Gondwana Research 19 (2011) 310–326

Contents lists available at ScienceDirect

Gondwana Research
j o u r n a l h o m e p a g e : w w w. e l s e v 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
a, b,c a d
Timothy P. Topper , Christian B. Skovsted , Glenn A. Brock , John R. Paterson
a Department of Biological Sciences, Macquarie University, NSW 2109, Australia
b Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala, Sweden
c 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

article info abstract

Article history: The oldest bradoriid fauna from Australia, occurring in the lower Cambrian Ajax and Wirrapowie limestones of the Flinders
Received 15 February 2010 Ranges, South Australia consists of eleven taxa, including one new genus and species, Quadricona madonnae gen. et sp. nov.
Received in revised form 30 April 2010 and two new species, Liangshanella circumbolina sp. nov. and Zepaera jagoi sp. nov. In the Ajax Limestone, Liangshanella
Accepted 23 May 2010 Available online 4
circumbolina sp. nov. occurs c. 20 m below the FAD of the zonal trilobite Abadiella huoi. This pre-trilobitic occurrence
June 2010
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),
Keywords:
Cambrian Pararaia tatei, P. bunyerooensis and P. janeae (all Botoman) trilobite biozones in South Australia indicates great potential for
Arthropoda future regional biostratigraphic correlation. Quantitative biogeographic analysis including new taxonomic data from the lower
Bradoriida Cambrian of South Australia, highlights the strong endemism displayed by early Cambrian bradoriid communities and
Phosphatocopida strengthens the close faunal affinities with South China and Antarctica.
South Australia

© 2010 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

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 Lagerstätten, including the Buen Formation (Siveter et al.,
1996), Qiongzhusi Formation (Hou and Bergström, 1991; Hou et al., 2002),
Burgess Shale Formation (Conway Morris, 1986; Siveter and Williams, 1997)
and the Alum Shale Formation (e.g. Müller, 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 five 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 (Öpik, 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
T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 311
et al., 2006; Topper et al., 2007) adding substantially to the diversity of
bivalved arthropods previously documented from the Parara Limestone in the
Bivalved
Stansbury arthropods
Basin co-occur
(Chapman, 1918;with the first
Bengtson trilobites
et al., 1990).on several continents
(Williams et al., 2007), but appear slightly earlier than the first 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 estab-lished the presence of species belonging
to the Hipponicharionidae (Albrunnicola bengtsoni Hinz-Schallreuter, 1993a)
and Monaster-iidae (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 first
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 Gond-wana, 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 provid-ed, 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 first stratigraphic section (AJX-
M) 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 30°35'49" S, 138°19'59.3"
E [WGS84] and is equivalent to section M of Gravestock (1984, fig. 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
silicified) 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 Wirrapo-wie
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 31°40'2.6"S;
138°26'11.5"E [WGS84]. The upper 47 m of massive, algal dominated,
mottled and ribboned carbo-nates 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 undefined 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 ratified (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 defined (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 first 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.
312 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326

Fig. 1. A, locality map showing study areas, the Mount Scott and Elder ranges, Flinders Ranges, South Australia. B, simplified 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 silicified, phos-phatic or
phosphatised macro- and microfossils, including species of trilobites,
lingulate and rhynchonelliform brachiopods, molluscs, tommotiids,
problematic small shelly fossils, spongio-morphs, 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, fig. 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 first 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 first appearance datum (FAD) of the index
trilobite taxon Abadiella huoi at 137 m true thickness above the base of the
section defines 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, defined by the FAD of the
eponymous species, is located at
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 313

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 difficult 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
reflecting 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)
314 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
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 significantly older that the bivalved arthropods recently
documen-ted 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 biostrati-graphic
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 pre-cludes 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 first 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 confined to the uppermost part of the section
which can be confidently assigned a Pararaia janeae Biozone biostrati-graphic
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 palaeobio-geographical
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, identified
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 identified by Shu (1981) is generally associated
with cool to cold water systems broadly mirroring the Olenellid trilobite
Province whilst the ‘eastern’ realm reflects 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 identified a number of distribution patterns and
assemblages, reflecting tropical and temperate climates as well as those
groups that had a cosmopolitan distribution.
Williams et al. (2007) provided a detailed account of the affinities and
biogeography of bradoriids for the duration of the Cambrian until their
eventual extinction in the mid-Ordovician. Williams et al. (2007) reviewed
number of palaeocontinental regions
the bradoriid faunas from a
including Laurentia, Avalonia, Siberia, South China, central
Asia and Gondwana commenting on the palaeogeographical
positions of these regions pertaining to
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 315

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

bradoriid distribution. Sedimentary successions from West Gond-wana 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 prolific faunas from
the lower Cambrian through to the Furongian of Australia (e.g. Öpik, 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.
associations between Avalonia, Laurentia, Baltica and West Gond-wana
(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

The large majority of East Gondwanan bradoriid taxa have been described
from the Ordian and succeeding Stages across central and northern Australia
(e.g. Öpik, 1968; Fleming, 1973; Hinz-Schallreuter, 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
suggest-ing 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 (Albrun-nicola
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, fig. 10) performed a pair-group cluster analysis using the statistical
package ‘PAST’, based on presence-absence 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, fig.
10, reproduced here in Fig. 3A) indicated a wide distribution of bradoriid
genera during the early Cambrian, with close faunal
Fig. 4. A, pair-group cluster analysis for presence-absence data (Raup-Crick similarity index,
correlation coefficient 0.6361) for 18 bradoriid genera from the early Cambrian, omitting
bradoriid genera that have been documented from only a single palaeoncon-tinental region. B,
pair-group cluster analysis for presence-absence data (Raup-Crick similarity index, correlation
coefficient 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.
316 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
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 Αlvarez 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 specific 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 undertak-en (Figs. 3B-C, 4). The first 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 limita-tions 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 first 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 Αlvarez et al., 2008) Acadobaltic Province sensu Sdzuy
(1972). The main discrep-ancy 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 publica-tions (e.g. Skovsted et al., 2006; Topper et al., 2007;
Zhang, 2007; Hinz-Schallreuter et al., 2008; Dies Αlvarez 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 significantly
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 superficial, 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 taxo-nomic diversity between faunal assemblages
can distort the outcomes. Only two non-endemic bradoriid genera have been
confirmed 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 Liangshanellaoccurring on all three continental
terranes, statistically, Antarctica will display a higher similarity
to Laurentia because of the higher taxonomic diversity of
Australian faunal assemblages.
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 317

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 μm, B, interior view of right valve, scale bar 500 μm, C, external surface ornament, scale bar 10 μm, D, detail of external surface ornament, scale bar 5 μm; E, detail of external
surface ornament, scale bar 5 μm, SAMP44790, sample AJX-M/209.2; F-I, SAMP44791, sample AJX-M/209.2; F, lateral view of left valve, scale bar 500 μm, G, interior view of left valve, scale bar
500 μm, H, detail of interior of left valve, scale bar 10 μm, I, detail of sealed, tubular structures, protruding from the interior of the valve, scale bar 10 μm.

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 definitive 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, figs. 1-8) to be conspecific with Mongolitubulus unispinosa Topper,
Skovsted, Brock and Paterson, 2007 (figs. 6A-J, 7A-C) and Beyrichona
chinensis Shu, 1990 documented by Zhang (2007, p. 145, pl. 17, figs. 9-16) to
be conspecific with Albrunnicola bengtsoni Hinz-Schallreuter, 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 reflect 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-fig. 2; 2003b, fig. 2) and Meert and
Lieberman (2004, fig. 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 non-mineralized arthropods (Hendricks and Lieberman, 2007) pro-
duced results that show significant 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, fig. 2) is supported by some of our results (Figs. 3B-C, 4A).
Hendricks and Lieberman (2007) provided a number of explanations for these
differences,
318 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326

Fig. 6. A-F, Zepaera jagoi sp. nov. All from the Ajax Limestone, Mount Scott Range, all scale bars 200 μm, 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 μm, C, detail of spinose margin, scale bar 20 μm; 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 μm; 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 μm; K, lateral view of right valve, scale bar 500 μm, SAMP44796, sample AJX-M/268.7; L-M, SAMP44797, sample AJX-M/271.3; L, dorsal view of shield, scale bar
500 μm, M, oblique view of shield, scale bar 500 μm.

but it is also worth noting that their analysis included taxa ranging from the
early Cambrian to latest Ordovician, potentially introduc-ing 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), comptalu-tids (Zepaera, Alutella), monasteriids
(Epactridion) and many other genera that are yet to be confidently assigned at
family level (e.g. Onagrocharion, Amphikeropsis and Spinospitella).
Williams et al. (2007) identified a number of faunal assem-blages that
reflected tropical and temperate climates during the
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 319

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 μm, 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 μm, 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 μm; 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 μm, K, detail of external surface ornament, scale bar 20 μm; L, Comptalutid gen et. sp. indet.,
lateral view of right valve, scale bar 100 μm, SAMP44804, sample AJX-M/266; M, Euzepaera sp., lateral view of right valve, scale bar 500 μm, 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 μm, SAMP44808, sample AJX-M/256.5.

Cambrian. The East Gondwana bradoriid assemblage in the early Cambrian
broadly conforms to the climatic distribution patterns identified 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 re-stricted 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.
320 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
4. Systematic palaeontology
All specimens described and illustrated herein are housed in the
palaeontological collection of the South Australian Museum (acro-nym:
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
Öpik, 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 prefix circum, circle and bolus, lump.
In reference to the distinctive circlet of raised papillate struc-tures on the
outer valve surface.
Diagnosis. Moderately to strongly inflated 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 perfora-tion 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 inflated 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 dis-tinct anterodorsal or
posterodorsal nodes or sulci. Valve surface consists of numerous, small
depressions, varying from 7 – 12 µm 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 µm (Fig. 5D-E). Internally, each depression
is sealed by raised cylindrical pillars that protrude from the valve some 10 -
13 µm (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 modified 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 difficult 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 Öpik, 1968. Such difficulties are
exempli-
fied 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 significance. 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 Newfound-land, 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, figs. 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 Trans-Baikal area in the Russian Far-
East and Greenland (Melnikova et al., 1997; Skovsted, 2006) lack an
anterodorsal node and has a valve profile similar to L. circumbolina sp. nov.,
but differ by the surface ornament of fine cancelate ridges. Liangshanella
circumbolina sp. nov. possesses a very distinctive exterior valve ornament
consis-ting of evenly distributed depressions each containing a raised circlet
of closely joined papillae surrounding a single, sub-micron 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 fine-grained siliciclastic rocks (e.g. Siveter and
Williams, 1997; Hou et al., 2002), a mode of preservation not ideal for
preserving fine, 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, fig. 2). However, the exact functional
significance 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, fig. 7) indicated that the lower
levels of the Parara Limestone at Curramulka
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
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, figs 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 Öpik, 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 con-tributions
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 defined, ornamented by a
series of regularly spaced, short, tapering spines.
Description. Valves small (maximum height 810 µm, maximum length
960 µm), postplete, with a well developed, straight hinge line. Valves have a
thin (approximately 7.5 µm in width) later-oadmarginal ridge, entire between
cardinal corners. Lateroadmar-ginal ridge separated from the lateral surface
by a continuous, but weakly defined shallow depression (Fig. 6C). Valve
margin well defined, ornamented by a series of regularly spaced, short,
tapering spines (Fig. 6C). Spines are approximately 10 µm in length and vary
between 13-19 µm in width and are continuous along the valve margin,
between cardinal corners (Fig. 6A). Lateral surface moderately to strongly
inflated, 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, flat
circular welt-like structures, varying in diameter from 26 µm to 63 µm (Fig.
6B). Interior displays a well defined duplicature, entire between the cardinal
corners (Fig. 6F).
Remarks. Comptalutids, like svealutids, frequently exhibit mor-
phologically 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
morpho-logical information will undoubtedly play an important role in the
taxonomic resolution of many comptalutid taxa. New soft-part preservation
has assisted in the taxonomic classification of the comptaluid Kunyangella
Huo, 1965 documented from the Chengjiang Lagerstätte 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
321
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-Schall-reuter
(1999) examined 240 specimens of Zepaera rete and 221 specimens of
Flemingopsis dua for ontogenetic analysis. The difficulties 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 µm, 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 spine-like 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 Beyrichoni-dae. The beyrichonids can be
discriminated from the hipponichar-ionids by having a less inflated shield,
with subdued anterior and posterior lobes that are frequently less ventrally
extended (Hinz-Schallreuter, 1993a, c; Siveter and Williams, 1997). The close
morphological similarity between the two families was recognised by Öpik
(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),
322 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
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, figs. 9-10). Based on the differences in shield inflation 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 inflated shield and
prominent lobation displayed by Quadricona madonnae gen. et. sp. nov.
supports des-ignation to the Hipponicharionidae.
Quadricona gen. nov.
Etymology. Derived from the Latin prefix 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 lateroad-marginal 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 defined 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 Hinz-Schallreuter, 1993a,
Andresia Hinz-Schallreuter, 1993a and Neokun-mingella Zhang, 1974,
typically exhibit confluent anterior and pos-terior 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, fig. 10F-G).
However, the degree and development in shield lobation between the two
genera is strikingly different. The discrete short, nipple-like projections dis-
played by Quadricona gen. nov. is a unique morphological character-istic 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, lateroadmar-ginal ridge is entire between cardinal corners.
Anterodorsal and posterodorsal nodes are discrete, short, sharp, nipple-like
pro-tuberances (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 µm 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 unequiv-ocally 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 150.26-151.71
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 fieldwork 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 field 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 field 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, fig. 10, reproduced here in Fig. 3A). The manuscript benefited from the
constructive reviews of two anonymous reviewers.
Appendix A
Global distribution of bradoriid genera of the major palaeoconti-nents
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 informa-tion from Morocco (Hinz-Schallreuter, 1993a),
Australia (Opik, 1968; Fleming, 1973; Jones and McKenzie, 1980; Hinz-
Schallreuter,
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326 323

1999; Skovsted et al. 2006; Topper et al., 2007), Antarctica (Rode et al., 2003; and Bassler, 1931; Dies Αlvarez et al., 2008), West Avalonia (Siveter and
Wrona, 2009), Armorica (Gozalo et al., 2004; Vannier et al., 2005), Siberia Williams, 1997), East Avalonia (Williams and Siveter, 1998), South China
(Melnikova et al., 1997), Laurentia (Siveter et al., 1996; Siveter and Williams, (Hou et al. 2002; Zhang, 2007) and Kazakhstan (Melnikova et al. 1997).
1997; Skovsted 2006), Baltica (Ulrich
324 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
References
Babcock, L.E., Peng, S., Geyer, G., Shergold, J.H., 2005. Changing perspectives on Cambrian
chronostratigraphy and progress toward subdivision of the Cambrian System. Geosciences
Journal 9, 101–106.
Babcock, L.E., Peng, S., 2007. Cambrian chronostratigraphy: Current state and future plans.
Palaeogeography, Palaeoclimatology, Palaeoecology 254, 62–66.
Babcock, L.E., Peng, S., Sanchi Ellwood, B.B., 2009. Progress towards completion of a
Cambrian chronostratigraphic scale. 14th International Field Conference of the Cambrian
Stage Subcommission Working Group. Maly Karatau Range, Kazakhstan, pp. 7–8
(Abstracts and Short papers).
Bengtson, S., Conway Morris, S., Cooper, B.J., Jell, P.A., Runnegar, B.N., 1990. Early
Cambrian fossils from South Australia. Memoirs of the Association of Australasian
Palaeontologists 9, 1–364.
Brasier, M.D., Cowie, J., Taylor, M., 1994. Decision on the Precambrian-Cambrian boundary
startotype. Episodes 17, 3–8.
Brock, G.A., Alexander, E.M., Paterson, J.R., Jago, J.B., Gatehouse, C.G., 2006. Mt Scott
Range and Ajax Mine. In: Jago, J.B., Zhang, Wenlong (Eds.), South Australia 2006. XI
International Conference of the Cambrian Stage Subdivision Working Group. Geological
Society of Australia. South Australian Division, Adelaide.
Brock, G.A., Engelbretsen, M.J., Jago, J.B., Kruse, P.D., Laurie, J.R., Shergold, J.H., Shi, G.R.,
Sorauf, J.E., 2000. Palaeobiogeographic affinities of Australian Cambrian faunas. Memoirs
of the Association of Australasian Palaeontologists 23, 1–61.
Chapman, F., 1918. Ostracoda from the Upper Cambrian Limestone of South Australia.
Proceedings of the Royal Society of Victoria 31, 108–112 (new series).
Conway Morris, S., 1986. The community structure of the middle Cambrian Phyllopod Bed
(Burgess Shale). Palaeontology 29 423-267.
Copeland, M.J., 1986. Bullaluta kindlei n. gen., n. sp. (Ostracoda, Archaeocopida) from Zone 5
(Late Cambrian, Cedaria-Crepicephalus) of the Cow Head Group, western Newfoundland.
Current Research, Part B. Geological Survey of Canada (Paper 86-1B).
Dies Αlvarez, M.E., Gozalo, R., Cederstrom, P., Ahlberg, P., 2008. Bradoriid arthropods from
the lower-middle Cambrian of Scania, Sweden. Acta Palaeontologica Polonica 53 (4), 647–
656.
Evans, K.R., Rowell, A.J., 1990. Small Shelly Fossils from Antarctica: An Early Cambrian
Faunal Connection with Australia. Journal of Paleontology 64, 692–700.
Fleming, P.J., 1973. Bradoriids from the Xystridura Zone of the Georgina Basin, Queensland.
Publications of the Queensland Geological Survey 356: Professional Papers, 31, pp. 1–9.
Gozalo, R., Hinz-Schallreuter, I., 2002. Biostratigraphy and palaeobiogeography of the
Cambrian genus Hipponicharion (Ostracoda). Paläontologishe Zeitscrift 76, 65–74.
Gozalo, R., Dies, M.A., Chirivella, J.B., 2004. New occurrence of the family Hipponichar-
ionidae (Bradoriida, Arthropoda), in the lower and middle Cambrian of the Cadenas
Ibéricas, Spain. Geobios 37, 191–197.
Gravestock, D., 1984. Archaeocyatha from the lower parts of the Cambrian carbonate sequence
in South Australia. Memoirs of the Association of Australasian Palaeontologists 2, 1–139.
Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001. PAST: palaeontological statistics software
package for education and data analysis. Palaeontologia Electronica 4, 1–9.
Hendricks, J.R., Lieberman, B.S., 2007. Biogeography and the Cambrian radiation of
arachnomorph arthropods. Memoirs of the Association of Australasian Palaeontol-ogists
34, 461–471.
Hinz, I., 1991. On Capricambria cornucopiae gen. et sp. nov. Stereo-Atlas of Ostracod Shells 18
(16), 65–68.
Hinz, I., 1992a. On Pejonesia sestina (Fleming). Stereo-Atlas of Ostracod Shells 19 (2), 5–8.
Hinz, I., 1992b. On Monasterium oepiki Fleming. Stereo-Atlas of Ostracod Shells 19 (29),
123–130.
Hinz, I., 1993. Evolutionary trends in archaeocopid ostracods. In: McKenzie, K.G., Jones, P.J.
(Eds.), Ostracoda in the Earth and life Sciences. A.A. Balkema, Rotterdam, pp. 3–12.
Hinz, I., Jones, P.J., 1992. On Tubupestis tuber Hinz and jones gen. et sp. nov. Stereo Atlast of
Ostracod Shells 19, 9–12.
Hinz-Schallreuter, I., 1993a. Cambrian ostracods mainly from Baltoscandia and Morocco.
Archiv für Geschiebekunde 1 (369, 370), 385–448.
Hinz-Schallreuter, I., 1993b. Ostracodes from the Middle Cambrian of Australia. Neues
Jahrbuch für Geologie und Paläontologie Abhandlungen 188, 305–326.
Hinz-Schallreuter, I., 1993c. Ein Mittelkambrischer hesslandonider Ostracod sowie zur
Morphologie und systematischen Stellung der Archaeocopa. Archiv für Geschie-bekunde 1
(359-350).
Hinz-Schallreuter, I., 1999. Systematics and phylogeny of the Oepikalutidae (Cambrian
Ostracoda). Greifswalder Geowissenschaftliche Beiträge 6, 23–53.
Hinz-Schallreuter, I., Jones, P.J., 1994. Gladioscutum lauriei n. gen. et n. sp. (Archae-ocopida)
from the Middle Cambrian of the Georgina Basin, central Australia. Paleontologicheskii
Zhurnal 68, 361–375.
Hinz-Schallreuter, I., Gozalo, R., Linan, E., 2008. New bradorid arthropods from the Lower
Cambrian of Spain. Micropalaeontology 53 (6), 497–510.
Hinz-Schallreuter, I., Schallreuter, R., 2009. Phylogeny of Phosphatocopa. Memoirs of the
Association of Australasian Palaeontologists 37, 151–164.
Hirschmann, N., 1909. Beitrag zur Kenntnis der Ostracodenfauna des Finnischen Meerbusens.
Mediterranean Society Fauna flora Fennica 35, 282–296.
Holmer, L.E., Popov, L.E., Wrona, R., 1996. Early Cambrian lingulate brachiopods from glacial
erratics of King George Island (South Shetland Islands), Antarctica. In: Gaździck, A.i (Ed.),
Palaeontological Results of the Polish Antarctic Expedition, Part II: Palaeontologia
Polonica, 55, pp. 37–50.
 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
Hou, Xianguang, Bergstrom, J., 1991. The arthropods of the Lower Cambrian Chengjiang fauna,
with relationships and evolutionary significance. In: Simo-netta, A.M., Conway Morris, S.
(Eds.), The early evolution of Metazoa and the significance of problematic taxa. Cambridge
University Press, Cambridge, pp. 179–187.
Hou, Xianguang, Siveter, D.J., Williams, M., Xianghong, Feng, 2002. A monograph of the
bradoriid arthropods from the Lower Cambrian of SW China. Transactions of the Royal
Society of Edinburgh, Earth Science 92, 347–409.
Hou, Xianguang, Siveter, D.J., Williams, M., Waloszek, D., Bergström, J., 1996. Appendages of
the arthropod Kunmingella from the early Cambrian of China: its bearing on the systematic
position of the bradoriids and the fossil record of the Ostracoda. Philosophical Transactions
of the Royal Society of London B 351, 1131–1145.
Hou, Xianguang, Williams, M., Siveter, D.J., Siveter, D.J., Aldridge, R.A., Sansom, R., 2010.
Soft-part anatomy of the Early Cambrian bivalve arthropods Kunyangella and
Kunmingella: significance for the phylogenetic relationships of Bradoriida. Proceedings of
the Royal Society, Series B. doi:10.1098/rspb.2009.2194.
Huo, Shicheng, 1956. Brief notes on Lower Cambrian Archaeostraca from Shensi and Yunnan.
Acta Palaeontologica Sinica 4, 425–445 (In Chinese with English abstract).
Huo, Shi-cheng, 1965. Additional notes on Lower Cambrian Archaeostraca from Shensi and
Yunnan. Acta Palaeontologica Sinica 13, 291–307.
Huo, Shi-cheng, Shu, De-gan, Zhang, Xi-guang, Cui, Zhi-lin, Tong, Hao-wen, 1983. Notes on
Cambrian bradoriids from Shaanxi, Yunnan, Sichuan, Guizhou, Hubei and Guangdong.
Journal of Northwest University 39, 56–75.
Jago, J.B., Zang, Wen-long, Sun, X., Brock, G.A., Paterson, J.R., Skovsted, C.B., 2006. A
review of the Cambrian biostratigraphy of South Australia. Palaeoworld 15, 406–423.
Jell, P.A., Jago, J.B., Gehling, J.G., 1992. A new conocoryphid trilobite from the Lower
Cambrian of the Flinders Ranges, South Australia. Alcheringa 16, 189–200.
Jiang, Z.W., 1982. Archaeostracoda. In: Luo, H.L., Jiang, Z.W., Wu, X.C., Song, X.L., Ouyang,
L. (Eds.), The Sinian-Cambrian boundary in eastern Yunnan, China, Yunnan Peoples
Publishing House, Yunnan, 46-47, 211-215.
Jones, P.J., Kruse, P.D., 2009. New Middle Cambrian bradoriids (Arthropoda) from the
Georgina Basin, central Australia. Memoirs of the Association of Australasian
Palaeontologists 37, 55–86.
Jones, P.J., Laurie, J.R., 2006. Bradoriida and Phosphatocopida (Arthropoda) from the Arthur
Creek Formation (Middle Cambrian), Georgina Basin, central Australia. Memoirs of the
Association of Australasian Palaeontologists 32, 205–223.
Jones, P.J., McKenzie, K.G., 1980. Queensland Middle Cambrian Bradoriida (Crustacea): new
taxa, palaeobiogeography and biological affinities. Alcheringa 4, 203–225.
Kobayshi, T., Kato, F., 1951. On the ontogeny and the ventral morphology of Redlichia
chinensis with description of Alutella nakamurain new gen. and sp. Japanese Journal of
Geology and Geography 11, 161–169.
Landing, E., 1994. Precambrian-Cambrian boundary global stratotype ratified and a new
perspective of Cambrian time. Geology 22, 179–182.
Landing, E., Peng, S.C., Babcock, L.E., Geyer, G., Moczydlowska-Vidal, M., 2007. Global
standard names for the lowermost Cambrian series and stage. Episodes 30, 287–289.
Laurie, J., 2006. Early Middle Cambrian trilobites from Pacific Oil & Gas Baldwin 1 well,
southern Georgina Basin, Northern Territory. Memoirs of the Association of Australasian
Palaeontologists 32, 127–204.
Lee, Yu-wen, 1975. On the Cambrian Ostracoda with new material from Sichuan, Yunnan and
southern Shaanxi, China. Professional papers on Stratigraphy and Palaeontology 2, 37–72.
Lieberman, B.S., 1997. Early Cambrian paleogeography and tectonic history: A biogeographic
approach. Geology 25, 1039–1042.
Lieberman, B.S., 2003a. Biogeography of the Trilobita during the Cambrian radiation: deducing
geological processes from trilobite evolution. Special Papers in Palaeon-tology 70, 59–72.
Lieberman, B.S., 2003b. Taking the pulse of the Cambrian radiation. Integrative and
Comparative Biology 43, 229–237.
Maas, A., Waloszek, D., Müller, K.J., 2003. Morphology, ontogeny and phylogeny of the
Phosphatocopina (Crustacea) from the Upper Cambrian “Orsten” of Sweden. Fossils and
Strata 49, 1–238.
Mansuy, H., 1912. Etude Geologique de Yun-Nan Oriental. Pt. 2, Palaeontologie.
Memoire service geologie 1'Indochine 1, 1–23.
Martinsson, A., 1979. Albrunnicola, a new name for the Cambrian ostracode genus Longispina
Andres, 1969. Lethaia 12, 27.
Matthew, G.F., 1886. On the Cambrian faunas of Cape Breton and Newfoundland.
Transactions of the Royal Society of Canada 4, 147–157.
Matthew, G.F., 1902. Ostracoda of the basal Cambrian rocks in Cape Breton. Canadian Record
of Science 8, 437–470.
McKenzie, K.G., Aangel, M.V., Becker, G., Hinz-Schallreuter, I., Kontrovitz, M., Parker, A.R.,
Schallreuter, R.E.L., Swanson, K.M., 1999. Ostracodes. In: Savazzi, E. (Ed.), Functional
Morphology of the Invertebrate Skeleton. John Wiley & Sons Ltd., Chichester, pp. 459 –
507.
McKerrow, W.S., Scotese, C.R., Brasier, M.D., 1992. Early Cambrian continental
reconstructions. Geological Society of London Journal 149, 599–606.
Meert, J.G., Lieberman, B.S., 2004. A palaeomagnetic and palaeobiogeographical perspective on
latest Neoproterozoic and Early Cambrian tectonic events. Journal of the Geological
Society of London 161, 1–11.
Melnikova, L.M., 1988. Nekotoryye bradoriidy(Crustacea) iz botomskogo yarusa vostochnogo
Zabaykal’ya. Paleontologicheskiy Zhurnal 114–117.
Meert, J.G., Lieberman, B.S., 2008. The Neoproterozoic assembly of Gondwana and its
relationship to the Ediacaran-Cambrian radiation. Gondwana Research 14, 5–21.
325
Melnikova, L.M., Siveter, D.J., Williams, M., 1997. Cambrian Bradoriida and Phospha-tocopida
(Arthropoda) of the former Soviet Union. Journal of Micropalaeontology 16, 179–191.
Missarzhevsky, V.V., 1977. Konodonty (?) i fosfatnye problematiki kembriya Mongolii i Sibiri.
Bespozvonochnye Paleozoya Mongolii,. Trudy Sovmestaya Sovetsko-Mon-golskaya
Paleontologicheskaya Ekspeditsiya, pp. 10–19.
Müller, K.J., 1964. Ostracoda (Bradoriina) mit phosphatischen Gehäusen aus dem
Oberkmabrium von Schweden. Neues Jahrbuch für Geologie und Paläontologie
Abhandlungen 121, 1–46.
Müller, K.J., 1979. Phosphatocopine ostracodes with preserved appendages from the Upper
Cambrian of Sweden. Lethaia 12, 1–27.
Müller, K.J., 1982. Hesslandona unisulcata sp. nov. with phosphatized appendages from Upper
Cambrian Orsten of Sweden. In: Bate, R.H., Robinson, E., Sheppard, L.M. (Eds.), Fossil
and Recent Ostracoda. Ellis Horwood, Chichester, pp. 276–304.
Öpik, A.A., 1961. The geology and palaeontology of the headwaters of the Burke River,
Queensland. Bulletin of the Bureau of Mineral Resources. Geology and Geophysics 53, 1–
249.
Öpik, A.A., 1963. Early Upper Cambrian fossils from Queensland. Bureau of Mineral
Resources, Geology and Geophysics 64, 1–133.
Öpik, A.A., 1967. The Mindyallan fauna of northwestern Queensland. Bureau of Mineral
Resources, Australian Bulletin, 74 (1), 1-404, 74 (2), 1-167.
Öpik, A.A., 1968. Ordian (Cambrian) Crustacea Bradoriida of Australia. Bulletin. Bureau of
Mineral Resources. Geology and Geophysics 103, 1–44.
Palmer, A.R., 2005. East-Gondwana/Laurentia trilobite connections — what do they tell us?
Geosciences Journal 9 (2), 75–79.
Palmer, A.R., Rowell, A.J., 1995. Early Cambrian trilobites from the Shackleton Limestone of
the Central Transantarctic Mountains. Paleontological Society. Memoir 45, 1–28.
Paterson, J.R., 2005. Revision of Discomesites and Estaingia (Trilobita) from the Lower
Cambrian Cymbric Vale Formation, western New South Wales: Taxonomic, biostratigraphic
and biogeographic implications. Proceedings of the Royal Society of New South Wales 126,
81–93.
Paterson, J.R., Brock, G.A., 2007. Early Cambrian trilobites from Angorichina, Flinders Ranges,
South Australia, with a new assemblage from the Pararaia bunyerooensis Zone. Journal of
Paleontology 81, 116–142.
Paterson, J.R., Jago, J.B., 2006. New trilobites from the Lower Cambrian Emu Bay Shale
Lagerstätte at Big Gully, Kangaroo Island, South Australia. Memoirs of the Association of
Australasian Palaeontologists 32, 43–57.
Peng, S.C., Babcock, L.E., 2008. Cambrian Period. In: Ogg, J.G., Ogg, G., Gradstein, F.M.
(Eds.), A Concise Geologic Time scale. Cambridge University Press, pp. 37–46.
Pillola, G.L., 1993. The Lower Cambrian trilobite Bigotina and allied genera.
Palaeontology 36, 855–881.
Raymond, P.E., 1935. Leanchoilia and other mid-Cambrian Arthropoda. Bulletin of the Museum
of Comparative Zoology 76, 205–230.
Richter, R., Richter, E., 1940. Die Saukianda – Stufe von Andalusien, eine fremde Fauna im
europaeischen, Ober-Kambrium (Studien im Palaeozoikum der Mittelmeer-Laender 5).
Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft. (88 p).
Rode, A.L., Lieberman, B.S., Rowell, A.J., 2003. A new Early Cambrian bradoriid (Arthropoda)
from East Antarctica. Journal of Paleontology 77, 691–697.
Sdzuy, K., 1972. Das Kambrium der acadobaltischen Faunen-provinz. Zentralblatt für Geologie
und Paläontologie II (1-2), 1–91.
Shergold, J.H., 1996. Cambrian (Chart 1). In: Young, G.C., Laurie, J.R. (Eds.), An Australian
Phanerozoic Timescale. Oxford University Press, Melbourne, pp. 63–76.
Shu, D., 1981. Bradoriids from Guizhou with discussion of several of the principal related
problems. Masters Thesis. Northwest University, 123–129.
Shu, D., 1990. Cambrian and Lower Ordovician Bradoriida from Zhenjiang. Hunnan and
Shaanxi Provinces. Northwest University Press, Xian. (90 p).
Shu, D., Chen, L., 1994. Cambrian palaeobiogeography of Bradoriida. Journal of Southeast
Asian Earth Sciences 9 (3), 289–299.
Shu, D., Vannier, J., Huilin, Luo, Ling, Chen, Xingliang, Zhang, Shixue, Hu., 1999. Anatomy
and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil
Lagerstätte (lower Cambrian; Southwest China). Lethaia 32, 279–298.
Siebold, C.T.E., Stannius, H., 1845. Lehrbuch der Vergleichenden Anatomie. Con Veit, Berlin.
Siveter, D.J., Williams, M., Peel, J.S., Siveter, D.J., 1996. Bradoriida (Arthropoda) from the
early Cambrian of North Greenland. Transactions of the Royal Society of Edinburgh,
Earth Science 86, 113–121.
Siveter, D.J., Williams, M., 1997. Cambrian bradoriid and phosphatocopid arthropods of North
America. Special Papers in Palaeontology 57, 1–69.
Siveter, D.J., Waloszek, D., Williams, M., 2003. An Early Cambrian phosphatocopid crustacean
with three-dimensionally preserved soft parts from Shropshire, England. Special Papers in
Palaeontology 70, 9–30.
Skovsted, C.B., 2006. Small Shelly Fauna from the upper Lower Cambrian Bastion and Ella
Island Formations, North-East Greenland. Journal of Paleontology 80, 1087–1112.
Skovsted, C.B., Brock, G.A., Paterson, J.R., 2006. Bivalved arthropods from the Lower
Cambrian Mernmerna Formation of South Australia and their implications for the
identification of Cambrian ‘small shelly fossils’. Memoirs of the Association of Australasian
Palaeontologists 32, 7–41.
Skovsted, C.B., Brock, G.A., Holmer, L.E., Paterson, J.R., 2009. First report of the early
Cambrian stem group brachiopod Mickwitzia from East Gondwana. Gondwana Research
16, 145–150.
Steiner, M., Zhu, M., Weber, B., Geyer, G., 2001. The Lower Cambrian of Eastern Yunnan:
Trilobite-based biostratigraphy and related faunas. Acta Palaeontologica Sinica 40, 63–79
(Supp).
Stigall, A.L., 2008. Bicarinellata a new name for Bicarinella Rode, Lieberman and Rowell, 2003
(Arthropoda, Bradoriida) preoccupied by Bicarinella Waterhouse, 1966
326 T.P. Topper et al. / Gondwana Research 19 (2011) 310–326
(Mollusca, Gastropoda) and Bicarinella Akopyan, 1976 (mollusca, Gastropoda).
Journal of Paleontology 82 (6), 1220.
Streng, M., Ebbestad, J.O.R., Moczydlowska, M., 2008. A Walcottella-like bradoriid
(Arthropoda) from the lower Cambrian of Sweden. GFF 130, 11–19.
Sylvester Bradley, P.C., 1961. Archaeocopida. Q100–10.5. In: Moore, R.C. (Ed.), Treatise on
Invertebrate Paleontology, Pt. Q, Arthropoda 3. Geological Society of America. Boulder
and University of Kansas Press, Lawrence.
Tate, R., 1892. The Cambrian fossils of South Australia. Transactions of the Royal Society of
South Australia 15, 183–189.
Topper, T.P., Brock, G.A., Skovsted, C.B., Paterson, J.R., 2009. Shelly fossils from the lower
Cambrian Pararaia bunyerooensis Zone, Flinders Ranges, South Australia. Memoirs of the
Association of Australasian Palaeontologists 37, 199–246.
Topper, T.P., Skovsted, C.B., Brock, G.A., Paterson, J.R., 2007. New bradoriids from the lower
Cambrian Mernmerna Formation, South Australia: systematics, biostratigraphy and
biogeography. Memoirs of the Association of Australasian Palaeontologists 33, 67–100.
Ulrich, E.O., Bassler, R.S., 1931. Cambrian bivalved Crustacea of the Order Conchostraca.
Proceedings of the US National Museum 78 (4), 1–130.
Vannier, J., Williams, M., Álvaro, J.J., Vizcaïno, D., Monceret, S., Monceret, E., 2005. New Early
Cambrian bivalved arthropods from southern France. Geological Magazine 142, 751–763.
Williams, M., Siveter, D.J., 1998. British Cambrian and Tremadoc bradoriid and
phosphatocopid arthropods. Monograph of the Palaeontolographical Society London 1–49
(Publ. No. 607, part of Vol. 153 for 1998).
Williams, M., Siveter, D.J., Popov, L.E., Vannier, J.M.C., 2007. Biogeography and affinities of
the bradoriid arthropods: Cosmopolitan microbenthos of the Cambrian seas.
Palaeogeography, Palaeoclimatology, Palaeoecology 248, 202–232.
Williams, M., Siveter, D.J., Jose Salas, M., Vannier, J.M.C., Popov, L.E., Ghobadi Pour, M.,
2008. The earliest ostracods: the geological evidence. Senckenbergiana Lethaea 88
(1), 11–21.
Whittaker, J.E., 1973. On Elofsonia baltica. In: Sylvester-Bradley, P.C., Siveter, D.J. (Eds.), A
Stereo-Atlas of Ostracod Shells, Volume 1. Department of Geology in the University of
Leicester, England (Part 3, August 31st).
Wrona, R., 2003. Early Cambrian mollusks from glacial erratics of King George Island, West
Antarctica. Polish Polar Research 24, 181–216.
Wrona, R., 2004. Cambrian microfossils from glacial erratics of King George Island, Antarctica.
Acta Palaeontologica Polonica 49, 13–56.
Wrona, R., 2009. Early Cambrian bradoriidae and phophatocopidae arthropods from King
George Island, West Antarctica; Biogeographic implications. Polish Polar Research 30 (4),
347–377.
Zang, W.-L., Jago, J.B., Lin, T.-R., 2001. Early Cambrian acritarchs, trilobites and
archaeocyathids from Yalkalpo 2, eastern Arrowie Basin, South Australia. Report Book,
2001/00002. Department of Primary Industry and Resources, South Australia, p. 41.
Zhang W.-T., 1974. Bradoriida. In: Nanjing Institute of Geology and Palaeontology, Academica
Sinica (Ed.), Handbook of Stratigraphy and Palaeontology of Southwest China. Science
Press, Beijing, 107-111
Zhang, W.-T., 2003. Cambrian correlation between North America and China based on trilobite
and conodont faunas. Acta Palaeontologica Sinica 42, 305–316.
Zhang, W.-T., Lu, Y.-H., Zhu, Z.-L., Qian, Y.-Y., Lin, H.-L., Zhou, Z.-Y., Zhang, S.-G., Yuan, J.-
L., 1980. Cambrian trilobite faunas of southwestern China. Palaeontologia Sinica 159, 1–
497 (new series B, no. 16).
Zhang, X.-G., 1987. Moult stages and dimorphism of Early Cambrian bradoriids from Xichuan,
Henan, China. Alcheringa 11, 1–19.
Zhang, X.G., 2007. Phosphatised Bradoriids (Arthropoda) from the Cambrian of China.
Palaaeontographica Abteilung A: Palaeozoologie-Stratigraphie 281, 93–173.
Zhang, X.G., Pratt, B.R., 1993. Early Cambrian ostracode larvae with a univalved carapace.
Science 262, 93–94.
Zhang, Z., Robson, S.P., Emig, C., Shu, D., 2008. Early Cambrian radiation of brachiopods:
A perspective from South China. Gondwana Research 14, 241–254.
Zhuravlev, A. Yu., 1995. Preliminary suggestions on the global Early Cambrian zonation. In:
Geye, G., Landing, E. (Eds.), MOROCCO ’95—The Lower-Middle Cambrian standard of
western Gondwana, Beringeria Special Issue 2. Freunde der Würzburger
Geowissenschaften e. V., Wrzberg, pp. 147–160.
Zhuravlev, A. Yu., Gravestock, D.I., 1994. Archaeocyaths from Yorke Peninsula, South
Australia and archaeocyathan Early Cambrian zonation. Alcheringa 18, 1–54.