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Gondwana Research 19 (2011) 310326

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Gondwana Research
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / g r

The oldest bivalved arthropods from the early Cambrian of East Gondwana: Systematics, biostratigraphy and biogeography
Timothy P. Topper a,, Christian B. Skovsted b,c, Glenn A. Brock a, John R. Paterson d
a

Department of Biological Sciences, Macquarie University, NSW 2109, Australia Department of Earth Sciences, Uppsala University, Villavgen 16, SE-752 36 Uppsala, Sweden Department of Palaeozoology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden d Division of Earth Sciences, School of Environmental & Rural Science, University of New England, Armidale NSW 2351, Australia
b c

a r t i c l e

i n f o

a b s t r a c t
The oldest bradoriid fauna from Australia, occurring in the lower Cambrian Ajax and Wirrapowie limestones of the Flinders Ranges, South Australia consists of eleven taxa, including one new genus and species, Quadricona madonnae gen. et sp. nov. and two new species, Liangshanella circumbolina sp. nov. and Zepaera jagoi sp. nov. In the Ajax Limestone, Liangshanella circumbolina sp. nov. occurs c. 20 m below the FAD of the zonal trilobite Abadiella huoi. This pre-trilobitic occurrence represents the oldest bivalved arthropod hitherto known from East Gondwana and suggests a lower Cambrian (Series 2, Stage 3) age for the assemblage. The recognition of distinct bradoriid assemblages associated with the Abadiella huoi (Atdabanian), Pararaia tatei, P. bunyerooensis and P. janeae (all Botoman) trilobite biozones in South Australia indicates great potential for future regional biostratigraphic correlation. Quantitative biogeographic analysis including new taxonomic data from the lower Cambrian of South Australia, highlights the strong endemism displayed by early Cambrian bradoriid communities and strengthens the close faunal afnities with South China and Antarctica. 2010 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

Article history: Received 15 February 2010 Received in revised form 30 April 2010 Accepted 23 May 2010 Available online 4 June 2010 Keywords: Cambrian Arthropoda Bradoriida Phosphatocopida South Australia

1. Introduction Bradoriids are small, bivalved, marine arthropods that formed an important component of Cambrian faunal assemblages before their extinction in the middle Ordovician (e.g. Melnikova et al., 1997; Siveter and Williams, 1997; Hou et al., 2002; Vannier et al., 2005; Zhang, 2007; Jones and Kruse, 2009). The group had a worldwide distribution and are present in all of the major Cambrian Lagersttten, including the Buen Formation (Siveter et al., 1996), Qiongzhusi Formation (Hou and Bergstrm, 1991; Hou et al., 2002), Burgess Shale Formation (Conway Morris, 1986; Siveter and Williams, 1997) and the Alum Shale Formation (e.g. Mller, 1964, 1979, 1982; Maas et al., 2003). Historically, bradoriids were suggested to be the ancestors of ostracods (Sylvester Bradley, 1961; Hinz-Schallreuter, 1993a, b, 1999; Zhang and Pratt, 1993; McKenzie et al., 1999; Gozalo and Hinz-Schallreuter, 2002), but investigation of appendage morphology in exceptionally preserved specimens of Kunmingella Huo, 1956 and Kunyangella Huo, 1956 from China (Hou et al., 1996, 2002; Shu et al., 1999; Hou et al., 2010) suggests that bradoriids share few synapomorphies with

Corresponding author. Tel.: +61 2 9850 7719; fax: +61 2 9850 6053. E-mail address: timothy.topper@mq.edu.au (T.P. Topper).

ostracods and are also not directly related to the other widespread group of Cambrian bivalved arthropods, the Phosphatocopida (see also Hinz-Schallreuter and Schallreuter, 2009). Current evidence suggests phosphatocopids are a sister group to the Eucrustacea, a group that includes all crustacean taxa with extant derivatives (Maas et al., 2003; Jones and Laurie, 2006; Williams et al., 2007, 2008). The possession of ve head appendages in Kunmingella, accompanied by endopods that consist of fewer than seven podomeres indicate that bradoriids most probably represent stem group crustaceans (Hou et al., 1996, 2010; Shu et al., 1999; Williams et al., 2007, 2008). A total of 41 bradoriid and 12 phosphatocopid genera have been described from latest early and middle Cambrian [Series 2 (Stage 4) and Series 3 (Stage 5 to Guzhangian)] equivalent successions across central and northern Australia (pik, 1961, 1963, 1967, 1968; Fleming, 1973; Jones and McKenzie, 1980; Hinz, 1991, 1992a,b, 1993; Hinz and Jones, 1992; Hinz-Schallreuter, 1993a, 1999; Hinz-Schallreuter and Jones, 1994; Jones and Laurie, 2006; Jones and Kruse, 2009). In stark contrast, the diversity and abundance of bivalved arthropods from the thick lower Cambrian successions of South Australia have, until recently, been seriously neglected. The authors have recently documented 13 bradoriid and a single phosphatocopid species from the lower Cambrian (Series 2, Stage 4) Mernmerna Formation in the Arrowie Basin (Skovsted

1342-937X/$ see front matter 2010 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.gr.2010.05.012

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et al., 2006; Topper et al., 2007) adding substantially to the diversity of bivalved arthropods previously documented from the Parara Limestone in the Stansbury Basin (Chapman, 1918; Bengtson et al., 1990). Bivalved arthropods co-occur with the rst trilobites on several continents (Williams et al., 2007), but appear slightly earlier than the rst recorded trilobite (Abadiella) in southern China (Hou et al., 2002; Williams et al., 2007; Zhang, 2007; Zhang, et al., 2008). The faunal abundance and early stratigraphic occurrence of bivalved arthropods in South China has prompted some authors (Hou et al., 2002; Williams et al., 2007) to suggest that bradoriids may have originated in the South China region. The oldest bradoriids previously documented from South Australia were possible svealutids from the lower Cambrian (Atdabanian equivalent) Parara Limestone at Curramulka Quarry on Yorke Peninsula (Chapman, 1918). Studies of small shelly fossil (SSF) assemblages from the Parara Limestone on Yorke Peninsula have also established the presence of species belonging to the Hipponicharionidae (Albrunnicola bengtsoni Hinz-Schallreuter, 1993a) and Monasteriidae (Epactridion portax Bengtson in Bengtson et al., 1990). These bradoriids occur within the Abadiella huoi trilobite Biozone, which broadly correlates with the Abadiella trilobite Zone in South China (Jell in Bengtson et al., 1990; Steiner et al., 2001; Zang et al., 2001; Paterson and Brock, 2007) and the Atdabanian Stage of Siberia (Zhuravlev and Gravestock, 1994; Zhuravlev, 1995). The oldest phosphatocopid in Australia (Indianidae gen. et sp. indet. A) has also been documented from the Abadiella huoi Biozone in the Ajax Limestone, South Australia (Bengtson et al., 1990). Detailed collections along measured stratigraphic sections through the Ajax Limestone reveal that the oldest bradoriid occurs c. 20 m below the rst appearance datum (FAD) of the zonal trilobite Abadiella huoi. This pre-trilobitic occurrence suggests an early Series 2, Stage 3 age and represents the oldest bivalved arthropod hitherto known from the lower Cambrian of East Gondwana. In a further contribution to the understanding of early Cambrian bivalved arthropod biodiversity from East Gondwana, three new bradoriid species are described herein from the lower Cambrian Ajax Limestone (Series 2, Stage 3) in the Mount Scott Range and the upper part of the Wirrapowie Limestone (Series 2, Stages 3-4) in the Elder Range. The new data, when combined with the recent documentation of a number of South Australian bivalved arthropod assemblages (e.g. Skovsted et al., 2006; 2009; Topper et al., 2007) provides the opportunity to shed new light on the biogeography, biodiversity and evolution of early Cambrian South Australian assemblages. A quantitative analysis of Australian and East Gondwanan bradoriid biogeography is provided, using a pair-group cluster analysis for presence-absence data (Raup-Crick similarity index) utilising the statistical package PAST (Hammer et al., 2001). 2. Localities, biostratigraphy and age 2.1. Localities The bivalved arthropods described herein were collected from two lower Cambrian (Series 2, Stages 3-4) stratigraphic sections in the Arrowie Basin (Figs. 1, 2). The rst stratigraphic section (AJXM) was measured through the Ajax Limestone outcropping in the Mount Scott Range, northern Flinders Ranges, South Australia (Figs. 1B, 2A). The base of the AJX-M section is located at coordinates 3035'49" S, 13819'59.3" E [WGS84] and is equivalent to section M of Gravestock (1984, g. 2). The stratigraphy and sedimentology of the Ajax Limestone at section AJX-M has been summarised by Brock et al. (2006) and Skovsted et al. (2009); the latter paper documents specimens of the early Cambrian stem group brachiopod Mickwitzia from the upper part of the section.

The carbonate-dominated Ajax Limestone is approximately 280 m thick at AJX-M and conformably overlies the siliciclastic Parachilna Formation (Fig. 2A). The lower 120 m of AJX-M section is dominated by massive stromatolitic boundstones and dolomitised bioclastic and cryptalgal, laminated limestones. Shelly fossils are generally absent from this part of the section. The upper 160 m of section consists of richly fossiliferous (frequently silicied) beds including grey to buff coloured, thin nodular limestone beds and red, massive, bioclastic limestone with minor nodular interbeds (Skovsted et al., 2009). The second section, ER-9, was measured through the Wirrapowie Limestone and conformably overlying Mernmerna Formation outcropping on the eastern limb of a north plunging syncline in the northern Elder Range (Figs. 1C, 2B). The base of the ER-9 section is at coordinates 3140'2.6"S; 13826'11.5"E [WGS84]. The upper 47 m of massive, algal dominated, mottled and ribboned carbonates of the Wirrapowie Limestone was sampled at section ER-9, with sampling continuing into the Mernmerna Formation. The conformably overlying Mernmerna Formation reaches a maximum thickness of 191 m in ER-9 and is dominanted by argillitic to nodular limestones interbedded with calcareous mudstone in the lower two-thirds of the exposure, grading into a calcareous mudstone with minor limestone nodules in the upper 60 m of outcrop. The majority of bivalved arthropod specimens are derived from the upper Wirrapowie Limestone; only a few specimens are present in the overlying Mernmerna Formation (Fig. 2B).

2.2. Biostratigraphy and age The International Submission on Cambrian Stratigraphy (ISCS) has recently adopted a four-Series, ten-Stage framework for the Cambrian chronostratigraphic timescale (Babcock et al., 2005; Babcock and Peng, 2007; Peng and Babcock, 2008; Babcock et al., 2009). Whilst the majority of stages are undened and key horizons are yet to be resolved (Babcock and Peng, 2007), two Series (Terreneuvian and Furongian) and four stages (Fortunian, Drumian, Guzhangian and Paibian) have been ratied (Babcock et al., 2009). The traditional lower Cambrian has been replaced by two Series and four Stages, with only the base of the Cambrian system, currently dened (Brasier et al., 1994; Landing, 1994; Landing et al., 2007). The fossiliferous part of the Ajax Limestone as represented in section AJX-M broadly coincides with the currently unnamed Cambrian Series 2, Stages 3-4. These stages are approximately equivalent to the Atdabanian and Botoman of the Siberian stage nomenclature. The FAD of Abadiella huoi, the eponym of the oldest Australian trilobite Biozone, at 137 m above the base of the section marks the rst occurrence of a trilobite taxon in the Ajax Limestone, and most likely falls within Cambrian Series 2, Stage 3 (Babcock et al., 2005; Babcock and Peng, 2007; Babcock et al., 2009). Jell (in Bengtson et al., 1990) has described fragmentary specimens of Redlichiid indet. 3 possibly representing a species of Redlichia (cf. Paterson and Brock, 2007) from a stratigraphic level that coincides with the Pararaia tatei Biozone in the Ajax Limestone. Thus, the upper Ajax Limestone that equates to the P. tatei Biozone may be early Stage 4 in age, given that one of the proposed candidate horizons to mark the base of this stage is the FAD of Redlichia (Babcock et al., 2005; Babcock and Peng, 2007). Consequently, the entire Ajax Limestone at section AJX-M may potentially span from the late Terreneuvian Series, Stage 2 through to the unnamed Cambrian Series 2, Stage 4. Ongoing investigations to determine the FAD and LAD of SSFs, brachiopods and molluscs (see Topper et al., 2009), in conjunction with chemostratigraphic analyses, will help to constrain the position of Series and Stage boundaries in the lower Cambrian succession of South Australia.

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Fig. 1. A, locality map showing study areas, the Mount Scott and Elder ranges, Flinders Ranges, South Australia. B, simplied geological map of the Mount Scott Range showing the positions of section AJX-M through the Ajax Limestone. C, geological map of the Elder Range showing position of the ER9 section through the Hawker Group.

The AJX-M section contains an abundance of silicied, phosphatic or phosphatised macro- and microfossils, including species of trilobites, lingulate and rhynchonelliform brachiopods, molluscs, tommotiids, problematic small shelly fossils, spongiomorphs, plus eight species of bivalved arthropods documented herein. Archaeocyaths from the AJX-M section were described by Gravestock (1984; his section M) and Bengtson et al. (1990, g. 6) documented a variety of taxa including trilobites, molluscs, sponge spicules and SSFs from spot localities along this creek section. Recently, Skovsted et al. (2009) described the rst East Gondwanan occurrence of the stem group brachiopod Mickwitzia from this locality. The stratigraphic ranges of trilobites and

bivalved arthropods documented through the entire AJX-M stratigraphic section are plotted against the lithostratigraphic column in Fig. 2A. Two distinct trilobite biozones can be differentiated in the AJX-M section (see Jago et al., 2006 for the most recent review of the biostratigraphy of the lower Cambrian of South Australia). The rst appearance datum (FAD) of the index trilobite taxon Abadiella huoi at 137 m true thickness above the base of the section denes the base of the eponymous biozone. The LAD of A. huoi occurs at 197.67 m above the base of the section indicating that the entire A. huoi Biozone is 60.67 m thick in the section. The base of the succeeding Pararaia tatei trilobite Biozone, dened by the FAD of the eponymous species, is located at

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Fig. 2. A, stratigraphic ranges of trilobite and bivalved arthropod taxa through the Ajax Limestone in the AJX-M section, Mount Scott Range. B, stratigraphic ranges of stem group brachiopod, trilobite, and bivalved arthropod taxa through the upper Wirrapowie Limestone and Mernmerna Formation in the ER-9 section in the Elder Range.

199.63 m (true thickness) above the base of the section and 1.96 m above the last appearance datum (LAD) of Abadiella huoi. This leaves a small stratigraphic gap of 1.96 m, where it is difcult to ascertain the exact boundaries of the two trilobite biozones. The Pararaia tatei Biozone represents at least a 39.7 m stratigraphic interval in the upper half of the section, with the LAD of the eponymous species occurring at 239.33 m above the base of the section, which equates with a distinctive erosional surface undoubtedly reecting a short hiatus in sedimentation. No trilobite specimens were recovered from the red and grey dolomotised,

stromatolitic limestones representing the top 40 metres of section AJX-M, and the presence of the normally much older tommotiids Micrina etheridgei and Dailyatia ajax in these beds strongly suggests substantial reworking (or fault repetition) in this interval. The FAD of the oldest bradoriid taxon, Liangshanella circumbolina sp. nov., occurs at 117 m above the base of the section, c. 20 m below the FAD of Abadiella huoi. Despite detailed sampling throughout the AJX-M section, we acknowledge that pervasive dolomitisation has destroyed original fabrics, and so uncertainties remain about the presence (or not) of trilobites (and other fossils)

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in the lower dolomitised part of the Ajax Limestone succession. If the pre-trilobitic occurrence of Liangshanella circumbolina sp. nov. is accepted, then this suggests an early Series 2, Stage 3 age for this taxon, representing the oldest bivalved arthropod hitherto described from the lower Cambrian succession of South Australia. The rest of the bivalved arthropod assemblage (7 species in total see Fig. 2A) from AJX-M, correlates with the A. huoi Biozone and is still signicantly older that the bivalved arthropods recently documented from the Mernmerna Formation (P. bunyerooensis and P. janeae biozones) in the central Flinders Ranges by Skovsted et al. (2006) and Topper et al. (2007). Only two species, Mongolitubulus squamifer Missarzhevsky, 1977 and Zepaera sp. range into the succeeding Pararaia tatei Biozone (Fig. 2A). Two species that occur in the Ajax Limestone have been previously documented from South Australia: Dabashanella hemicyclica Huo, Shu and Fu in Huo et al., 1983, from the Pararaia bunyerooensis Biozone of the Mernmerna Formation (Skovsted et al., 2006) and Mongolitubulus squamifer from the P. janeae Biozone of the Mernmerna Formation (Topper et al., 2007). Unfortunately, since both are long-ranging taxa they have limited biostratigraphic value. Other long-ranging taxa with low biostratigraphic utility in the AJX-M and ER-9 sections include generic forms Euzepaera, Zepaera. and Mongolitubulus. Specimens of each genus from AJX-M and ER-9 are either too poorly preserved (e.g. Euzepaera sp. and Zepaera sp.) or display substantial morphological variation and fragmentation (e.g. Mongolitubulus sp.) that precludes assessment at species level. The bradoriid assemblage at section ER-9 is mainly restricted to the Wirrapowie Limestone, which equates to the basal 47 m (true thickness) of the section. The dearth of trilobite specimens throughout the ER-9 stratigraphic section does not allow precise placement of the zonal trilobite boundaries. The rst trilobite recovered in the section, Alanisia guillermoi Richter and Richter, 1940 occurs 106 m above the base of the section and is generally taken to represent the Pararaia tatei Biozone (Jell in Bengtson et al., 1990). Although no trilobites indicative of the Abadiella huoi Biozone were recovered from the basal 47 m of the ER-9 section, these beds do contain typical Atdabanian representatives including the tannuoliniid tommotiid Micrina etheridgei Tate, 1892 and the paterinid brachiopod Askepasma cf. toddense, which widely co-occur with Abadiella huoi elsewhere in the Arrowie Basin and provide strong evidence in support of an A. huoi Biozone equivalent age for the bradoriid assemblage in the Wirrapowie Limestone at section ER-9. The occurrence of Quadricona madonnae gen. et sp. nov. provides additional correlation with section AJX-M and supports a A. huoi equivalent biostratigraphic age for the assemblage. Two species, Alutella sp. and Spinospitella coronata Skovsted, Brock and Paterson, 2006 are also present in the overlying Mernmerna Formation at ER-9. Spinospitella coronata co-occurs with the P. tatei Biozone trilobite A. guillermoi and continues to range 25 m (true thickness) above the occurrence of A. guillermoi. Alutella sp. is conned to the uppermost part of the section which can be condently assigned a Pararaia janeae Biozone biostratigraphic age, based on the presence of the eponym along with Serrodiscus gravestocki Jell in Bengtson et al., 1990, Hebediscina yuqingensis Zhang in Zhang et al., 1980, Atops rupertensis Jell, Jago and Gehling, 1992, and a possible edelsteinaspidid. 3. Biogeographic implications The distribution of bradoriids, and to a much lesser extent phosphatocopids, has often been utilised in Cambrian palaeobiogeographical analyses (e.g. Melnikova et al., 1997; Siveter and Williams, 1997; Williams and Siveter, 1998; Hou et al., 2002). Shu (1981), in an early biogeographical review of bradoriids, identied

a western faunal realm, characterized by the genera Indiana and Bradoria and an eastern faunal realm, characterized by the so called alutids (e.g. Zepaera, Liangshanella and Kunmingella). The term alutids was used to describe bradoriids with apparent outer border structures on their shields (Shu and Chen, 1994). However, recent taxonomic revisions have demonstrated that the nominate taxa Aluta should be considered a nomen dubium and the term alutids be abandoned (Siveter and Williams, 1997; Williams et al., 2007). Palaeobiogeographic studies have tended to concentrate on comparisons of Cambrian bradoriid biogeographical patterns with that of trilobites (e.g. Melnikova et al., 1997; Hou et al., 2002; Williams et al., 2007; Zhang, 2007). The western faunal realm identied by Shu (1981) is generally associated with cool to cold water systems broadly mirroring the Olenellid trilobite Province whilst the eastern realm reects warm water, largely equatorial communities equivalent to the Redlichiid trilobite province (Melnikova et al., 1997; Hou et al., 2002; Williams et al., 2007). Bradoriid taxa characteristic of the Redlichiid trilobite province include Shangsiella, Comptaluta and Zepaera, whereas others, such as Indiana, Walcottella and Dielymella, epitomize the Olenellid trilobite province. The trilobite-based Acadobaltic Province sensu Sdzuy (1972), a region that includes Baltica, Avalonia, Armorica and Morocco also displays strong provinciality. This is supported by the presence of genera such as Beyrichona Matthew, 1886, Indiana and Wimanicharion Hinz-Schallreuter, 1993a, which are almost exclusively known from these areas in the late early and middle Cambrian (Streng et al., 2008). However, uncertainties have arisen regarding the degree of differentiation between the olenellid and redlichiid provinces (Zhang, 2003; Palmer, 2005). Zhang (2003) and Palmer (2005) have noted that some early Cambrian trilobites from both provinces display similarities that suggest that faunal provinciality was not as well developed in the early Cambrian. The use of the term western and eastern was abandoned by Shu and Chen (1994) who established the European realm and the 4A realm, based predominantly on temperature-latitude gradients. The European faunal realm consisted of Baltica, Avalonia, northern France and North Africa and represented a cool to cold water fauna and the 4A faunal realm consisted of Laurentia, Asia (North and South China), Siberia, Tarim, Kazakhstan, Mongolia, Australia and Antarctica, representing an equatorial, warm water fauna (Shu and Chen, 1994). Shu and Chen (1994) suggested a number of possible dispersal routes for bradoriids, including migration from South China to Australia during the late early Cambrian, based on the presence of Zepaera on both palaeocontinents. The late early Cambrian timing of the migration was based solely on the lack of bradoriid taxa from the early Cambrian of Australia. Zepaera has subsequently been discovered in the early Cambrian of Australia (Skovsted et al., 2006; Topper et al., 2007; herein) in conjunction with a diverse suite of bradoriid genera, suggesting that if a migration event did occur between South China and Australia, it must have transpired much earlier than previously recognised. Williams et al. (2007) broadly supported the idea of western and eastern faunal realms; however, stressed that the patterns of bradoriid biogeography was much more complicated than previous works indicated; they identied a number of distribution patterns and assemblages, reecting tropical and temperate climates as well as those groups that had a cosmopolitan distribution. Williams et al. (2007) provided a detailed account of the afnities and biogeography of bradoriids for the duration of the Cambrian until their eventual extinction in the mid-Ordovician. Williams et al. (2007) reviewed the bradoriid faunas from a number of palaeocontinental regions including Laurentia, Avalonia, Siberia, South China, central Asia and Gondwana commenting on the palaeogeographical positions of these regions pertaining to

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Fig. 3. A, reproduced cluster analysis of early Cambrian bradoriid genera conducted by Williams et al. (2007, g. 10A, page 224). B, pair-group cluster analysis for presence-absence data (Raup-Crick similarity index, correlation coefcient 0.6587) for 60 bradoriid genera from the early Cambrian. C, pair-group cluster analysis for presence-absence data (RaupCrick similarity index, correlation coefcient 0.6684) with Australia and Antarctica as amalgamated provinces of the palaeocontinental region of East Gondwana.

bradoriid distribution. Sedimentary successions from West Gondwana have yielded cambriids from the lower Cambrian of France (Vannier et al., 2005), hipponicharionids from Spain (Gozalo and Hinz-Schallreuter, 2002; Gozalo et al., 2004) and a number of taxa from the middle Cambrian of Morocco (Hinz-Schallreuter, 1993a). East Gondwana yields taxa from the lower Cambrian of Antarctica (Rode et al., 2003; Wrona, 2009) along with prolic faunas from the lower Cambrian through to the Furongian of Australia (e.g. pik, 1968; Jones and McKenzie, 1980). According to Williams et al. (2007, see table 2), at the generic level, Australia produces one of the most diverse bradoriid assemblages (23 genera documented in total) during the Cambrian to Early Ordovician interval. The large majority of East Gondwanan bradoriid taxa have been described from the Ordian and succeeding Stages across central and northern Australia (e.g. pik, 1968; Fleming, 1973; HinzSchallreuter, 1999). Previously, the Australian Ordian Stage was considered to be early Middle Cambrian (e.g. Shergold, 1996 and references therein), however evidence continues to amass suggesting that this stage should be regarded as latest Early Cambrian (see Laurie, 2006 and Paterson and Brock, 2007 for discussion). Consequently, all bradoriid genera described from the Ordian Stage of Australia are treated as early Cambrian in age and incorporated into the following analyses. The taxonomic diversity and biogeographic analysis of bivalved arthropods reported by Williams et al. (2007) only included two lower Cambrian (Cambrian Stage 2, Series 3-4) taxa from South Australia (Albrunnicola bengtsoni Hinz-Schallreuter, 1993a and Epactridion portax Bengtson in Bengtson et al., 1990). Recent work by Skovsted et al. (2006), Topper et al. (2007), Wrona (2009) and herein document another thirteen bivalved arthropod genera from the lower Cambrian of South Australia and Antarctica, greatly increasing the database of early Cambrian bivalved arthropod biodiversity from East Gondwana. As part of their original evaluation of bradoriid biogeography, Williams et al. (2007, g. 10) performed a pair-group cluster analysis using the statistical package PAST, based on presenceabsence data (Raup-Crick similarity index) of 32 bradoriid genera from the early Cambrian and 33 genera from the middle Cambrian. The cluster analysis presented by Williams et al. (2007, g. 10, reproduced here in Fig. 3A) indicated a wide distribution of bradoriid genera during the early Cambrian, with close faunal

associations between Avalonia, Laurentia, Baltica and West Gondwana (Morocco and Amorica). Bradoriid assemblages from South China clustered with those of Kazakhstan as part of an eastern hemisphere, warm water fauna, whilst Australian bradoriid data (based on two taxa) for the early Cambrian display only weak statistical similarity to South China, Kazakhstan and Siberia (Fig. 3A). We have performed a number of presence-absence cluster analyses (Fig. 3B, C) using PAST (Raup-Crick similarity index), employing the biogeographical distribution data provided by Williams et al. (2007, table 2 and appendix A), but also incorporating new taxonomic data published in recent years that

Fig. 4. A, pair-group cluster analysis for presence-absence data (Raup-Crick similarity index, correlation coefcient 0.6361) for 18 bradoriid genera from the early Cambrian, omitting bradoriid genera that have been documented from only a single palaeoncontinental region. B, pair-group cluster analysis for presence-absence data (Raup-Crick similarity index, correlation coefcient 0.6876) for 18 bradoriid genera from the early Cambrian, omitting bradoriid genera that have been documented from only a single palaeoncontinental region and with Australia and Antarctica as amalgamated provinces of the palaeocontinental region of East Gondwana.

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was not used in the Williams et al. (2007) analysis (e.g. Skovsted, 2006; Skovsted et al., 2006; Topper et al., 2007; Zhang, 2007; Dies lvarez et al., 2008; Hinz-Schallreuter et al., 2008; Wrona, 2009), in addition to data provided herein. Any bradoriid taxon that was questionably assigned at generic level have been omitted from the following analysis. Given the scope of our data, the biogeographic cluster analyses were limited to the early Cambrian; results discussed below. 3.1. Bradoriid assemblages from East Gondwana At the specic level the bradoriid assemblages from the early Cambrian of East Gondwana are largely endemic. Strong faunal provinciality among bradoriid species is not unusual at this time and has been recognised by numerous authors (e.g. Siveter and Williams, 1997; Williams et al., 2007), in particular faunas from South China frequently display high levels of endemism (Hou et al., 2002; Zhang, 2007). However, several genera from East Gondwana exhibit a wider distribution in the early Cambrian, including Liangshanella Huo, 1956 that has a near cosmopolitan distribution. As a result of particular palaeocontinental regions displaying high levels of endemism during the early Cambrian (e.g. East Gondwana and South China), two cluster analyses were undertaken (Figs. 3B-C, 4). The rst analysis (Fig. 3B-C) incorporates the presence-absence of 60 bradoriid genera from the early Cambrian. The second analysis (Fig. 4) only incorporates bradoriid genera that have been documented from two or more early Cambrian palaeocontinental regions, thereby removing all endemic taxa. We have also run the analysis with Australia and Antarctica as separate entities during the early Cambrian (Figs. 3B, 4A) and as amalgamated provinces of the palaeocontinental region of East Gondwana (Figs. 3C, 4B). We note that differences in taxonomic diversity between the analysed faunas do impose some limitations on the cluster analysis. Some regions remain inadequately explored in terms of bradoriid diversity (see Appendix), for example, Antarctica only has four described genera for the early Cambrian (e.g. Rode et al., 2003; Stigall, 2008; Wrona, 2009). Bradoriid diversity in other regions is much higher, such as Australia where 21 bradoriid genera have been documented from the early Cambrian (e.g. Skovsted et al., 2006; Topper et al., 2007; herein). The South China terrain hosts the most diverse bradoriid faunal assemblage during the early Cambrian with 32 genera documented. The rst pair-group cluster analysis (Fig. 3B) displays many similarities with the original cluster analysis (Fig. 3A) produced by Williams et al. (2007). Williams et al. (2007) recognised a wide distribution of congeners in the western hemisphere, between Avalonia, Baltica, Laurentia and West Gondwana, a view supported by data presented here. Both analyses also illustrate an association between South China and Kazakhstan. Close faunal connections exist between West Avalonia and West Gondwana (Morocco and Armorica) based on the presence of taxa including Hipponicharion Matthew, 1886 and Matthoria Siveter and Williams, 1997. The bradoriid fauna of Baltica displays close similarities with the fauna of West Avalonia, but also with Armorica and Morocco, supporting the previously recognised (Gozalo et al., 2004; Dies lvarez et al., 2008) Acadobaltic Province sensu Sdzuy (1972). The main discrepancy between the analyses is the close faunal association between Siberia, Laurentia and East Avalonia (Fig. 3C) based on the presence of taxa including Liangshanella Huo, 1956 and Matthoria, a grouping not recognised in the analysis presented by Williams et al. (2007). Australia, similar to data presented Williams et al. (2007), displays only weak statistical similarity to Siberia, Laurentia, East Avalonia and Antarctica. Incorporating the early Cambrian faunal assemblages of Australia and Antarctica to form an East Gondwana assemblage

(Fig. 3C) produces a dendrogram that is very similar to the results obtained by Williams et al. (2007). The only noticeable difference is the close association of Siberia with Laurentia and East Avalonia, discussed above and West Avalonia displaying a stronger faunal association with Armorica and Morocco (Fig. 3C). Recent publications (e.g. Skovsted et al., 2006; Topper et al., 2007; Zhang, 2007; Hinz-Schallreuter et al., 2008; Dies lvarez et al., 2008; Jones and Kruse, 2009; Wrona, 2009 and herein) have greatly increased the database of early Cambrian bivalved arthropod biodiversity, especially from East Gondwana, where nine new genera have been established. Despite this increase in faunal diversity, the resulting cluster analysis (Fig. 3C) produced only minor changes to the results of the original analysis conducted by Williams et al. (2007). A possible explanation for the limited variation in analyses is the high levels of endemism displayed in recently described bradoriid assemblages (Skovsted et al., 2006; Topper et al., 2007; Zhang, 2007; Wrona, 2009). Recent documentation of early Cambrian bradoriid assemblages from South Australia contain a high portion of endemic genera, including Spinospitella Skovsted, Brock and Paterson, 2006, Amphikeropsis Topper, Skovsted, Brock and Paterson, 2007, Onagrocharion Topper, Skovsted, Brock and Paterson, 2007 and Quadricona gen. nov. A diverse early Cambrian fauna from the Yu'anshan and Shuijingtuo formations of South China documented by Zhang (2007) included three new endemic genera, Retaluta, Spinaluta and Yucola. To highlight the strong endemism displayed by South Chinese faunal assemblages during the early Cambrian, of the total 32 genera documented, 19 are entirely restricted to the South China province. The addition of endemic taxa into a quantitative cluster analysis would not signicantly change the resulting biogeographic groupings of the palaeocontinental regions; merely change the similarity index between them. To gain a better understanding of the association of faunal assemblages during the early Cambrian we decided to conduct the cluster analysis again by omitting bradoriid genera that have only been documented from a single lower Cambrian palaeocontinental region, effectively removing all endemic taxa. The resulting cluster analysis (Fig. 4A) displays four distinct biogeographic groupings for the early Cambrian. Similar to previous results, close faunal associations exist between the palaeocontinental regions of the Acadobaltic Province (sensu Sdzuy, 1972), including West Avalonia, West Gondwana (Morocco and Armorica) and Baltica. Siberia has been biogeographically linked with Kazakhstan based solely on the occurrence of Alutella Kobayshi and Kato, 1951 on both palaeocontinents. Laurentian faunal assemblages have curiously been closely associated with Antarctica, based on the presence of Albrunnicola Martinsson, 1979 and Liangshanella Huo, 1956. However, this appears supercial, considering the strong evidence supporting the close early Cambrian faunal associations of Australia and Antarctica, based on trilobites (Palmer and Rowell, 1995; Brock et al., 2000; Paterson, 2005; Paterson and Jago, 2006), brachiopods (Holmer et al., 1996; Brock et al., 2000), archaeocyaths (Zhuravlev and Gravestock, 1994), molluscs (Wrona, 2003) and small shelly fossils (Evans and Rowell, 1990; Wrona, 2004). The apparent close relationship between Laurentia and Antarctica highlights one of the challenges with utilising this quantitative technique as differences in taxonomic diversity between faunal assemblages can distort the outcomes. Only two non-endemic bradoriid genera have been conrmed from Antarctica (Albrunnicola and Liangshanella), six genera have been documented from Laurentia, whereas twelve genera have been documented from the early Cambrian of Australia (see Appendix). Despite Albrunnicola and Liangshanella occurring on all three continental terranes, statistically, Antarctica will display a higher similarity to Laurentia because of the higher taxonomic diversity of Australian faunal assemblages.

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Fig. 5. Liangshanella circumbolina sp. nov. All specimens from the Ajax Limestone, Mount Scott Range. A-D, holotype, SAMP44789, sample AJX-M/209.2; A, lateral view of right valve, stereo pair, scale bar 500 m, B, interior view of right valve, scale bar 500 m, C, external surface ornament, scale bar 10 m, D, detail of external surface ornament, scale bar 5 m; E, detail of external surface ornament, scale bar 5 m, SAMP44790, sample AJX-M/209.2; F-I, SAMP44791, sample AJX-M/209.2; F, lateral view of left valve, scale bar 500 m, G, interior view of left valve, scale bar 500 m, H, detail of interior of left valve, scale bar 10 m, I, detail of sealed, tubular structures, protruding from the interior of the valve, scale bar 10 m.

Australia and South China are shown to have a very strong faunal association, with a similarity index of 0.94 (Fig. 4A), a marked contrast with the information presented by Williams et al. (2007) and by our previous analysis (Fig. 3A-B). Incorporating the early Cambrian faunal assemblages of Australia and Antarctica to form an East Gondwana assemblage (Fig. 4B) barely changes the original dendrogram. East Gondwana and South China again display a very denitive faunal association, with a slightly stronger similarity index than previously of 0.96 (Fig. 4B). This result is not surprising, as bradoriid taxa such as Zepaera Fleming, 1973, Haoia Shu, 1990, Hipponicharion , Liangshanella, Parahoulongdongella Shu, 1990, Albrunnicola (Albrunnicola bengtsoni =Beyrichona chinensis Zhang, 2007, p. 145) and Mongolitubulus (Mongolitubulus unispinoa = Spinella unialata) documented from the Mernmerna Formation (Skovsted et al., 2006; Topper et al., 2007) and the Ajax and Wirrapowie limestones herein (Figs. 5-7), have all been documented from the lower Cambrian (Eoredlichia-Wutingaspis Biozone) of China (e.g. Shu, 1990; Hou et al., 2002; Zhang, 2007). Based on similar shield morphologies, we consider Spinella unialata Zhang, 2007 (pl. 17, gs. 1-8) to be conspecic with Mongolitubulus unispinosa Topper, Skovsted, Brock and Paterson, 2007 (gs. 6A-J, 7A-C) and Beyrichona chinensis Shu, 1990 documented by Zhang (2007, p. 145, pl. 17, gs. 9-16) to be conspecic with Albrunnicola bengtsoni HinzSchallreuter, 1993a. The biogeographic ties between the two palaeoncontinental regions of South China and East Gondwana has been previously recognised based on trilobites (see Paterson and Brock, 2007). However, the results of the original cluster analysis (Fig. 3C) show a low similarity index between South China and East Gondwana (0.3). This evidently appears to be a direct result of differences in taxonomic diversity between the analysed faunas, due to high levels of endemism. Ten out of the total 12 non-endemic bradoriid genera documented from the early Cambrian of South Australia are

present in the early Cambrian of South China, further strengthening the biogeographic ties between the two palaeocontinental regions. The biogeographic groupings shown in Fig. 4B strongly reect the distribution of trilobites during the early Cambrian. South China and East Gondwana are characterised by trilobites that distinguish the Redlichiid trilobite Realm, whilst Laurentia, Avalonia and Baltica are characterised by trilobites distinguishing the Olenellid trilobite Realm. Intervening regions, such as Siberia, Morocco, Armorica and potentially Kazakhstan are characterised by trilobites of the transitional Bigotinid trilobite Realm (McKerrow et al., 1992; Pillola, 1993). Early Cambrian trilobites and other non-mineralized arthropods have recently been subjected to a number of quantitative biogeographic analyses (e.g. Lieberman, 1997, 2003a, b; Meert and Lieberman, 2004; 2008; Hendricks and Lieberman, 2007; Z. Zhang et al., 2008). In particular, the results of Lieberman (2003a, text-g. 2; 2003b, g. 2) and Meert and Lieberman (2004, g. 4) show similar major biogeographic groupings to our results, especially when compared with the cluster analysis that omitted endemic bradoriid genera and treated Australia and Antarctica as a single region (Fig. 4B). However, it is important to note that some discrepancies may be the result of the type of analysis conducted (phylogeographic vs. cluster, or a cladistic vs. phenetic approach, respectively) and the fact that the trilobite analyses were strongly biased towards the inclusion of olenelline taxa, which are absent in East Gondwana and South China. The analysis involving nonmineralized arthropods (Hendricks and Lieberman, 2007) produced results that show signicant differences in area relationships to those produced by bradoriid and trilobite data, although the grouping of Baltica and Africa and the close relationship of Siberia and Laurentia (Hendricks and Lieberman, 2007, g. 2) is supported by some of our results (Figs. 3B-C, 4A). Hendricks and Lieberman (2007) provided a number of explanations for these differences,

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Fig. 6. A-F, Zepaera jagoi sp. nov. All from the Ajax Limestone, Mount Scott Range, all scale bars 200 m, unless otherwise stated. A-C, holotype, SAMP44792, sample AJX-M/266; A, lateral view of right valve, stereo pair, B, detail of external surface ornament, scale bar 20 m, C, detail of spinose margin, scale bar 20 m; D, lateral view of left valve, SAMP44793, sample AJX-M/256; E-F, SAMP44794, sample AJX-M/266; E, internal view of left valve, F, internal view showing duplicature, scale bar 20 m; G-M, Quadricona madonnae gen. et sp. nov. G-J, holotype from the Mernmerna Formation, Elder Range, SAMP44795, sample ER-9/67.3; G, lateral view of left valve, H, oblique lateral view of left valve, I, oblique lateral view from posterior, J, detail of external surface ornament, scale bar 10 m; K, lateral view of right valve, scale bar 500 m, SAMP44796, sample AJX-M/268.7; L-M, SAMP44797, sample AJX-M/271.3; L, dorsal view of shield, scale bar 500 m, M, oblique view of shield, scale bar 500 m.

but it is also worth noting that their analysis included taxa ranging from the early Cambrian to latest Ordovician, potentially introducing noise to the biogeographic signal. Recent palaeogeographic reconstructions of the early Cambrian indicate that Australia and the South China block occupied the tropical carbonate development zone (30 5 north and south latitudes) (Brock et al., 2000; Meert and Lieberman, 2004; Williams et al., 2007). During the early Cambrian in South China, the dominant groups of bivalved arthropods are the kunmingellids and the comptalutids. Kunmingellids frequently cover bedding surfaces (e.g. in the Heilinpu Formation), with the type species, Kunmingella

douvillei (Mansuy, 1912) accounting for over 80% of recovered individuals in the Chengjiang biota (Hou and Bergstrom, 1991; Hou et al., 2002). In contrast, East Gondwanan bradoriid assemblages contain an equal proportion of many different groups, including the svealutids (Liangshanella), hipponicharionids (Hipponicharion and Albrunnicola), beyrichonids (Parahoulongdongella), comptalutids (Zepaera, Alutella), monasteriids (Epactridion) and many other genera that are yet to be condently assigned at family level (e.g. Onagrocharion, Amphikeropsis and Spinospitella). Williams et al. (2007) identied a number of faunal assemblages that reected tropical and temperate climates during the

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Fig. 7. A-G, Parahoulongdongella bashanensis Shu, 1990. From the Mernmerna Formation, Elder Range and the Ajax Limestone, Mount Scott Range, all scale bars 200 m, unless otherwise stated. A, lateral view of right valve, SAMP44798, sample ER-9/16.5; B-D, SAMP44799, sample ER-9/16.5; B, lateral view of right valve, C, oblique lateral view of right valve, D, detail of external surface ornament, scale bar 10 m, E-G, SAMP44800, sample ER-9/53.5; E, lateral view of right valve, F, oblique lateral view of right valve, G, detail of external surface ornament, scale bar 50 m; H, Zepaera sp., lateral view of right valve, SAMP44801, sample AJX-M/410; I, Mongolitubulus sp., lateral view of spine, SAMP44802, sample ER-9/10.8; J-K, Mongolitubulus squamifer Missarzhevsky, 1977. J-K, SAMP 44803, sample AJX-M/415; J, lateral view of spine, scale bar 60 m, K, detail of external surface ornament, scale bar 20 m; L, Comptalutid gen et. sp. indet., lateral view of right valve, scale bar 100 m, SAMP44804, sample AJX-M/266; M, Euzepaera sp., lateral view of right valve, scale bar 500 m, SAMP44805, sample ER-9/53.5; N, Spinospitella coronata Skovsted, Brock and Paterson, 2006, lateral view of valve fragment, SAMP44806, sample ER-9/0; O, Dabashanella hemicyclica Huo and Shu in Huo et al., 1983, lateral view of left valve, SAMP44807, sample AJX-M/274.4; P, Indet. phosphatocopid, lateral view of left valve, scale bar 100 m, SAMP44808, sample AJX-M/256.5.

Cambrian. The East Gondwana bradoriid assemblage in the early Cambrian broadly conforms to the climatic distribution patterns identied by Williams et al. (2007). Comptalutids, prevalent in South Chinese and Australian assemblages are interpreted as warm water tropical and sub-tropical bradoriids that cover a latitudinal range from about 30 North to 15 South (Williams et al., 2007). Species of Liangshanella have been documented from early Cambrian faunal assemblages in East Gondwana, South China, Avalonia, Laurentia and Baltica and thus seems capable of surviving in both low and mid-latitudes (Williams et al., 2007). The beyrichonids and hipponicharionids have been recognised as mid- to high latitude bradoriids (further north and south than 30), based on their preference for Avalonia and Baltica during the Cambrian (Williams et al., 2007). Regardless, some hipponicharionids, like Neokunmingella Zhang, 1974, Meishucunella Jiang, 1982 and occur

amongst the early Cambrian faunal assemblages of South China (Hou et al., 2002). Hipponicharion, a genus previously thought to be restricted to the Acadobaltic Province (sensu Sdzuy, 1972; Gozalo and Hinz-Schallreuter, 2002; Gozalo et al., 2004) has since been documented from South China and South Australia (Topper et al., 2007; Zhang, 2007), extending the biogeographic range of the genus into low latitude, tropical regions. The beyrichonids, Beyrichona Matthew, 1886 and Parahoulongdongella have also been described from the low latitude faunas of southern Kazakhstan (Melnikova et al., 1997), South Australia and South China (Topper et al., 2007; Zhang, 2007). While both groups may be more abundant in Avalonian and Baltic faunas, holistically, representatives of the Hipponicharionidae and Beyrichonidae, were evidently able to disperse between the warm climates of South Australia and South China and the cooler climates of Avalonia and Baltica.

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4. Systematic palaeontology All specimens described and illustrated herein are housed in the palaeontological collection of the South Australian Museum (acronym: SAMP). Stratigraphic ranges for all taxa are provided in terms of true thickness above the base of the AJX-M and ER-9 sections. The terminology used to describe the bivalved arthropods largely follows that employed to describe Ordovician and later ostracods (as adopted in Siveter and Williams, 1997; Williams and Siveter, 1998; Hou et al., 2002). We follow Siveter et al. (2003, p. 13) in using the more neutral term head-shield, or shield instead of the ostracod term carapace (see also Maas et al., 2003). The majority of additional bivalved arthropod taxa in the Ajax and Wirrapowie assemblages (see Figs. 2, 7) have been described previously from the Mernmerna Formation; refer to Skovsted et al. (2006) and Topper et al. (2007) for descriptions and discussions. Phylum Arthropoda Siebold and Stannius, 1845 Order Bradoriida Raymond, 1935 Family Svealutidae pik, 1968 Liangshanella Huo, 1956 Type species. Liangshanella liangshanensis Huo, 1956 Liangshanella circumbolina sp. nov. (Fig. 5) Type material. Holotype: SAMP44789 (Fig. 5A-D) from sample number AJX-M/209.2, 117 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range. Paratypes: SAMP44790 (Fig. 5E) and SAMP44791 (Fig. 5F-I) from sample number AJX-M/ 209.2, 117 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range. Etymology. Dervied from the Latin prex circum, circle and bolus, lump. In reference to the distinctive circlet of raised papillate structures on the outer valve surface. Diagnosis. Moderately to strongly inated postplete valves with a straight hinge line. Latero-admarginal ridge present, entire between cardinal corners, anterodorsal or posterodorsal lobation or sulci absent. Outer surface with numerous, shallow depressions each hosting a submicron sized, centrally located circular perforation girt by a circlet of raised, closely packed papillate structures; individual depressions are sealed on the internal valve surface by raised, cylindrical pillars. Description. Moderately to strongly inated valves, with a straight hinge line. Postplete valves exhibiting a distinct lateroadmarginal ridge, entire between cardinal corners and separated from the lateral surface by a continuous furrow. Maximum dimensions of shield are 1.48 mm in length and 1.27 mm in height. Lateral surface lacks distinct anterodorsal or posterodorsal nodes or sulci. Valve surface consists of numerous, small depressions, varying from 7 12 m in diameter that are evenly distributed over the entire shield (Fig. 5C). Each individual depression hosts a submicron sized, centrally located circular perforation (Fig. 5D-E) surrounded by a circlet of raised, closely packed papillate structures; raised circlet structure has an approximate diameter of 4 m (Fig. 5D-E). Internally, each depression is sealed by raised cylindrical pillars that protrude from the valve some 10 - 13 m (Fig. 5H-I); neighbouring pillars can be fused or isolated (Fig. 5H, I), depending on space between individual depressions. Remarks. Postplete bradoriid valves from the Cambrian that display little or no lobation are often referred to the genus Liangshanella Huo, 1956. Originally described for svealutids that exhibit no lobation (Huo, 1956), the generic diagnosis has since been modied slightly to incorporate an anterior lobe that may range from well developed to virtually lacking (Siveter and Williams, 1997; Hou et al., 2002). The simplicity of shield morphology makes it difcult to distinguish Liangshanella from other bradoriid genera that also possess a non-lobate shield, such as Indiana Matthew, 1902, Hanchungella Huo, 1956, Ovaluta Zhang, 1987 and Indota pik, 1968. Such difculties are exempli-

ed by Hou et al. (2002) who synonymised both Hanchungella and Ovaluta with Liangshanella citing similar shield morphologies. Hou et al. (2002) considered that the alleged differences in anterior cardinal angle traditionally used to discriminate these genera was the result of deformational processes, and thus of no taxonomic signicance. Zhang (2007) disputed this synonymy and suggested that a lack of ontogenetic information made such claims premature. Zhang (2007) considered all generic names valid, and noted that the presence of an anterior lobe can be used to distinguish Hanchungella from a lobeless Liangshanella. Zhang (2007) was also unconvinced that Ovaluta represented a juvenile instar of Liangshanella, instead noting the strong similarities in shield size and features of Ovaluta with Bullaluta Copeland, 1986, an upper Cambrian (Furongian) bradoriid of western Newfoundland, and suggested that the former may be a junior synonym. Liangshanella circumbolina sp. nov. displays the postplete shield, lateroadmarginal rim and lack of lobation typical of the genus and is similar in general shield outline to L. liangshanensis Huo, 1956, L. rotundata Huo, 1956 and L. yunnanensis Zhang, 1974 described from South China and L. burgessensis Siveter and Williams, 1997 from the Burgess Shale in Canada. However, all four of these species have a weakly to moderately developed anterodorsal node, a feature absent in L. circumbolina sp. nov. Liangshanella baensis described by Zhang (2007, pl. 15, gs. 1-5) from the Shuijingtuo Formation of southern China differs by possessing a broad, shallow sulcus underneath the anterodorsal margin and a narrow marginal rim. Liangshanella sayutinae (Melnikova, 1988) from the TransBaikal area in the Russian Far-East and Greenland (Melnikova et al., 1997; Skovsted, 2006) lack an anterodorsal node and has a valve prole similar to L. circumbolina sp. nov., but differ by the surface ornament of ne cancelate ridges. Liangshanella circumbolina sp. nov. possesses a very distinctive exterior valve ornament consisting of evenly distributed depressions each containing a raised circlet of closely joined papillae surrounding a single, submicron sized central perforation (Fig. 5C-E). This highly distinctive exterior valve ornament has not been previously documented from any Liangshanella species, nor indeed from any other bradoriid species. However, it should be noted that the majority of Liangshanella species have been described based on crack-out macrofossil specimens from ne-grained siliciclastic rocks (e.g. Siveter and Williams, 1997; Hou et al., 2002), a mode of preservation not ideal for preserving ne, microstructural detail. Minute surface perforations have been previously interpreted as potential openings for sensory setae (Skovsted et al., 2006; Topper et al., 2007), analogous to those found in modern ostracods. The function of the raised circlet of papillae surrounding the central perforation is unknown, but may have provided some protection to the base of sensory hairs that may have developed inside the sealed cylindrical pillars on the inner surface of the valve. Similar structures are found in the recent ostracod, Elofsonia baltica (Hirschmann, 1909), which possesses sieve-pores that have a central sensory hair, surrounded by a ring of mound-like papillae (see Whittaker, 1973, pl. 1:37:200, g. 2). However, the exact functional signicance of the mound-like papillae in this taxon also remains mysterious. The FAD of Liangshanella circimbolina sp. nov. occurs some 20 m below the FAD of the zonal trilobite Abadiella huoi in the Ajax Limestone. The pre-trilobitic occurrence of this species means that, potentially, Liangshanella circumbolina sp. nov. represents the oldest bradoriid taxon hitherto documented from the early Cambrian of East Gondwana. Previously, the oldest bradoriids documented from East Gondwana were possible svealutids from the lower Cambrian (Atdabanian equivalent) Parara Limestone at Curramulka Quarry on Yorke Peninsula described by Chapman (1918) as ostracods. Jell in Bengtson et al. (1990, g. 7) indicated that the lower levels of the Parara Limestone at Curramulka

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contained A. huoi, and thus can be correlated with the A. huoi Biozone. Jones and Laurie (2006) suggest that the drawings by Chapman (1918, pl. 9, gs 1, 2, 5) bear some resemblance to the Svealutidae, however without a more accurate representation of the specimens, comparison with L. circumbolina sp. nov. is not possible. Distribution. Ajax Limestone, lower Cambrian (Series 2, Stage 3), Flinders Ranges, Arrowie Basin, South Australia. Stratigraphic range. All specimens recovered from a single stratigraphic level, 117 m above the base of the AJX-M section. Family Comptalutidae pik, 1968 Zepaera Jones and McKenzie, 1980 Type species. Zepaera rete Jones and McKenzie, 1980 Zepaera jagoi sp. nov (Fig. 6A-F) Type material. Holotype: SAMP44792 (Fig. 6A-C) from sample number AJX-M/266, 148.75 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range. Paratypes: SAMP44793 (Fig. 6D) from sample number AJX-M/255, 143 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range and SAMP44794 (Fig. 6E-F), from sample number AJX-M/266, 148.75 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range. Etymology. Named for Dr. Jim Jago, in recognition for his contributions to South Australian Cambrian geology, palaeontology and biostratigraphy. Diagnosis. Valves postplete, with a marked retral swing and a well developed, straight hinge line. Lateroadmarginal ridge present, entire between cardinal corners with valves showing weak anterior sulcation. Valve exterior with widely spaced, circular punctae; surface ornament consists of irregularly concentric wrinkled texture. Valve margin well dened, ornamented by a series of regularly spaced, short, tapering spines. Description. Valves small (maximum height 810 m, maximum length 960 m), postplete, with a well developed, straight hinge line. Valves have a thin (approximately 7.5 m in width) lateroadmarginal ridge, entire between cardinal corners. Lateroadmarginal ridge separated from the lateral surface by a continuous, but weakly dened shallow depression (Fig. 6C). Valve margin well dened, ornamented by a series of regularly spaced, short, tapering spines (Fig. 6C). Spines are approximately 10 m in length and vary between 13-19 m in width and are continuous along the valve margin, between cardinal corners (Fig. 6A). Lateral surface moderately to strongly inated, without distinct anterodorsal or posterodorsal nodes. A weak, V-shaped centrodorsal depression is present in some specimens (Fig. 6D). Surface is infrequently punctate (Fig. 6C) ornamented with an irregularly concentric wrinkled texture (Fig. 6B). Some specimens exhibit low, at circular welt-like structures, varying in diameter from 26 m to 63 m (Fig. 6B). Interior displays a well dened duplicature, entire between the cardinal corners (Fig. 6F). Remarks. Comptalutids, like svealutids, frequently exhibit morphologically simple shields that result in taxonomic confusion between some early Cambrian bradoriids such as Houlongdongella Lee, 1975, Flemingopsis Jones and McKenzie, 1980, Zepaera Jones and McKenzie, 1980 and Quetopsis Hinz-Schallreuter, 1999. The exact taxonomic status and relationship of these taxa has been the focus of frequent discussion, but remains controversial (Shu, 1990; Hinz-Schallreuter, 1999; Skovsted et al., 2006; Zhang, 2007). Due to the simplicity of the comptalutid shield, soft-part morphological information will undoubtedly play an important role in the taxonomic resolution of many comptalutid taxa. New softpart preservation has assisted in the taxonomic classication of the comptaluid Kunyangella Huo, 1965 documented from the Chengjiang Lagersttte of China (Hou et al., 2010). A specimen of Kunyangella cheni Hou, Williams, Siveter, Siveter, Aldridge and Sansom, 2010 interpreted as a penultimate instar displays

remnants of three head and two trunk appendages. Based on the ontogentic patterns of segmentation in Crustacea, Hou et al. (2010) reconstructed the adult as having four head appendages. However, to date only two bradoriid species (Kumingella douvelli and Kunyangella cheni) have been described with their soft-part anatomy preserved (Hou et al., 1996, 2010; Shu et al., 1999). In the absence of soft-part preservation, Zhang (2007) suggested detailed ontogenetic assessment may provide a solution for elucidating the taxonomic status of comptalutids. Whilst the various growth stages of some bradoriids, such as Zepaera rete Jones and McKenzie, 1980, Flemingopsis dua (Jones and McKenzie, 1980) and Houlongdongella xichuanensis Zhang, 1987, have been well illustrated, the ontogeny of the vast majority of bradoriids remain obscure. This is primarily due to a lack of available material, since an accurate ontogenetic framework requires a considerable catalogue of specimens. Zhang (1987), for example, investigated 131 specimens of Houlongdongella xichuanensis and Hinz-Schallreuter (1999) examined 240 specimens of Zepaera rete and 221 specimens of Flemingopsis dua for ontogenetic analysis. The difculties associated with charting bradoriid ontogeny using external features of the shield was recently highlighted by Hou et al. (2010) who described pre-adult and adult specimens of Kumingella douvelli and Kunyangella cheni that displayed changes in soft-part anatomy between growth stages, but little or no morphologic change was manifested in shield morphology. The genus Zepaera was originally reserved for comptatulid bradoriids with distinctive lobes forming a characteristic omega-shaped ridge on the anterior part of the valve. However, the omega-shaped ridge was found to be only distinct in juvenile stages, gradually becoming effaced, until it fades completely in adult stages (Jones and McKenzie, 1980; Hinz-Schallreuter, 1999). This ontogenetic variation has also been observed in other comptalutids (e.g. Flemingopsis dua Hinz-Schallreuter, 1999). Zepaera jagoi sp. nov. lacks these characteristic ridges and with the large majority of specimens approaching a maximum length of approximately 900 m, all specimens are herein considered to represent fully mature adults. Zepaera jagoi sp. nov. displays the typical postplete shield outline and marked retral swing characteristic of adult Zepaera species. Zepaera jagoi sp. nov. displays a weak V-shaped anterodorsal sulcus but lacks any obvious anterior lobation that characterises the type species, Z. rete. The irregularly concentric, wrinkled ornament of Z. jagoi sp. nov. also distinguished it from the well developed reticulate ornament in the type species. There are a number of Cambrian bradoriid genera with short spinelike protuberances from their shield; however the regularly spaced, short spines that adorn the valve margin of Z. jagoi sp. nov. (Fig. 6A, C) have not been previously described in any known bradoriid taxon. Distribution. Ajax Limestone, lower Cambrian (Series 2, Stage 3), Abadiella huoi Biozone, Flinders Ranges, Arrowie Basin, South Australia. Stratigraphic range. A total of 5.6 m between 143.15-148.75 m above the base of the section AJX-M. Family Hipponicharionidae Sylvester Bradley, 1961 Remarks. A subtriangular shield in conjunction with distinct, anterodorsal and posterodorsal lobes is indicative of two families within the Bradoriida, the Hipponicharionidae and the Beyrichonidae. The beyrichonids can be discriminated from the hipponicharionids by having a less inated shield, with subdued anterior and posterior lobes that are frequently less ventrally extended (HinzSchallreuter, 1993a, c; Siveter and Williams, 1997). The close morphological similarity between the two families was recognised by pik (1968), who regarded the Hipponicharionidae as a subjective synonym of the Beyrichonidae based on the comparable morphology of Hipponicharion Matthew, 1886 and Beyrichona Matthew, 1886, the type genera of their respective families. This view has not been widely accepted. Hinz-Schallreuter (1993a),

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Siveter and Williams (1997) and Williams et al. (2007) all emphasized the different lobation between the families, in particular the anterior and posterior lobes that ventrally extend further in the Hipponicharionidae than in the Beyrichonidae. In addition, Zhang (2007) noted that the posterior node or linear lobe is distinct in the Hipponicharionidae, whereas it often forms as sulcation in the Beyrichonidae, a morphological feature that characterises the type species, Beyrichona papilio Matthew, 1886 (see Siveter and Williams, 1997, pl.4, gs. 9-10). Based on the differences in shield ination and lobation, both families are recognised as valid, separate taxonomic groupings. Quadricona madonnae gen. et. sp. nov. displays the amplete shield, subtriangular shield outline and distinct lateroadmarginal ridge reminiscent of the Hipponicharionidae and Beyrichonidae, however the lobation is strikingly different. Quadricona madonnae gen. et. sp. nov. displays short, pointed anterior and posterior nodes (Fig. 6E-F, L), unlike the rounded nodes, or ridge-like lobes frequently encountered in the Beyrichonidae and the Hipponicharionidae. The anterior and posterior nodes are prominent in all specimens with no examples of posterior sulcation observed, a feature frequently displayed by the beyrichonids. The moderately inated shield and prominent lobation displayed by Quadricona madonnae gen. et. sp. nov. supports designation to the Hipponicharionidae. Quadricona gen. nov. Etymology. Derived from the Latin prex quattuor, four and conus, cone. In reference to the four, short nodes on the complete shield of the bradoriid. Diagnosis. Valves small, subamplete, with a subtriangular shield outline and well developed, straight hinge line. Narrow lateroadmarginal ridge, entire between cardinal corners. Anterodorsal and posterodorsal nodes are discrete, short, nipple-like protuberances. Surface with many small, closely spaced circular pores, each in a well dened u-shaped depression; some pores with raised, circular margin. Surface sculpture partially wrinkled, more prominent towards the valve margin. Remarks. Discrimination of genera within the Hipponicharionidae is predominantly related to shield shape, size and ventral extent of the anterior and posterior lobes. For example, Wimanicharion HinzSchallreuter, 1993a, Andresia Hinz-Schallreuter, 1993a and Neokunmingella Zhang, 1974, typically exhibit conuent anterior and posterior lobes. Albrunnicola Martinsson, 1979, one of the most common bradoriids in early Cambrian assemblages from South Australia, is characterised by a weakly developed posterior lobe, with both lobes restricted to the dorsal half of the valve. Quadricona gen. nov. is similar in shield outline to Albrunnicola and also has a very similar surface sculpture (see Topper et al., 2007, g. 10F-G). However, the degree and development in shield lobation between the two genera is strikingly different. The discrete short, nipple-like projections displayed by Quadricona gen. nov. is a unique morphological characteristic that easily distinguishes it from other genera within the Hipponicharionidae. Quadricona madonnae gen. et sp. nov. (Fig. 6G-M) Type material. Holotype: SAMP44795 (Fig. 6G-J) from sample number ER-9/67.3, 38.6 m true thickness above the base of the section, Wirrapowie Limestone, Elder Range. Paratypes: SAMP44796 (Fig. 6K) from sample number AJX-M/268.7, 150.2 m true thickness above the base of the section and SAMP 44797 (Fig. 6 L-M) from sample number AJX-M/271.3, 151.7 m true thickness above the base of the section, Ajax Limestone, Mount Scott Range. Etymology. Named for the American entertainer Madonna; in reference to the nodes on each valve resembling her conical brassiere made famous during the 1980s and 1990s, particularly her Blond Ambition tour in 1990. Diagnosis. As for genus, by monotypy. Description. Valves subamplete with a subtriangular outline and a well developed, straight hinge line. The valves are small in size,

measuring on average 1.19 cm in length and 0.922 cm in height, with a maximum length of 1.3 cm and height of 1.05 cm. Valves are moderately convex with generally rounded posterior and anterior margins. The anterior and posterior borders extend slightly beyond the hinge line (Fig. 6G). A distinct, narrow, lateroadmarginal ridge is entire between cardinal corners. Anterodorsal and posterodorsal nodes are discrete, short, sharp, nipple-like protuberances (Fig. 6E-F, L). Posterodorsal node is positioned slightly more dorsally than the anterodorsal node (Fig. 6G). A slight central, dorsal swelling close to the dorsal valve margin, slightly anterior of the midline forms a weak and indistinct central lobe (Fig. 6G). Exterior surface with numerous small u-to v-shaped indentations, hosting a single, minute perforation with a distinctive, but simple, raised rim approximately 3 m in diameter (Fig. 6J). Surface sculpture partially wrinkled, especially towards the margins of the valves (Fig. 6I). Interior surface smooth. Remarks. Quadricona madonnae gen. et sp. nov. is based on eight specimens recovered from the Ajax Limestone in the Mount Scott Range and the upper Wirrapowie Limestone in the Elder Range. Quadricona madonnae gen. et sp. nov. is the only taxon unequivocally common to both localities and the short stratigraphic ranges in both sections (Fig. 2) may have potential for future regional correlation. The shield outline and surface ornament of Q. madonnae is closely comparable to a number of Cambrian bradoriids such as the hipponicharionid Albrunnicola bengtsoni Hinz-Schallreuter, 1993a and the beyrichonid Parahoulongdongella bashanensis Shu, 1990 (see Figs. 6J, 7D). This suggests that similar types of surface ornamentation may appear in many different bradoriid lineages, restricting the value of this feature in higher level bradoriid taxonomy. Distribution. Ajax and Wirrapowie Limestones, Early Cambrian (Series 2, Stage 3), Abadiella huoi Biozone, Flinders Ranges, Arrowie Basin, South Australia. Stratigraphic range. A total interval range of 1.45 m from 150.26151.71 m above the base in section AJX-M (Ajax Limestone) and a total of 4.71 m from 38.6-43.31 m above the base in section ER-9 (Wirrapowie Limestone). Acknowledgements Financial support towards eldwork and laboratory costs come from a Macquarie University Research Development Grant to GAB and CBS. Funding support for CBS was also provided by a postdoctoral grant from the Swedish Research Council (VR). We extend warm thanks to Mr Graham Ragless, owner of Beltana Station and Mr Glen Gabe, Manager at Mt. Little Station for access to the eld areas. We are indebted to B. Jonak, P. Cockle, L.E. Holmer, J.B. Jago, R. Smart, M. Smith, T. Bradley, R. Callow and M. Fuller for their assistance during the eld seasons when this material was collected. We are indebted to Dean Oliver (Dean Oliver Graphics) for drafting Figs. 1-2 with characteristic skill and speed. We also acknowledge Elsevier for permission to reproduce the cluster analysis presented by Williams et al. (2007, g. 10, reproduced here in Fig. 3A). The manuscript beneted from the constructive reviews of two anonymous reviewers. Appendix A Global distribution of bradoriid genera of the major palaeocontinents during the early Cambrian. Presence of bradoriid genera indicated by shaded box. Taxa that have only been documented from a single palaeocontinental region and subsequently removed from the second cluster analysis (Fig. 4) are indicated with an asterix. Principal sources of revised taxonomy and biogeographic information from Morocco (Hinz-Schallreuter, 1993a), Australia (Opik, 1968; Fleming, 1973; Jones and McKenzie, 1980; Hinz-Schallreuter,

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1999; Skovsted et al. 2006; Topper et al., 2007), Antarctica (Rode et al., 2003; Wrona, 2009), Armorica (Gozalo et al., 2004; Vannier et al., 2005), Siberia (Melnikova et al., 1997), Laurentia (Siveter et al., 1996; Siveter and Williams, 1997; Skovsted 2006), Baltica (Ulrich

and Bassler, 1931; Dies lvarez et al., 2008), West Avalonia (Siveter and Williams, 1997), East Avalonia (Williams and Siveter, 1998), South China (Hou et al. 2002; Zhang, 2007) and Kazakhstan (Melnikova et al. 1997).

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