, 2015; Larsson et al., 2014). 54 The original description of Paterimitra pyramidalis by Laurie (1986) from the Todd 55 River Dolostone, Amadeus Basin in central Australia was based on 10-12 specimens of a 56 large pyramidal sclerite type, subsequently recognised as the S1 sclerite by Skovsted et al. 57 (2009). Large collections comprising more than 1400 isolated sclerites from rocks of the 58 Hawker Group in the Arrowie Basin, South Australia have revealed that this taxon bore three 59 sclerite morphotypes, the S1, S2 and L sclerites (Skovsted et al., 2009, figs. 1, 2; Larsson et 60 al., 2014, fig. 2). Significantly, partially articulated scleritome composites indicate that P. 61 pyramidalis secreted a cone-shaped skeleton consisting of a single pair of conjoined 62 bilaterally symmetrical S1 and S2 sclerites, and an unresolved number of laterally compressed 63 L sclerites fused around the distal margin of the open cone formed by the S1-S2 composite. 64 An organic, pedicle-like structure for epifaunal attachment is interpreted to have emerged 65 through the foramen-like posterior opening formed between the S1 and S2 sclerites. The 66 cone-like scleritome is assumed to have housed a filter-feeding lophophorate organism (see 67 Larsson et al., 2014, fig. 21 for reconstruction). Similar fixed epifaunal scleritomes have been 68 demonstrated for Eccentrotheca helenia (Skovsted et al., 2011) and Kulparina rostrata 69 (Skovsted et al., 2015). 70 Like many tommotiids, Paterimitra pyramidalis had been regarded as an East 71 Gondwanan endemic (Skovsted et al., 2011, 2015; Larson et al., 2014). Here we describe P. 72 pyramidalis from the Xinji Formation in the Shanxi Province, greatly extending the 73 palaeogeographic range of this taxon. This is the first tommotiid ever reported from the North 74 China Platform, which has significant implications for understanding stem group brachiopod 75 larval dispersal patterns, biostratigraphic correlation between Australia and North China, and 3 76 for testing a wide range of contradictory interpretations regarding the palaeogeographic 77 position of the North China Platform and Cambrian East Gondwana. 78 79 2. Materials and methods 80 All isolated sclerites of Paterimitra pyramidalis from the Xinji Formation are more or 81 less fragmentary, but micro-ornament and other diagnostic characters are generally well 82 preserved. Many specimens are encrusted with adhering quartz grains. Whilst some grains 83 have been carefully removed, it is not possible to remove all of them without the risk of 84 damaging the specimens. In total, 22 specimens have been recovered including 6 fragments of 85 S1 sclerites, 1 S2 sclerite and 15 L sclerites. 86 All samples were treated in 6% acetic acid. Selected specimens were placed on pin-type 87 stubs, gold coated and photographed using the Scanning Electron Microscopy facility (LEO 88 1530VP) at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of 89 Sciences (NIGPAS), Nanjing. All illustrated specimens are housed and cataloged at storage 90 facilities of NIGPAS. 91 92 3. Geological setting 93 3.1 Locality and stratigraphy 94 All specimens of Paterimitra pyramidalis are derived from the Xinji Formation, located 95 at Ruicheng County, Shanxi Province. (Fig. 1A). The Xinji Formation represents the 96 lowermost Cambrian strata in this region, disconformably overlying Precambrian strata. The 97 Xinji Formation consists of siliciclastic rocks intercalated with carbonates, and mainly crops 98 out on the southwestern to southern margin of the North China Platform (Liu et al., 1991, 99 1994). The thickness of the Xinji Formation decreases from south to north, along the southern 100 margin of the North China Platform (Liu, 1986; Liu et al., 1991; Yan et al., 1993). In the 4 101 Shuiyu section, the Xinji Formation disconformably overlies Precambrian strata; either the 102 Luoquan Formation, or the Longjiayuan Formation (Palaeoproterozoic or Mesoproterozoic, 103 gray dolostones) (Fig. 1B). A karst surface is developed between the Luoquan Formation and 104 the Longjiayuan Formation. The Xinji Formation at the Shuiyu section is about 39.8 m thick, 105 and can be subdivided into two parts: The lower part, which is dominated by grey-black 106 phosphatic conglomerates, purple-red shale and phosphatic sandstone, and the upper part, 107 which consists of red quartz sandstone intercalated with argillaceous dolostone (Bureau of 108 Geology and Mineral Resources of Shanxi Province, 1989). The basal 10.7 m of the section 109 was sampled for shelly fossils since the upper part of the section is covered by alluvium and 110 vegetation. There are abundant trace fossils in the lowermost rocks, and the trace fossil 111 assemblage belongs to a typical Cruziana ichnofacies (Miao and Zhu, 2014), indicating a 112 subtidal environment for the basal Xinji Formation at the Shuiyu section. The brachiopod 113 Kutorgina sinensis Rong in Lu 1979 had been reported from the lower part of this section (Lu, 114 1979, pl. Ⅴ, figs. 9-11). Paterimitra was collected from a strongly weathered bed of calcitic 115 phosphatic quartz siltstone, located 3.4-3.8 m above the base of the Xinji Formation (Fig. 1B). 116 117 3.2. Biostratigraphy 118 The Xinji Formation and the coeval Houjiashan Formation along the southern margin of 119 North China Platform have both yielded a trilobite fauna with low diversity, consisting of 120 Estaingia (Bergeroniellus) lonanensis Hsiang in Lu et al., 1965 (Luonan County and Huoqiu 121 County), Estaingia (Hsuaspis) houchiuensis Chang in Hsiang, 1963 (in Luonan County and 122 Huoqiu County), and Redlichia cf. R. nanjiangensis Zhang and Lin in Lee, 1978 (in Queshan 123 County) (Zhang and Zhu, 1979; Zhang et al., 1979; Miao, 2014). Based on the trilobite 124 assemblage, the Xinji Formation was correlated with the Drepanuloides Biozone of the 125 middle Tsanglangpuan stage (Cambrian Stage 4) on the Yangtze Platform (Zhang et al., 1979; 5 126 Zhang and Zhu, 1979; He et al., 1984; He and Pei, 1985; Miao, 2014). The Estaingia trilobite 127 assemblage from the Xinji Formation and the Houjiashan Formation of North China was also 128 correlated with the Pararaia janeae trilobite Zone (Stage 4) assemblage of South Australia by 129 Miao (2014). 130 In South Australia, Redlichia occurs in Stage 3 (Pararaia bunyerooensis Zone in 131 Paterson and Brock, 2007; Dailyatia odyssei Zone of Betts et al., 2016) rather than only in 132 Stage 4 as in South China.
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