Early Cambrian Trilobites From Angorichina, Flinders Ranges, South Australia, With A New Assemblage From The Pararaia Bunyerooensis Zone

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EARLY CAMBRIAN TRILOBITES FROM ANGORICHINA, FLINDERS RANGES, SOUTH AUSTRALIA, WITH A NEW ASSEMBLAGE
FROM THE PARARAIA BUNYEROOENSIS ZONE
Paterson, John R
ABSTRACT-
Trilobites from the Lower Cambrian succession at Angorichina in the eastern Flinders Ranges, South Australia, are described. Silicified material
from the Mernmerna Formation reveals the presence of a new assemblage from the Pararaia bunyerooensis Zone, including the eponymous
species. Yorkella aff. australis, Eoredlichia sp., Redlichia sp., and the new species Wutingaspis euryoptilos and Yunnanocephalus macromelos.
Trilobites of the Pararaia bunyerooensis Zone show a strong affinity with those from the Yu'anshan Member of the Heilinpu Formation in
Chengjiang and Jinning Counties, Yunnan Province, southwest China. The Pararaia bunyerooensis Zone is correlated with the Yunnanocephalus
Assemblage subzone (upper Eoredlichia-Wutingaspis Zone) of the Chiungchussuan (= Qiongzhusian) Stage of China. Additional trilobites from
Angorichina include Elicicola calva from the Wilkawillina Limestone, Estaingia occipitospina (Jell) new combination from the Oraparinna Shale,
and Redlichia guizhouensis Zhou from the Wirrealpa Limestone. Australian Early Cambrian trilobite biozonation is reviewed, with discussion of
distinct assemblages within the Pararaia janeae Zone that have the potential for zonal subdivision, and evidence to support the placement of the
northern Australian Ordian/Early Templetonian Stage within the late Early Cambrian.
A possible paedomorphic lineage between Pararaia bunyerooensis and P. janeae is proposed. Adult specimens of P. janeae retain juvenile
characteristics of the progenitor P. bunyerooensis. Retardation in onset of maturity in P. janeae resulted in the attainment of a larger adult size
than in P. bunyerooensis, indicating the former species evolved via neoteny.
INTRODUCTION
EARLY CAMBRIAN trilobites were first described from the Flinders Ranges over a century ago (Etheridge, 1905), yet there remains a dearth of
literature on the abundant and diverse faunas of this region. Early taxonomic works include those of Etheridge (1905, 1919) and Walcott (in
Howchin, 1920). Daily (1956) provided the first significant biostratigraphic scheme for the Cambrian succession in South Australia when he
established 12 informal "faunal assemblages" that have subsequently been used for intra- and intercontinental correlations for many years.
Unfortunately, Daily provided no formal taxonomic treatment of the largely unpublished faunas, nor did he include any precise stratigraphic
ranges of the diagnostic taxa. More recently. Jell (in Bengtson et al., 1990) provided the most comprehensive study to date on the Early
Cambrian trilobites of South Australia. Jell recorded 18 genera and 32 species, many of which were left under open nomenclature, and erected
four trilobite zones: Abculiellci huoi (base), Pararaia ialei, Pararaia humerooensis. and Paruraia janeae.
Other taxonomic studies of Early Cambrian trilobites from the Flinders Ranges include Pocock (1970), Jell et al. (1992), Zhang et al. (2001), and
Paterson and Edgecombe (2006). In erecting the family, genera, and species of the Emuellidae, Pocock (1970) described Balcnracania flintlerxi
Pocock, 1970 from the lower Billy Creek Formation, the type locality of which occurs at Angorichina Station. In a systematic revision of the
Emuellidae, Paterson and Edgecombe (2006) regarded Balcoracania flinders! to be a junior subjective synonym of B. dailyi Pocock, 1970. Jell et
al. (1992) described the conocoryphid Atops rupertensis Jell, Jago, and Gehling, 1992 from the upper Mernmerna Formation and basal
Oraparinna Shale at various localities in the Flinders Ranges; this species was previously referred to as Atops sp. nov. by Jell (in Bengtson et al.,
1990). In revising Chinese, Moroccan, and Australian species of Ptirabadiellii Chang, 1966 and Abadiella Hupé, 1953, Zhang et al. (2001)
considered that specimens of Abiuliella huoi (Chang. 1966) illustrated by Jell (hi Bengtson et al., 1990) belong to a new species of Wutingaspis
Kobayashi, 1944, W. jelli Zhang et al., 2001, and thus proposed that the Australian Abadiella huoi Zone be changed to the Wutingaspis jelli Zone.
However, Jago et al. (2002a) subsequently contested the conclusions of Zhung et al. (2001) and advocated that the Australian species belongs to
Abacliella huoi as originally assigned by Jell (in Bengtson et al.. 1990). In recent years there have been several studies on the biostratigraphy.
correlation, and paleobiogeography of the Early Cambrian trilobites of the Flinders Ranges and other areas of Australia, including Brock et al.
(2000), Gravestock and Shergold (2001 ), Zang et al. (2001 ), Jenkins et al. (2002), and Jago et al. (2002b).
This study focuses on the taxonomy and biostratigraphy of the trilobite faunas from the Lower Cambrian succession at Angorichina Station in the
eastern Flinders Ranges, South Australia. Silieified material from the Mernmerna Formation reveals the presence of a new assemblage from the
Paruraia bunyerooensis Zone, including the new species Wutingaspis euryoptilos and Yunnanocephalus nuicromelos.
LOCALITY AND STRATIORAPHY
Trilobites were collected from the Wilkawillina Limestone, Mernmerna Formation. Oraparinna Shale and Wirrealpa Limestone outcropping at
Angorichina Station in the Wirrealpa area of the eastern Flinders Ranges, South Australia. Angorichina Station is situated approximately 11 km
east of Blinman and borders with Wirrealpa Station to the east and the Flinders Ranges National Park to the south. The Cambrian carbonate and
clastic sediments in this area represent part of the Arrowie Basin in the Adelaide Geosyncline (Priess, 1999); a detailed sequence stratigraphie
framework has been developed for the Cambrian of the Arrowie Basin (see Gravestock and Hibburt, 1991: Gravestock and Cowley, 1995;
Boucher, 1997; Gravestock and Shergold, 2001; Zang, 2003; Zang et al., 2004) (Fig. 1 ).
Specimens were sampled from measured stratigraphie sections through the Wilkawillina Limestone and Mernmerna Formation (section MMF),
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Oraparinna Shale (section OS), and Wirrealpa Limestone (section WL). The 60 m thick Mernmerna Formation at section MMF (including one
sampled horizon in the underlying Wilkawillina Limestone. MMF/0.0) (base of section coordinates: 31°11'38.4''S, 138°52'28.7''E; map datum:
WGS84), is located on the eastern side of The Bunkers, approximately 1 km south of Balcoracana Creek (Figs. 2, 3). The Oraparinna Shale section
OS (coordinates: 31°12'26.7''S, 138°52'38.9''E; map datum: WGS84) is also located on the eastern side of The Bunkers, approximately 2.5 km
south of Balcoracana Creek and 1.5 km south of section MMF (Fig. 2). The Wirrealpa Limestone section WL (base of section coordinates:
31°09'21.8''S. 138°53'23.7''E; map datum: WGS84), coincides with section 1 of Youngs (1977, 1978) at Balcoracana Creek (Figs. 2, 3).
The stratigraphy and sedimentology of the Mernmerna Formation at section MMF have been recently documented by Brock and Paterson
(2004). They correlated section MMF with the Third Plain Creek Member of the middle Mernmerna Formation at Wilkawillina Gorge (type
section in the Bunkers Graben) based on similar lithologies, sedimentary structures, and the occurrence of the trilobite Pararaia bunyerooensis
Jell in Bengtson et al., 1990, and concluded that the middle Mernmerna Formation at section MMF unconformably overlies the Wilkawillina
Limestone and is disconformably overlain by the Bunkers Sandstone (Clarke, 1986a, 1989, 1990: Gravestock and Cowley, 1995). Further
evidence to support the unconformity between the Wilkawillina Limestone and Mernmerna Formation at the MMF section includes the presence
of a thin red microstromatolite horizon at the contact of these units, representing the Flinders Unconformity (James and Gravestock, 1990;
Gravestock and Cowley, 1995), in addition to the co-occurrence of the trilobite Elicicola calva Jell (in Bengtson et al., 1990) and brachiopod
Askepasma Laurie, 1986 in horizon MMF/0.0 (discussed below). This indicates that a considerable hiatus exists at the MMF section with the
absence of sequence 61.2 transgressive deposits of Gravestock and Cowley (1995, p. 23), i.e., Second Plain Creek Member of the Wilkawillina
Limestone (Clarke. 1986b) and the Six Mile Bore and Linns Springs members of the lower Mernmerna Formation (Clarke, 1986a, 1990). The
Mernmerna Formation at section MMF is dominated by interbedded black turbiditic wackestone-packstone and laminated lime silt and mud with
rare sandstone beds (Fig. 4). Grain flow deposits are common and contain intraclasts, ooids, and peloids. Irregular and nodular bedding is
relatively common and slump structures are also evident. Based on Clarke's (1990) depositional model, the Third Plain Creek Member of the
Mernmerna Formation was deposited in anaerobic and dysaerobie zones of the lower slope (= Sequence euro61.2 highstand deposits of Gravestoek
and Cowley, 1995). The MMF section contains an abundance of silicified and phosphatic macro-and microfossils, including a variety of trilobites
(documented herein), the helcionellid mollusc Tannuella elinorae Brock and Paterson, 2004, in addition to other molluscs, brachiopods,
archaeocyaths, chancelloriids, sponge spicules, and other enigmatic small shelly fossils which will be described in future studies.
The 42 m thick exposure of Oraparinna Shale at section OS (Figs. 2, 4) is relatively thin compared to the thickness of ~200 m at Wilkawillina
Gorge; however, little or no outcrop of this unit is typical along the eastern side of The Bunkers (P. S. Moore, 1979: Gravestoek and Cowley,
1995). The Oraparinna Shale at section OS conformably overlies the Bunkers Sandstone, with the lower 38 m (true thickness) representing fissile
green-, gray- and buff-colored shales with common limestone concretions containing rare trilobite and brachiopod fragments. The remaining 4 m
consists of buff-colored siltstones containing abundant specimens of the trilobite Estuingia ocdpitiixpina (Jell in Bengtson et al., 1990) along with
various centimeter-sized molluscs. The Edeowie Limestone Member overlies the Oraparinna Shale at section OS; the relationship between these
units has been the source of much debate over the last 40 years. Dalgarno and Johnson (1962, 1965) defined the dolomitic Edeowie Limestone
Member as the basal unit of the Billy Creek Formation. P. S. Moore (1979) later reassigned the member to the Oraparinna Shale. Priess (1999)
reinstated the Edeowie Limestone Member as the basal member of the Billy Creek Formation based on P. S. Moore's (1979, fig. 6) mapping, which
showed that the member overlies a low-angle truncation surface in the Hawker Group. This suggests that the Edeowie Limestone Member
unconformably overlies the Oraparinna Shale at the OS section.
The stratigraphy and sedimentology of the Wirrealpa Limestone have been documented in great detail by Youngs (1977, 1978). The Wirrealpa
Limestone is relatively consistent in thickness (105-140 m) in the Flinders Ranges, conformably overlying red beds of the Billy Creek Formation,
and is conformably overlain by the micaceous shales, siltstones, and sandstones of the Moodlatana Formation (Youngs, 1977, 1978; Brock and
Cooper, 1993; Gravestoek and Cowley, 1995). The unit represents a transgressive-regressive carbonate sequence displaying lagoonal and ooid
bank megafacies. The lagoonal facies association is characterized by lime mudstone and wavy-bedded to nodular limestone in a calcareous silly
matrix, with subordinate skeletal, peloidal and oolitic beds, calcimicrobe build-ups, and stromatolite and columnar thrombolite bioherms. The
ooid bank facies association predominantly consists of oolitic and oncolitic lithologies (Youngs, 1977, 1978). Section WL (Figs. 2-4) is equivalent
with Section 1 of Youngs (1977, 1978) at Balcoracana Creek. This section was chosen because of its completeness, representing a 'typical section'
of the Wirrealpa Limestone (Youngs, 1978. p. 68); it also contains an abundance of fossils, including trilobites. brachiopods, small shelly fossils,
hyoliths, and stromatolites. As noted by Youngs (1978). this section shows an upward progression from mostly line-grained carbonates, nodules,
and calcareous siltstones to stromatolite bioherms and the first oolitic beds, followed by predominantly allochemical rocks with stromatolites and
minor micrite beds. The upper part of the section consists of alternating thin oosparites with thicker beds of nodules in silts, and the up-permost
beds are represented by calcareous green-gray siltstones that grade into the red beds of the Moodlatana Formation. Although fossils have not been
formally described from this section, a few studies have documented faunas of the Wirrealpa Limestone from surrounding areas, e.g., the trilobite
Redlichia guizhouensis Zhou (in Lu et al., 1974; Jell HI Bengtson et al., 1990). cyanobacterial-archaeocyathan-radiocyathan bioherms (Kruse,
1991b), and a variety of shelly fossils (Brock and Cooper, 1993).
BIOSTRATIGRAPHY AND CORRELATION
Review of Early Cambrian trilobite biozonation in South Australia.-The Early Cambrian trilobite zones established by Jell (in Bengtson et al.,
1990) were introduced as a preliminary biozonation in an effort to provide a basis for future biostratigraphic studies. Jell (in Bengtson et al.,
1990, p. 14) hoped "that future work will refine this zonation, fill in the obvious gaps and test its applicability to other areas." These zones have
now been widely accepted and utilized in global syntheses on the Cambrian timescale (e.g., Zhuravlev, 1995; Shergold, 1997; Geyer and
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Shergold, 2000; Zhuravlev and Riding. 2001; Zhang, 2003), although these syntheses show considerable disparity in the correlation of the
South Australian zones with other regions. Early Cambrian intercontinental correlation has been a major problem for decades; complicating
factors include the high endemicity of Early Cambrian trilobite faunas related to biofacies control, paucity of biostratigraphically useful faunas
in key regions, substantial hiatuses in the stratigraphie record, and inconsistencies in taxonomic nomenclature (Palmer, 1998b; Geyer, 2001).
The Early Cambrian trilobite faunas from South Australia have closest affinities with those of South China, with IO congeneric and six conspecific
occurrences (Jell in Bengtson et al., 1990; described herein). Correlation of the Early Cambrian stages and trilobite zones of Siberia, China, and
South Australia is presented in Figure 5.
The Abadiella huoi Zone represents the oldest zone in South Australia and contains the following species: Abadiella huoi, Elicicola calva, Yorkella
australis (Woodward, 1884), Alanisia gnillermoi (Richter and Richter, 1940). and Eoredlichia sp. Taxa from this zone have been recorded from
the Parana Liinestone at Curramulka. Horse Gully (Ardrossan), and Kulpara in the Stansbury Basin on Yorke Peninsula, from the Ajax
Limestone (Mt Scott Range) and Wilkawillina Limestone (Wilkawillina Gorge, Wirrealpa Mine, and Yalkalpo-2 drillhole) in the Arrowie Basin
(Jell in Bengtson et al., 1990; Zang et al., 2001; Gravestock et al., 2001). Elicicola culva also occurs in the Wilkawillina Limestone immediately
below the red microstromatolite horizon of the Flinders Unconformity, which marks the contact with the Mernmerna Formation at the MMF
section (Figs. 2-4). Although the biostratigraphic ranges of most species of the Abadiella huoi Zone (excluding Eoredlichia sp.) extend up into the
younger Pararaia tatet Zone (Jell in Bengtson et al., 1990), the occurrence of E. calva at the base of the MMF section is taken to represent the A.
huoi Zone. This is further supported by the co-occurrence of the paterinid brachiopod Askepasina in the same horizon (i.e., MMF/ 0.0). The last
appearance datum (LAD) of Askepasina in the Stansbury Basin occurs at the top of the Pelagiella subangulata mollusc assemblage, which
correlates with the A. huoi Zone (Gravestock et al., 2001). This confirms the suggestion by Brock and Paterson (2004, p. 136) that the
Mernmerna Formation unconformably overlies the Wilkawillina Limestone at the MMF section.
The South Australian A. huoi Zone correlates with the Chinese Parabadiella Zone based on the occurrence of A. huoi in both regions (Zhang, 1985,
2003; Jell in Bengtson et al., 1990; Steiner et al., 2001; Zhang et al., 2001). These zones correlate with the latest Atdabanian-earliest Botoman
of Siberia, based on correlation of South Australian and Siberian archaeocyathan faunas (Zhuravlev and Gravestock, 1994; Zhuravlev, 1995).
The Pararaia tatei Zone comprises 10 species, including four overlapping species from the older Abadiella huioi Zone (Jell in Bengtson et al.,
1990). The base of the P. tatei Zone corresponds to the first appearance datum (FAD) of the eponymous species. This zone occurs in the Parara
Limestone at Curramulka, Horse Gully (Ardrossan), and Kulpara in the Stansbury Basin, and in the Ajax Limestone (Mt Scott Range),
Mernmerna Formation (Willochra, Wilkawillina Gorge and Yalkalpo-1 drillhole), and Wilkawillina Limestone (Wirrealpa Mine and west of
Wirrealpa Springs) in the Arrowie Basin (Jell in Bengtson et al., 1990; Zang et al., 2001; Gravestock et al., 2001). Jell (in Bengtson et al., 1990,
p. 16) correlated the P. tatei Zone with the Chinese Eoredlichia-Wutingaspis Zone based on the occurrence of Eoredlichia shensiensis (Chang,
1966) in both regions. These zones appear to correlate with the early Botoman (Bergeroniellus micmacciformis-Erbiella Zone) of Siberia based on
the correlation of older (Abadiella huoi Zone equivalent) archaeocyathan faunas (discussed above). However, Jell (in Bengtson et al., 1990, p.
17) considered that the Pararaia tatei Zone should be considered Atdabanian based on the genera Prouktaspis Repina (in Khomentovskii and
Repina, 1965), Egyngolia Korobov, 1980, and Pararaia Kobayashi, 1942 ( = Tanniiolaspis Zadorozhnaya in Zhuravleva et al., 1967, fide Jell in
Bengtson et al., 1990). According to Korobov (1989) and Zhuravlev (1995), Egyngolia from Mongolia occurs together with Botoman
archaeocyaths and the trilobites Binodaspis Lermontova, 1951, Erbiopsis Lermontova, 1940, Inouyina Poletaeva, 1936, Jakutus Lermontova,
1951, Kadyella Pokrovskaya, 1959, and Proerbia Lermontova, 1940, thus supporting an early Botoman age for the P. tatei Zone.
The Pararaia bunyerooensis Zone has been the most poorly understood of the Early Cambrian trilobite zones of South Australia. This was largely
due to its supposed low diversity, consisting of the eponym and an indeterminate redlichiid, and its limited occurrence in the Flinders Ranges.
This zone has only been previously recorded from the Mernmerna Formation at Bunyeroo Creek, and from the Third Plain Creek Member of the
middle Mernmerna Formation at Wilkawillina Gorge (Jell in Bengtson et al., 1990; Gravestock and Cowley, 1995). A new assemblage from the P.
bunyerooensis Zone consisting of six species has now been discovered at Angorichina and is described below.
The Pararaia janeae Zone has the highest diversity of all of the South Australian trilobite zones, containing 19 species (Jell in Bengtson et al.,
1990). Jell (in Bengtson et al., 1990) based this zone primarily on the faunule at Bunyeroo Creek, containing the species Pararaia janeae Jell (in
Bengtson et al., 1990), Hebediscina yuqingensis (Zhang in Zhang et al., 1980). Serrodiscus gravestocki Jell (in Bengtson et al., 1990), Alops
rupertensis, Kootenia diutina Fritz, 1972, Estaingia bilobata Pocock, 1964, and Paleofossusl sp. For convenience, Jell extended this zone beyond
this assemblage to incorporate the stratigraphie ranges of Estaingia species, i.e., E. bilobata and E. occipitospina. This included the trilobite
faunas of the White Point Conglomerate and Emu Bay Shale on Kangaroo Island (Pocock, 1964, 1970; Jell in Bengtson et al., 1990; Nedin, 1995).
Jell also suspected that the emuellid Balcoracania dailyi from the White Point Conglomerate was synonymous with B. flindersi from the
Warragee Member of the lower Billy Creek Formation at Balcoracana Creek. Angorichina-this has now been confirmed by Paterson and
Edgecombe (2006)-and thus incorporated these species into the P. janeae Zone. This implies that the trilobites Estaingia bilobata and
Balcoracania dailyi have considerable stratigraphic ranges and may be of limited biostratigraphic utility if the P. janeae Zone were to be further
subdivided. The trilobite fauna from the Cymbric Vale Formation in western New South Wales, originally deseribed by Öpik (1975b), is
considered to be of P. janeae Zone age (Jago et al., 1997; Paterson, 2005).
There is, in fact, evidence to suggest further refinement of the Pararaiu janeae Zone. A trilobite assemblage containing the species Estaingia
occipitoxpina and Korobovia ocellata Jell (in Bengtson et al., 1990) from the upper Oraparinna Shale apparently does not overlap with the
Pararaia janeae assemblage (Jell in Bengtson et al., 1990). This has also been observed in continuous stratigraphic sections through the
Mernmerna Formation and Oraparinna Shale sampled by the authors in the Elder Range (personal data). Further detailed biostratigraphic work
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of these assemblages is needed before any future subdivision is attempted. Jell (in Bengtson et al., 1990) also documented an additional
assemblage containing the trilobites Xela drena Jell (in Bengtson et al., 1990), Micmaccopsis separata Jell (in Bengtson et al., 1990),
Micmaccopsis alba Jell (in Bengtson et al., 1990), Redlichia endoi Lu, 1950, and Yorkella sp. nov., referred to herein as the 'Micmaccopsis
assemblage,' from an archaeocyathid bioherm within the upper Wilkawillina Limestone at Wirrealpa Mine, which he tentatively assigned to the
P. janeae Zone. Three lines of evidence indicate the temporal separation of the Pararaia janeae assemblage and the Micmaccopsis assemblage.
Firstly, the occurrence of Hebediscina yuqingensis in the Pararaia janeae assemblage permits correlation with the early Tsanglangpuan of
China, based on the occurrence of H. yuqingensis in the Jiumengchong ( = Niutitang) Formation in eastern Guizhou (Zhang et al., 1980; Peng
and Babcock, 2001); the occurrence of Atops Emmons, 1844 (=Ivshiniellus Korobov, 1966, fide Jell et al., 1992) from the latest Aldanian ( =
early Botoman, sensu Rozonov and Sokolov, 1984) in Siberia also supports this correlation (Korobov, 1966, 1973). Secondly, the occurrence of
Redlichia enden in the Micmaccopsis assemblage allows correlation with the Chinese Palaeolenus Zone (Zhang et al., 1980; Zhou and Yuan,
1980); the Palaeolenus Zone equates with the latest Botoman Bergeroniaspis ornata Zone of Siberia based on correlation of archaeocyathan
faunas (Yuan and Zhang, 1981; Zhuravlev, 1995; Zhang, 2003). Thirdly, Xela Jell (in Bengtson et al., 1990) has been described from the Lesser
Himalaya in India by Jell and Hughes (1997) and is considered to be of late Tsanglangpuan (Palaeolenus Zone) age (Hughes and Jell, 1999;
Hughes et al., 2005).
Trilobites from younger horizons in the Early Cambrian of South Australia are rare, but include Recllichia ffiiizliouenxix from the Wirrealpa
Limestone, and Onaraspis rubra Jell (in Bengtson et al., 1990) from the Moodlatana Formation. Previously, the stratigraphic range of R.
guizhouensis in South Australia was poorly understood, since it had been recorded from only a single horizon in the Wirrealpa Limestone by Jell
(in Bengtson et al., 1990). However, specimens are now known from a new locality (WL section) at Angorichina, which demonstrates that R.
guizhouensis ranges throughout the majority of the Wirrealpa Limestone (Fig. 4). Redlichia guizhouensis occurs in the latest Lungwangmiaoan
Stage in South China (Zhou in Lu et al., 1974; Yin and Li, 1978; Zhou and Yuan, 1980; Peng and Babcock, 2001), suggesting that the Wirrealpa
Limestone is of similar age. Zhang (1985, 2003) and Jell (in Bengtson et al., 1990) both considered that R. guizhouensis may be a synonym of R.
nobilis Walcott, 1905, suggesting that the R. guizhouensis Zone is equivalent to the R. nobilis Zone of China.
Jell (in Bengtson et al., 1990) described Onaraspis rubra from the Moodlatana Formation at Wilkawillina Gorge and a locality northwest of
Wirrealpa (NMVPL89). He tentatively correlated the rocks containing O. rubra with the northern Australian Ordian Stage-now regarded as part
of the Ordian/Early Templetonian Stage (see Kruse et al., 2004, p. 18-19 for detailed discussion)-based on the occurrence of congeners O.
somniurna Öpik, 1968 and O. adusta Öpik, 1968 from central and Western Australia, respectively (Öpik, 1968).
Previously, the Australian Ordian/Early Templetonian Stage (=Xystridura templetonensis/Redlichia chinensis Zone) was considered to be early
Middle Cambrian (e.g., Shergold, 1996; references therein), but mounting evidence suggests that this stage should be regarded as latest Early
Cambrian, based on correlation with the Lungwangmiaoan Stage of China. Firstly, Öpik (1970) described Redlichia chinensis Walcott, 1905 from
the Ordian of northern Australia, which allows correlation with the chinensis Zone of the mid-Lungwangmiaoan in China (Zhang, 2003).
However, Kruse et al. (2004, p. 19) have questioned Öpik's (1970) assignment to this species, since he illustrated only three specimens.
Therefore, correlation of the Redlichia chinensis Zone between Australia and China remains tentative. More convincing evidence comes from the
occurrence of the Ungulate brachiopods Karathele napuru (Kruse, 1990) and Vandalotreta djagoran (Kruse, 1990) in the Wirrealpa and Ramsay
Limestones of South Australia (Brock and Cooper, 1993: Gravestock et al., 2001) as well as several Ordian/Early Templetonian successions in
northern Australia, including the Tindall Limestone in the Daly Basin, Montejinni Limestone in the Wiso Basin, and the Top Springs Limestone
and Gum Ridge Formation in the Georgina Basin (Kruse, 1990, 1991a, 1998). This permits direct correlation with the Redlichia guihouensis
Zone (latest Lungwangmiaoan) of China (discussed above). These species and other associated ungulate brachiopods co-occur with trilobites such
as Redlichia forresti (Etheridge in Foord, 1890), Redlichia gumridgensis Laurie (in Kruse et al., 2004), Xystridura negrina Öpik, 1975a, and X.
verticosa Öpik, 1975a (Kruse, 1990, 1991a, 1998; Kruse et al., 2004) that are considered characteristic of the Ordian/Early Templetonian
Stage. An unidentified species of Redlichia Cossmann, 1902 from the Tindall Limestone in the Northern Territory (Kruse, 1990, pl. 1) may be
conspecific with Redlichia guizhouensis, but the limited number and fragmentary nature of the specimens from the Tindall Limestone precludes
confirmation. The occurrence of Bathynotus holopygux (Hall, 1859) in the Ordian/Early Templetonian of central Australia (Shergold and
Whittington, 2000) and the Kaili Formation of Guizhou, South China (Yuan et al., 2002), provides additional evidence to support the correlation
of the Ordian/Early Templetonian with the Lungwangmiaoan. Yuan et al. (2002) record B. holopygus from the Ovatoryctocara
granulata-Kathynotus holopygus Zone, which they correlate with the late Lungwangmiaoan. This species is also considered to be of late Early
Cambrian (Olenellux Zone) age in Laurentia (Palmer, 1998a; Shergold and Whittington, 2000). However, until the Early-Middle Cambrian
boundary is selected and ratified, correlation of the Ordian/Early Templetonian Stage with other intercontinental Early Cambrian stages
remains tentative.
The Pararaia bunyerooensis Zone.-A new silicified assemblage of this zone is described from the Third Plain Creek Member of the Mernmerna
Formation at the MMF section, located on the eastern side of The Bunkers, approximately 1 km south of Balcoracana Creek on Angorichina
Station (Figs. 2-4). The assemblage contains the following species, in order of appearance: Pararaia bunyerooensis Jell (in Bengtson et al., 1990),
Yunnamocephalus macromelos n. sp., Yorkella aff. australis, Redlichia sp., Wutingaspis euryoptilos n. sp., and Eoredlichia sp. The base of this
zone corresponds to the FAD of the eponymous species. At the MMF section, the FAD of P. bunyerooensis corresponds to horizon MMF/8.8, 5.1 m
(true thickness) above the base of the section (Fig. 4).
The trilobites from the MMF section are interpreted as being an allochthonous assemblage deposited by turbiditic conditions in a lower slope
environment, following Clarke's (1990) depositional model of the Mernmerna Formation at Wilkawillina Gorge. Taphonomic data are limited;
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but trilobites from the MMF section commonly occur in highly concentrated silicified 'trash bands' and larger sclerites (>10 mm) are often
fragmentary. Syndepositional breakage is also evident in crushed specimens of the large helcionellid mollusc Tannuclla elinorae that are
preserved in silicified crusts. Thin sections reveal the presence of graded beds containing randomly oriented trilobite cuticle. Thin (
Trilobites of the Pararaia bunyerooensis Zone show a strong affinity with those of the Chiungchussuan (=Qiongzhusian) Stage of China, especially
taxa from the Yu'anshan Member of the Heilinpu Formation in Chengjiang and Jinning Counties, Yunnan (Zhang, 1987; Shu et al., 1995;
Babcock and Zhang, 2001; Steiner et al., 2001; Zhang et al., 2001). The association of the genera Eoredlichia Chang (in Lu and Dong, 1952),
Wutingaspis Kobayashi, 1944 and Yunnanocephalus Kobayashi, 1936 is considered to be an important age-diagnostic assemblage of the
Eoredlichia-Wutingaspis Zone in South China (Zhang, 1985, 2003; Steiner et al., 2001; Zhang et al., 2001). The genus Wutingaspis ranges
throughout the Eoredlichia-Wutingaspis Zone; however, Steiner et al. (2001) and Zhang et al. (2001) have observed that this zone can be
subdivided into distinct lower and upper generic assemblages. Zhang et al. (2001) subdivided the Eoredlichia-Wutinguspis Zone into two
subzones: the lower 'Tsunydiscus Assemblage-subzone'; and the upper 'Chengjiangaspis Assemblage-subzone.' Subsequently, Steiner et al. (2001)
also subdivided the Eoredlichia-Wutingaspis Zone into two subzones: the lower 'Tsunydiscus Taxon-range Subzone'; and the upper
'Yunnanocephalus Assemblage subzone.' The two subzones of Zhang et al. (2001) and Steiner et al. (2001) appear to represent the same
assemblages of the Yu'anshan Member, but the use of the name 'Yunnanocephalus Assemblage subzone' for the upper assemblage seems more
appropriate due to the rare occurrence of Chenjiangaspis Zhang and Lin (in Zhang et al., 1980) in eastern Yunnan (Steiner et al., 2001).
Therefore, the co-occurrence of Eoredlichia. Wutingaspis, and Yunnanocephalus in the South Australian Pararaia bunyerooensis Zone permits
correlation with the Yunnanocephalus Assemblage subzone (upper Eoredlichia-Wutingaspis Zone) of South China.
Jell (in Bengtson et al., 1990) considered cranidia referred to as Dolerolenus? sp. nov. by Öpik (1975b) to be synonymous with Abadiella huoi.
These cranidia are herein assigned to Wutingaspis euryoptilos from the Pararaia bunyerooensis Zone at Angorichina. Unfortunately, the precise
stratigraphic horizon of the specimens from the Parara Limestone at Kulpara is unknown. Jell (in Bengtson et al., 1990) believed them to have
come from a part of the Kulpara section containing taxa of the Abadiella huoi and Pararaia tatei zones, but it is possible that the specimens came
from higher in the section, above the range of Pararaia tatei (Woodward, 1884).
SYSTEMATIC PALEONTOLOGY
The morphological terminology employed follows Whittington and Kelly (1997). Registered specimens are housed in the paleontological
collections of the South Australian Museum. Adelaide (prefix SAMP).
Order REDLICHIIDA Richter, 1932
Suborder REDLICHIINA Richter, 1932
Superfamily REDLICHIOIDEA Poulsen, 1927
Family REDLICHIIDAE Poulsen, 1927
Genus REDLICHIA Cossmann, 1902
Type species.-Hoeferia noetlingi Redlich, 1899, Early Cambrian, Khussak Group, Salt Range, Pakistan.
REDLICHIA GUIZHOUENSIS Zhou in Lu et al., 1974
Figure 6
Redlichia guizhouensis ZHOU IN LU, CHANG, CHIEN, CHU, LIN, ZHOU, QIAN, ZHANG, AND YUAN, 1974, p. 85, pl. 31, fig. 10: YIN AND LI,
1978, p. 400, pl. 147, fig. 1; LU, 1981, pl. 1, fig. 20: JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 267,
fig. 179.
Olenellus sp. ETHERIDGE, 1905, p. 247, pl. 25, fig. 1.
Olenellus? sp. ETHERIDGE, 1919, p. 382, pl. 39. fig. 1.
Figured material.-Four fragmentary cranidia, SAMP40581-40584 (Fig. 6.1-6.5); three librigenae, SAMP40585-40587 (Fig. 6.6-6.11); one
rostral plate, SAMP40588 (Fig. 6.12, 6.13); one fragmentary thoracic pleura, SAMP40589 (Fig. 6.14).
Occurrence.-Wirrealpa Limestone, WL section horizons: WL/ 4, WL/12, WL/13, WL/17, WL/48, WL/52.3, WL/53.6, WL/56.2, WL/57.4,
WL/63.1, WL/65.2, WL/74.8, WL/83, WL/103.6, WL/ 147; 2.8-103.9 m (true thickness) above the base of the section (Fig. 4).
Discussion.-The occurrence of this species in the Wirrealpa Limestone has been adequately documented by Jell (in Bengtson et al., 1990). Large,
well-preserved silicified librigenae from the WL section display a wider (tr.) genal field compared to the small, fragmentary librigena described
and illustrated by Jell (in Bengtson et al., 1990, fig. 179d).
REDLICHIA sp.
Figure 7
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Redlichiid indet. 1 JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 273, fig. 182b-d.
Figured material.-Four fragmentary cranidia. SAMP40590-40593 (Fig. 7.1-7.4); three librigenae, SAMP40594-40596 (Fig. 7.5-7.9); three
pygidia, SAMP40597-40599 (Fig. 7.10-7.13).
Occurrence.-Pararaia bunyerooensis Zone, Third Plain Creek Member, Mernmerna Formation, MMF section horizons: MMF/ 44.5, MMF/54.6,
MMF/58.3, MMF/64.5, MMF/93, MMF/103; 25.5-59.1 m (true thickness) above the base of the section (Fig. 4).
Discussion.-This species shows affinities with Redlichia takooensis Lu, 1950, and other indeterminate redlichiid fragments described and
illustrated by Jell (in Bengtson et al., 1990). Shared characteristics with R. takooensis include: narrow (tr.) palpebral area; angle of divergence
and shape of anterior branches of facial suture; length (exsag.) and width (tr.) of preocular field; length (sag., exsag.) of anterior cranidial border:
librigena with broad (tr.) genal field and straight, wide (tr.) posterior margin: pygidium with broad axis not reaching posterior margin and
displaying a bilobed terminal piece; and a single pair of pygidial pleurae terminating as short marginal spines. However, R. takooensis is easily
distinguished in having a well-developed occipital spine and two pygidial axial rings.
Specimens referred to as 'Redlichiid indet. 1' by Jell (in Bengtson et al., 1990) are most certainly attributed to the species from the MMF section
based on the material illustrated by Jell (in Bengtson et al., 1990, fig. 182b-d) and its association with Pararaia bunyerooensis in the Mernmerna
Formation at Bunyeroo Creek. The two fragmentary cranidia of 'Redlichiid indet. 1' differ only in having a slightly narrower (tr.) palpebral area.
Pygidia of 'Redlichiid indet. 3' documented by Jell (in Bengtson et al., 1990, fig. 182j, 1, m) from the Ajax Limestone at Mt Scott Range also bear
striking similarity to pygidia from the MMF section, based on characteristics mentioned above.
Genus EOREDLICHIA Chang in Lu and Dong, 1952
Type species.-Redlichia intermediata Lu, 1940, Early Cambrian, Chiungchussu Formation, Yunnan, China.
EOREDLICHIA sp.
Figure 8.1-8.7
Figured material.-One fragmentary cranidium, SAMP40600 (Fig. 8.1, 8.2); one fragmentary thoracic pleura, SAMP40601 (Fig. 8.7); four
fragmentary pygidia, SAMP40602-40605 (Fig. 8.3-8.6).
MMF/64.5, MMF/75.2, MMF/93; 31.3-53.3 m (true thickness) above the base of the section (Fig. 4).
Discussion.-Fragmentary silicified material from the Mernmerna Formation makes species identification difficult, although specimens show
similarities with Eoredlichia shensiensis and E. intermediata (Lu, 1940). Jell (in Bengtson et al., 1990, p. 281) considered Pachyredlichia
(=Eoredlichia) zhangshanensis Lin and Yao (in Zhang et al., 1980) to be a junior subjective synonym of E. shensiensis; this synonymy is followed
herein. Palpebral lobe morphology (e.g., shape and exsagittal length) and pustulose ornament of the Angorichina material closely resembles that
of E. shensiensis (e.g., Zhang et al., 1980, pl. 38, figs. 9, 12, 14; Jell in Bengtson et al., 1990, fig. 185a-g). However, cranidia of E. shensiensis
from China (Chang, 1966, pl. 1, figs. 5, 6; Zhang et al., 1980, pl. 38, figs. 9-14; pl. 39, figs. 1, 2, 4) and South Australia (Jell in Bengtson et al.,
1990. fig. 185a-e, g) do not appear to display a stout occipital spine.
The fragmentary pygidia from the Mernmerna Formation (Fig. 8.3-8.6) bear close resemblance to pygidia of Eoredlichia intermediata from
Chengjiang, South China (Shu et al., 1995, figs. 4f, 7b). Similarities include: pygidial outline; sagittal length of axis: and the presence of two
axial rings and a semicircular terminal piece that is slightly narrower (tr.) than the axial rings.
Genus WUTINGASPIS Kobayashi, 1944
Wutingaspis KOBAYASHI, 1944, p. 130; HARRINGTON IN R. C. MOORE, 1959, p. 205; LU, 1961, p. 301; LI, 1978, p. 189; YIN IN YIN AND LI,
1978, p. 406: LI, 1980, p. 45; ZHANG AND LIN IN ZHANG, LU, ZHU, QIAN, LIN, ZHOU, ZHANG, AND YUAN, 1980, p. 159 (for additional
synonymy): LUO, 1981, p. 335; ZHOU, LI, AND QU, 1982, p. 223; LI AND ZHANG IN LI, KANG, AND ZHANG, 1990, p. 44; LUO IN LUO, JIANG,
AND TANG, 1994, p. 124; ZHANG, 1997, p. 441.
Type species.-Wutingaspis tingi Kobayashi, 1944, Early Cambrian, Chiungchussu Formation, Yunnan, China.
Discussion.-The close morphological similarity between the genera Abadiella, Parabadiella, and Wutingaspis has been the source of considerable
debate in recent years (Jell in Bengtson et al., 1990; Steiner et al., 2001; Zhang et al., 2001; Jago et al., 2002a). Jell (in Bengtson et al., 1990)
and Jago et al. (2002a) have been advocates of synonymizing Abadiella and Parabadiella, while Steiner et al. (2001) and Zhang et al. (2001)
consider them to be separate genera. Steiner el al. (2001, p. 70) also commented on the similarity in the cranidial and pygidial morphology of
Parabadiella and Wutingaspis, listing three discriminating criteria to separate them: 1) shape of the preglabellar ridge (=plectrum); 2)
exsagittal length of the palpebral lobes: and 3) posterolateral projection of the fxigenae. The first two criteria are unreliable diagnostic characters
at the generic level because they show interspecific variation. For example, Wutingaspis euryoptilos n. sp. has a very narrow (tr.) plectrum and
the length (exsag.) of the palpebral lobes is 30%-35% glabellar length (sag.), whereas W. tingi has a broad (tr.) plectrum and the length (exsag.)
of the palpebral lobes is 35%-45% glabellar length (sag.). There appears to he a consistent difference in the width (tr.) of the posterolateral
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projection of the fixigena between both genera. Species of Abadiella (=Parabadiella) have a very narrow (tr.) posterolateral projection, whereas
species of Wutingeispis possess a considerably wider (tr.) posterolateral projection. Whether this character should be given generic or species level
significance is uncertain. It is possible that Abadella (=Parabadiella) and Wutingaspis are synonymous, but this interpretation can only be
resolved by conducting a comprehensive revision of these genera, which is beyond the scope of this study.
We agree with the arguments of Jell (in Bengtson et al., 1990) and Jago et al. (2002a) regarding the synonymy of Abadiella and Parabadiella.
Furthermore, specimens described and illustrated as Abadiella huoi by Jell (in Bengtson et al., 1990) should be maintained as that species and
not assigned to a new species of Wutingaspis, W. jelli, as described by Zhang (in Zhang et al., 2001).
WUTINGASPIS EURYOPTILOS new species
Figures 8.8-8.17, 9.1-9.7
Dolerolenus? sp. nov. ÖPIK, 1975B, p. 41, pl. 7, tig. 2.
Diagnosis.-Wutingaspis with anterior sections of facial suture diverging anteriorly at 90°-100° between ? and ß glabella moderately tapering,
anterior width (tr.) at midlength of frontal lobe 60%-65% occipital ring width (tr.); palpebral lobe length (exsag.) 30%-35% glabellar length
(sag.): palpebral area width (tr.) at e 70%-80% adjacent glabellar width; small intergenal spine developed at distal extremity of posterior border.
Librigenal field width (tr.) approximately 70%-75% adjacent librigenal width. Pygidium with three axial rings and a short (sag.), rounded
terminal piece; second and third axial rings with medial depression.
Description.-Cranidium of moderate size, up to 19 mm in length (sag.); trapezoidal in outline; low convexity (sag., tr.). Anterior margin
moderately curved; posterior margin (excluding occipital ring) straight, directed posterolaterally. Anterior sections of facial suture curved,
diverging anteriorly at 90°-100° between ? and ß, then strongly convergent anteriorly between ß and a posterior sections of facial suture long,
curved anteriorly, widely divergent posteriorly, with well-developed suturai ridge. Glabella moderately tapering, anterior width (tr.) at
midlength of frontal lobe 60%-65% occipital ring width (tr.); frontal lobe rounded; moderate convexity (tr.), weak convexity (sag.); sagittal
length (including LO) approximately 15% cranidial length. Axial furrow shallow, narrow (tr.); preglabellar furrow shallow, narrow (sag.,
exsag.). Glabellar furrows well developed; S1 strongly impressed, deepening abaxially, proximal portion straight, becoming strongly convex
anteriorly at distal extremity, directed anterolaterally abaxially. width (tr.) approximately 30%-35% adjacent glabellar width (tr.); S2 and S3
same as for S1. Occipital ring of moderate convexity (tr.), flattened sagittally; length (sag.) 15%-20% glabellar length; short, stout posteromedial
occipital spine directed slightly posteriorly; posterior margin moderately convex posteriorly. SO shallow and transverse medially, then directed
anterolaterally and deepened into apodemal pits abaxially, wide (sag., exsag.). Preglabellar field flat, length (sag.) approximately 10% cranidial
length: plectrum well developed, narrow (tr.). Preocular field gently downsloping, flat to weakly convex (exsag.), maximum length (exsag.) 25%
cranidial length (sag.). Anterior border weakly convex, length (sag.) 10% cranidial length; anterior border furrow moderately curved, shallow,
narrow (sag., exsag.). Palpebral lobe well developed, strongly convex (tr.). length (exsag.) 30%-35% glabellar length (sag.), width (tr.) 25%-30%
lobe length, anterior tip situated opposite L3, posterior tip situated opposite midlength of L1; palpebral furrow shallow, narrow (tr.). Eye ridge
well developed, forming a continuation of the palpebral lobe, strongly convex (exsag.), gently curved anteriorly, ridges diverge posteriorly at
120°-130°, proximal end merges with lateral margin of frontal lobe with anteroproximal margin of eye ridge continuing onto anterior margin of
frontal lobe to form narrow (sag., exsag.) parafrontal band of low relief. Palpebral area weakly convex, width (tr.) at e 70%-80% adjacent
glabellar width. Postocular area short (exsag.), of subequal length (exsag.) to sagittal length of occipital ring. Posterolateral projection of fixigena
very wide, width (tr.) 20%-25% e-e, gently downsloping, strongly tapering abaxially. Posterior border strongly convex (exsag.), slightly
expanding abaxially; small intergenal spine developed at distal extremity of posterior border (Fig. 8.16); posterior border furrow deep, very wide
(exsag.), expanding abaxially.
Rostral plate and hypostome unknown.
Librigena of moderate size, up to 22 mm in length (including genal spine): width (tr.) approximately 40%-50% length (including genal spine);
lateral margin moderately curved to base of genal spine, then becoming straight and slightly deflected abaxially; posterior margin straight
before strongly curving onto genal spine. Genal field of low convexity, width (tr.) approximately 70%-75% adjacent librigenal width. Eye socle of
low dorsal elevation. Lateral border moderately convex, raised above genal field, widening (tr.) posteriorly, becoming widest at genal angle,
anteriormost dorsal portion of border sharply tapers to a point; lateral and posterior border furrows more of a change in slope than distinct
furrows. Posterior border weakly convex, short (tr.), narrow (exsag.), of uniform width (exsag.). Genal spine long, length approximately
30%-35% librigenal length (including spine): narrow base (tr.), strongly tapering posteriorly; straight. Librigenal doublure width (tr.) mimics
dorsal surface of lateral and posterior librigenal border: sharp ridge developed on lateral margin of doublure, terminating at base of genal spine,
representing a change in slope from rounded lateral margin to wide (tr.). slightly concave adaxial portion.
Thorax unknown.
Pygidium moderately small, up to 6 mm in length (sag.), weakly convex (sag., tr.). Axis with curved lateral margins, reaching maximum width
(tr.) at second axial ring; three axial rings and a short (sag.), rounded terminal piece; first and second axial rings separated by well-developed
axial furrow that is wide (sag.) medially, becoming narrower (exsag.) and deeper abaxially; second axial ring with medial depression: third axial
ring with medial depression that is wider (tr.) than depression in second axial ring; articulating half ring poorly preserved. Pleural regions with
at least one pair of distinct pleurae: pleural furrow narrow and of moderate depth. Borders and margins obscure.
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Entire dorsal surface of exoskeleton densely covered in granules. Preglabellar, preocular and librigenal fields covered in genal caeca.
Etymology.-Greek eurys, broad, widespread, and optilos, eye; referring to the wide (tr.) palpebral area of the fixigenae.
Type material.-Holotype: fragmentary cranidium, SAMP40606 (Fig. 8.8). Paratypes: six fragmentary cranidia, SAMP40607-40612 (Figs.
8.9-8.12, 8.16, 8.17, 9.1); five librigenae, SAMP40613-40617 (Figs. 8.13-8.15, 9.2-9.5); two fragmentary pygidia, SAMP40618-40619 (Fig.
9.6, 9.7).
Type locality.-Mernmerna Formation (MMF section), Hawker Group; base of section coordinates: 31°11'38.4''S, 138°52'28.7''E; map datum:
WGS84. on the eastern side of The Bunkers, approximately 1 km south of Balcoracana Creek on Angorichina Station, Flinders Ranges, South
Australia (see Figs. 2, 3).
Type stratum.-Third Plain Creek Member of the Mernmerna Formation, MMF section, horizon MMF/93; 53.3 m (true thickness) above the base of
the section (Fig. 4).
MMF/64.5, MMF/75.2, MMF/93; 31.3-53.3 m (true thickness) above the base of the section (Fig. 4).
Discussion.-Wutingaspis euryoptilos is remarkably similar to the type species of Wutingaspis, W. tingi Kobayashi, 1944. Cranidial features of W.
tingi that are shared with W. euryoptilos include: well-developed sutural ridge on posterior section of facial suture (Chang, 1962, pl. 1, fig. 4a;
Zhang et al., 1980, pl. 42, figs. 2, 4); short, stout posteromedial occipital spine: eye ridges diverge posteriorly at 120°-130°; anteroproximal
margin of eye ridge continues around frontal lobe to form narrow (sag., exsag.) parafrontal band (Zhang et al., 1980, pl. 42, fig. 2); very wide
(tr.) posterolateral projection of fixigenae, width (tr.) 20%-25% e-e; small intcrgcnul spine developed at distal extremity of posterior border
(Zhang et al., 1980, pl. 42, fig. 6; Shu et al., 1995, fig. 20. Glabellar furrow morphology of these species is also very similar, although some
cranidia of W. tingi display a medial connection of S1 (e.g., Chang, 1962, pl. 1, fig. 4a; Zhang et al., 1980, pl. 42. fig. 4; Shu et al., 1995, fig. 2f;
Steiner et al., 2001, pl. 2, fig. 6). However, it is important to note that these specimens of W. lingi are preserved as internal molds. The pygidium
of W. tingi also bears close similarity to that of W. euryoptilos in possessing three axial rings and a short (sag.), rounded terminal piece, with the
second and third axial rings showing a medial depression (Zhang et al., 1980, pl. 41, fig. 10; pl. 42, fig. 3; Steiner et al., 2001, pl. 2, fig. 7).
Wutingaspis tingi can be distinguished from W. euryoptilos, however, by the following features: anterior sections of facial suture diverge
anteriorly at 60°-70° (90°-100° in W. euryoptilos); glabella gently tapers, anterior width (tr.) at midlength of frontal lobe 75%-80% occipital
ring width (tr.) (60%-65% in W. euryoptilos); palpebral lobe length (exsag.) 35%-45% glabellar length (sag.) (30%-35% in W. euryoptilos); and
palpebral area width (tr.) at e 55%-65% adjacent glabellar width (tr.) (70%-80% in W. euryoptilos).
The majority of Wutingaspis species described from China are poorly known, in most cases represented by no more than two or three cranidia,
thus making species comparison difficult. The better known species that are represented by different sclerite types, and in some cases partially
articulated exoskeletons, e.g., W. conditus Kobayashi, 1944 (Chang, 1966, pl. 2, fig. 1), W. malungensis Lu, 1961 (Zhang et al., 1980, pl. 40, fig.
7), W. sichuanensis Li, 1978 (pl. 91, fig. 9), can be easily differentiated from W. euryoptilos by having a narrower (tr.) palpebral area, no
intergenal spine, narrower (tr.) librigenal field, and two pygidial axial rings.
Dolerolenus? sp. nov. described and illustrated by Öpik (1975b, p. 41, pl. 7, fig. 2) is considered herein to be synonymous with Wutingaspis
euryoptilos. The cranidia are almost indistinguishable, although the cranidium of Dolerolenus? sp. nov. displays stronger genal caeca on the
preocular field; this difference may be attributed to different styles of preservation. Particular characteristics of Dolerolenus? sp. nov. such as
glabellar shape and furrows, width (tr.) of the palpebral area, and the granulose ornamentation strongly support this synonymy. Jell (in
Bengtson et al., 1990) and Zhang et al. (2001) considered Dolerolenus? sp. nov. to be a synonym of Abadiella (=Parabadiella) huoi. The cranidia
of Dolerolenus? sp. nov. and Abadiella huoi are similar in many respects, but those of A. huoi differ in having a narrower (tr.) palpebral area
(e.g., Chang, 1966, pl. 1, figs. 1, 2; Zhang et al., 1980, pl. 46, figs. 1-4, 6; Jell in Bengtson et al., 1990, figs. 183a, c, d, j, 184s-u, w, x, z, ab).
Unfortunately, the posterolateral projection of the fixigena is not preserved in cranidia of Dolerolenus? sp. nov., making it impossible to
determine whether it has a wide (tr.) projection like Wutingaspis euryoptilos or narrow (tr.) projection like Abadiella huoi.
Genus YORKELLA Kobayashi, 1942
Yorkella KOHAYASHI, 1942, p. 492; JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 288; ZHANG, 1997, p.
444.
Type species.-Conocephalites australis Woodward, 1884, Early Cambrian, Parara Limestone, Yorke Peninsula, South Australia.
YORKELLA aff. AUSTRALIS (Woodward, 1884)
Figure 9.8-9.20
Figured material.-Two fragmentary cranidia, SAMP40620-40621 (Fig. 9.8, 9.9): four librigenae, SAMP40622-40625 (Fig. 9.10-9.14); four
pygidia, SAMP40626-40629 (Fig. 9.15-9.20).
MMF/54.6, MMF/64.5, MMF/82.6, MMF/93, MMF/103; 25.5-59.1 m (true thickness) above the base of the section (Fig. 4).
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Discussion.-The species of Yorkella from the MMF section bears close resemblance to the type species Y. australis (Woodward, 1884), which has
been well described and illustrated by Jell (in Bengtson et al., 1990, figs. 189, 190). Although only two fragmentary cranidia are known from
the MMF section (Fig. 9.8, 9.9). they are indistinguishable from the cranidia of Y. australis illustrated by Jell (in Bengtson et al., 1990, figs.
1891-q, 190a, b, 1-t). The librigenae of Y. australis differ only in having a narrower (tr.) lateral border and wider (tr.) genal field (Jell in
Bengtson et al., 1990, figs. 189s. t, 190k). The smallest illustrated librigena of Yorkella from the MMF section (Fig. 9.14) is similar to those of Y.
australis; however, larger librigenae (Fig. 9.10, 9.11) show that the lateral border widens and the genal field becomes narrower throughout
ontogeny. The only major morphological difference between pygidia from the MMF section and Y. australis is the ratio of sagittal length to width
(tr.), i.e., sagittal length 45% and 60% maximum width (tr.), respectively. Specimens from the MMF section most likely represent a new species.
However, until additional well-preserved cranidia are found, this species is left under open nomenclature.
Superfamily ELLIPSOCEPHALOIDEA Matthew, 1887
Family ESTAINGIIDAE Öpik, 1975a
Discussion.-The subfamily Estaingiinae Öpik, 1975a was recently elevated to family level by Jell (in Jell and Adrain, 2003, p. 334, n6) based on
the discovery of a nomenclatural error between the synonymous genera Estaingia Pocock, 1964 and Hsuaspis Chang in Lu et al., 1965.
Westrop and Landing (2000, p. 863) have recently discussed the differing classifications of ellipsocephaloid families, in particular those of Öpik
(1975b), Zhang et al. (1980), Jell (in Bengtson et al., 1990), and Geyer (1990). Westrop and Landing (2000) noted that the majority of these
classifications relied upon glabellar outline to diagnose ellipsocephaloid families and subfamilies. For example. Jell (in Bengtson et al., 1990)
assigned Hsuaspis (=Estaingia) to the Ichangiidae (=Estaingiidae, fide Jell in Jell and Adrain, 2003) and Pararaia to the Protolenidae on the basis
of glabellar outline. Westrop and Landing (2000) commented that there are limits to the utility of glabellar outline in the suprageneric
classification of ellipsocephaloids. They observed that Hsuaspis (=Extaingia) and Pararaia have strong similarities in cephalic and pygidial
morphology, as previously noted by Palmer (in Palmer and Rowell, 1995, p. 16), but that these characters were absent or differed greatly in
Protolenus Matthew, 1892 and related genera. Hence, Westrop and Landing (2000) concluded that similarities in glabellar outline between
"protolenids" and some species of Pararaia are homoplastic, and that both Hsuaspis (=Estaingia) and Pararaia should be assigned to the
Ichangiidae (=Estaingiidae).
Genus ESTAINGIA Pocock, 1964
Estaingia POCOCK, 1964, p. 462; ÖPIK, 1975b, p. K); PATERSON, 2005, p. 89.
Hsuaspis CHANG IN LU, CHANG, ZHU, QIAN, AND XIANG, 1965, p. 85; SUN IN ZHOU, LIU, MENG, AND SUN, 1977, p. 123; LI IN YIN AND LI,
1978, p. 427; ZHANG AND ZHU, 1979, p. 516; ZHU IN ZHANG, LU, ZHU, QIAN, LIN, ZHOU, ZHANG, AND YUAN, 1980, p. 244; LI IN ZHOU, LI,
AND QU, 1982, p. 227; ZHANG IN QIU, LU, ZHU, BI, LIN, ZHOU, ZHANG, QIAN, JU, HAN, AND WEI, 1983, p. 52; SUN, 1984, p. 347; JELL IN
BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 310; PALMER IN PALMER AND ROWELL, 1995, p. 16; NEDIN, 1995,
p. 36; JAGO, LIN, DAVIDSON, STEVENS, AND BENTLEY, 1997, p. 69.
Pseudichangia CHU AND ZHOU IN LU, CHANG, CHIEN, CHU, LIN, ZHOU, QIAN, ZHANG, AND YUAN, 1974, p. 93; ZHU IN ZHANG, LU, ZHU,
QIAN, LIN, ZHOU, ZHANG, AND YUAN, 1980, p. 239; SUN, 1984, p. 346.
Strenax ÖPIK, 1975b, p. 13.
Zhuxiella ZHANG AND ZHU IN ZHANG, LU, ZHU, QIAN, LIN, ZHOU, ZHANG, AND YUAN, 1980, p. 247; SUN, 1984, p. 348.
Type species.-Estaingia bilobata Pocock, 1964, Early Cambrian, Emu Bay Shale, Kangaroo Island, South Australia.
Discussion.-The synonymy of Estaingia, Hsuaspis, Pseudichangia, Strenax, and Zhuxiella has been discussed by Jell (in Bengtson et al., 1990, p.
310), Jago et al. (1997, p. 69-71), and Paterson (2005, p. 89).
ESTAINGIA OCCIPITOSPINA (Jell in Bengtson et al., 1990) new combination
Figure 10
Hsuaspis occipitospina JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 312, fig. 200.
Figured material.-Eight cranidia, SAMP40630-40637 (Fig. 10.1-10.8); one partial librigena, SAMP40638 (Fig. 10.10); two partial thoraces,
SAMP40639-40640 (Fig. 10.11, 10.14); one partial thoracic segment, SAMP40641 (Fig. 10.9); two pygidia, SAMP40642-40643 (Fig. 10.12,
10.13).
Occurrence.-Pararaia janeae Zone, Oraparinna Shale, OS section horizon: OS/76; 38 m (true thickness) above the base of the section (Fig. 4).
Discussion.-This species was originally described and referred to as Hsuaspis occipitospina by Jell (in Bengtson et al., 1990). However, regarding
the synonymy of Hsuaspis and Estaingia (see Jell in Jell and Adrain, 2003, p. 334), it should now be referred to as Estaingia occipitospina (Jell).
Specimens of Estaingia from the OS section undoubtedly belong to E. occipitospina based on Jell's (in Bengtson et al., 1990, p. 312) diagnosis and
description, although most cranidia illustrated herein (Fig. 10.1-10.6, 10.8) do not display the weakly impressed axial furrows otherwise
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characteristic of this species. This is likely due to preservation, since specimens from the OS section are preserved as exfoliated internal molds in
siltstone; this is further supported by the variation in the axial furrows between the internal mold and latex cast of the holotype of E.
occipitospina (Jell in Bengtson et al., 1990, fig. 200e, f). This would also explain the apparent absence of the occipital spine in many of the
specimens. Jago et al. (1997) and Paterson (2005) have also observed the same preservational bias of the occipital spine in specimens of Estaingia
cerastes (Öpik, 1975b) from the Cymbric Vale Formation, western New South Wales. Two cranidia from the OS section (Fig. 10.7, 10.8) also
demonstrate that the occipital spine in E. occipitospina is considerably longer and more vertically inclined than those illustrated by Jell (in
Bengtson et al., 1990, fig. 200f, o-r).
Genus PARARAIA Kobayashi, 1942
Pararaia KOBAYASHI, 1942, p. 492; JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEOAR, 1990, p. 302.
Proichangia ZHANG AND ZHU IN ZHANG, LU, ZHU, QIAN, LIN, ZHOU, ZHANG, AND YUAN, 1980, p. 241.
Type species.-Dolichometopus tatei Woodward, 1884, Early Cambrian, Parara Limestone, Horse Gully, Ardrossan, Yorke Peninsula, South
Australia.
PARARAIA BUNYEROOENSIS Jell in Bengtson et al., 1990
Figure 11
Pararaia bunyerooensis JELL IN BENGTSON, CONWAY MORRIS, COOPER, JELL, AND RUNNEGAR, 1990, p. 306, fig. 196.
Description.-Cranidium moderately small, up to 10 mm in length (sag.); subquadrate in outline: low convexity (sag., tr.). Anterior margin
strongly curved, width (tr.) equal to d-d; posterior margin (excluding occipital ring) straight, directed slightly posterolaterally abaxially to
fulcrum, then directed anterolaterally. Anterior sections of facial suture curved, diverging anteriorly at 50°-60° between ? and ß, then
converging between ß and a; posterior sections of facial suture very short. Glabella gently tapering, anterior width (tr.) at midlength of frontal
lobe 70%-80% occipital ring width (tr.); frontal lobe rounded: moderate convexity (tr.), low convexity (sag.); sagittal length (including LO)
70%-80% cranidial length. Axial furrow shallow, narrow (tr.), straight; preglabellar furrow shallow (exsag.), very shallow sagittally due to
plectrum, narrow (sag., exsag.). S1 weakly to moderately impressed, straight, directed anterolaterally abaxially (diverging anteriorly from
each other at 130°), width (tr.) approximately 25% adjacent glabellar width (tr.), narrow (exsag.); S2 weakly to moderately impressed, straight,
transverse, width (tr.) same as S1, narrow (exsag.); S3 obscure to moderately impressed, straight, transverse to directed posterolaterally
abaxially, width (tr.) same as S1, narrow (exsag.), Occipital ring of moderate convexity (tr.), flat-tened sagittally; length (sag.) 20%-25%
glabellar length; small posteromedial occipital node weakly developed; posterior margin strongly bowed. SO straight, narrow (sag., exsag.), of
moderate depth medially and becoming deeper abaxially. Preglabellar field moderately downsloping, weakly convex (sag.), length (sag.)
10%-20% cranidial length; plectrum well developed and of low relief, subtriangular in outline, expanding anteriorly, encroaching onto anterior
border. Preocular field moderately downsloping, weakly convex (exsag.), length (exsag.) approximately 25% cranidial length (sag.). Anterior
border flat to weakly convex, length (sag.) 10%-15% cranidial length, slightly expanding (exsag.) abaxially; anterior border furrow strongly
curved, more of a change in slope than distinct furrow, interrupted sagittally by plectrum. Palpebral lobe well developed, moderately convex
(tr.), length (exsag.) 30%-35% cranidial length (sag.), width (tr.) 25%-30% lobe length, anterior tip situated opposite S3, posterior tip situated
opposite SO or slightly anterior of SO; palpebral furrow shallow, narrow (tr.). Eye ridge well developed, forming a continuation of the palpebral
lobe, flat to weakly convex (exsag.), gently curved anteriorly, ridges diverge posteriorly at 140°, proximal end situated adjacent to posterior
portion of frontal lobe, separated from glabella by axial furrow; eye ridge divided into inner and outer bands by very weakly developed ocular
furrow. Palpebral area flat to weakly convex, width at e (tr.) 70%-85% adjacent glabellar width. Posterolateral projection of fixigena extremely
short (tr.), gently downsloping, broadly rounded distally. Posterior border moderately convex (exsag.), slightly expanding abaxially to fulcrum
that displays a rounded bulge, then sharply tapers to distal end of border; posterior border furrow deep, wide (exsag.), expanding abaxially.
Rostral plate strongly curved, following same curvature as anterior margin of cranidium, strongly convex dorsoventrally; uniform length (sag.,
exsag.), length approximately 15% width (tr.); rostral suture evenly curved; connective sutures straight, divergent anteriorly; posterior margin
smooth. Dorsal surface with shallow, very narrow (sag., exsag.) furrow developed around midlength (sag., exsag.), extending across entire width
(tr.). Ventral surface with ridge extending along midlength, representing change in slope.
Hypostome unknown.
Librigena moderately small, up to 14 mm in length (including genal spine); width (tr.) approximately 25%-30% length (including genal spine);
lateral margin gently curved, continuing evenly onto genal spine; posterior margin gently curved before strongly curving onto genal spine.
Genal field of low convexity and low relief, width (tr.) approximately 40%-50% adjacent librigenal width. Lateral border flat to weakly convex,
raised above genal field, widening (tr.) posteriorly, becoming widest at genal angle, anteriormost dorsal portion of border sharply tapers to a
point; epiborder furrow weakly developed near lateral margin and continues posteriorly onto entire length of genal spine, epiborder furrow also
developed on proximal margin of genal spine; lateral and posterior border furrows more of a change in slope than distinct furrows. Posterior
border flat to weakly convex, raised above genal field, short (tr.), narrow (exsag.), moderately expanding in width (exsag.) abaxially. Genal
spine long, length 50%-60% librigenal length (including spine) [at least 65% in smaller librigenae (Fig. 11.17, 11.18)]; broad base (tr.). strongly
tapering posteriorly, slight adaxial curvature. Librigenal doublure width (tr.) mimics dorsal surface of lateral and posterior librigenal border
and anterior cranidial border; sharp ridge developed on lateral portion of doublure terminating at base of genal spine (forming continuation of
10 of 19 8/12/2008 12:29 AM
ridge on ventral surface of rostral plate), representing u change in slope from rounded lateral margin to wide (tr.), slightly concave adaxial
portion.
Thoracic segments with axis of strong convexity. Axial ring width (tr.) approximately 80% width of pleura, slightly convex anteriorly with
lateral portions directed anterolaterally; small medial axial node weakly developed; articulating half ring of subequal length (sag.) to axial ring,
strongly tapering abaxially, slightly narrower (tr.) than axial ring; articulating furrow concave anteriorly, shallow medially, becoming deeper
abaxially. Pleura horizontal to fulcrum, then moderately downsloping to distal end; anterior pleural band rapidly expands abaxially with well
developed facet; posterior pleural band rapidly tapers abaxially; pleural furrow deep, extends diagonally across pleura from anterolateral corner
of axial ring to posterolateral corner of pleural tip.
Pygidium very small, up to 2.5 mm in length (sag.), strongly convex (sag., tr.), sagittal length 35%-45% maximum width (tr.). Axis gently
tapers posteriorly, almost reaching posterior margin, anterior width (tr.) 30%-40% maximum pygidial width; two axial rings and a broadly
rounded terminal piece, anterior and posterior axial rings separated by well developed pseudo-articulating half ring: axial ring furrow
separating posterior axial ring and terminal piece very shallow medially and deepened laterally; articulating half ring of uniform length (sag.,
exsag.), of subequal length to, and slightly narrower (tr.) than, anterior axial ring (Jell in Bengtson et al., 1990, fig. 196k). Pleural regions well
defined with three distinct pleurae, each terminating in short posteriorly directed marginal spines: pleurae separated by well developed
interpleural furrows; anterior pleura considerably wider (tr.) than posterior pleura; anterior band of anterior pleura rapidly expands abaxially,
posterior band of anterior pleura of uniform length (exsag.) and narrower (exsag.) than anterior band; anterior and posterior bands of posterior
pleura subequal in length (exsag.); anterior and posterior bands of pleurae separated by deep pleural furrows that terminate before marginal
spines. Third pair of marginal spines situated on posterior margin. Pygidial doublure narrow laterally, obscure medially.
Entire dorsal surface of exoskeleton smooth. Preglabellar and preocular fields covered in genal caeca.
Figured material.-Six cranidia, SAMP40644-40649 (Fig. 11.1, 11.2, 11.5-11.10); six librigenae, SAMP40650-40655 (Fig. 11.3, 11.4,
11.15-11.18, 11.22-11.24); one rostral plate, SAMP40656 (Fig. 11.11, 11.12); one fragmentary thoracic segment, SAMP40657 (Fig. 11.13,
11.14); two pygidia, SAMP40658-40659 (Fig. 11.19-11.21).
MMF/19, MMF/25.2, MMF/44.5, MMF/51.9, MMF/52.4, MMF/54.6, MMF/56.4, MMF/64.5, MMF/75.2, MMF/82.6, MMF/85.8, MMF/91.8,
MMF/93, MMF/103; 5.1-59.1 m (true thickness) above the base of the section (Fig. 4).
Discussion.-New silicified material of Pararaia bunyerooensis has enabled better understanding of the morphology of this species, especially
ventral structures, hence the revised description herein. Zang et al. (2001) tentatively assigned three fragmentary cranidia from the Yalkalpo-2
drillcore in the Arrowie Basin to Pararaia bunyerooensis. Unfortunately, the specimens of Pararaia? bunyerooensis from Yalkalpo-2 illustrated
by Zang et al. (2001, pl. 1, figs, g, h, k; reillustrated in Gravestock et al., 2001, pl. 1, figs. 7, 8, 11) are too incomplete, and the paucity of other
sclerites renders species identification impossible.
A possible paedomorphic lineage may exist between Pararaia bunyerooensis and P. jeineae based on new evidence from small specimens of P.
bunyerooensis from the MMF section. These species are considered to be sister taxa based on the following shared derived characters that are
lacking in other congeners: lateral border and genal spine of librigena with distinct epiborder furrow, and three pairs of pygidial marginal spines.
Cranidial characteristics in smaller specimens of P. bunyerooensis (e.g., Fig. 11.10) such as the longer (sag.) preglabellar field, wider (tr.)
palpebral area, and narrower (tr.) glabella are typical in large cranidia of P. janeae (Jell in Bengtson et al., 1990, fig. 197a, b, p, q). Smaller
librigenae of P. bunyerooensis (e.g., Fig. 11.17, 11.18) possess a considerably longer, narrow based (tr.) genal spine and narrower (tr.) lateral
border, characteristic of P. janeae librigenae (Jell in Bengtson et al., 1990, fig. 1971, m). Smaller pygidia of P. bunyerooensis (e.g., Fig. 11.19,
11.20) appear to have a similar sagittal length to transverse width ratio as pygidia of P. janeae (Jell in Bengtson et al., 1990, fig. 197f, g).
Furthermore, there is a considerable difference in the known maximum size ranges of these species, i.e., up to 10 mm cranidial length (sag.) in P.
bunyerooensis and up to 15 mm cranidial length (sag.) in P. janeae (Jell in Bengtson et al., 1990, fig. 197q). This indicates that retardation in
the onset of maturity in P. janeae resulted in the attainment of a larger adult size than in P. bunyerooensis. The above evidence suggests that P.
janeae evolved from the ancestral P. bunyerooensis via neoteny (s. McNamara, 1986).
Family YUNNANOCEPHALIDAE Hupé, 1953
Genus YUNNANOCEPHALUS Kobayashi, 1936
Yunnanocephalus KOBAYASHI, 1936, p. 101; HENNINGSMOEN IN R. C. MOORE, 1959, p. 212; LI, 1978, p. 196; ZHANG, 1987, p. 228 (for
additional synonymy); LUO IN LUO, JIANG, AND TANG, 1994, p. 137; SHU, GEYER, CHEN, AND ZHANG, 1995, p. 222 (for comprehensive
synonymy); PALMER IN PALMER AND ROWELL, 1995, p. 17.
Type species.-Ptychoparia yunnanensis Mansuy, 1912, Early Cambrian, Chiungchussu Formation, Yunnan, China.
YUNNANOCEPHALUS MACROMELOS new species
Figure 12
Diagnosis.-See Table 1 for summary of diagnostic characters.
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Description.-Cranidium moderately small, up to 8 mm in length (sag.); trapezoidal in outline; strongly convex (sag., tr.). Anterior margin
moderately curved, width (tr.) 85%-95% e-e; posterior margin (excluding occipital ring) straight and transverse to fulcrum, then directed
posterolaterally. Anterior sections of facial suture short, subparallel to slightly divergent anteriorly between ? and ß, then strongly convergent
anteriorly between ß and a; posterior sections of facial suture long, curved anteriorly, widely divergent posteriorly. Glabella gently tapering,
anterior width (tr.) at midlength of frontal lobe 70%-75% occipital ring width (tr.); frontal lobe truncate; strong convexity (sag., tr.); sagittal
length (including LO) 80%-85% cranidial length. Axial furrow deep, wide (tr.), straight; preglabellar furrow more of a change in slope than
distinct furrow. Glabellar furrows well developed; S1 moderately impressed, convex anteriorly, directed anterolaterally abaxially, width (tr.)
approximately 30%-35% adjacent glabellar width (tr.), narrow (exsag.); S2 moderately impressed, straight, transverse, width (tr.) same as S1,
narrow (exsag.); S3 weakly impressed, straight, directed posteroluterally abaxially, width (tr.) approximately 25%-30% adjacent glabellar
width (tr), narrow (exsag.). Occipital ring of strong convexity (tr.), flattened sagittally; length (sag.) 15%-20% glabellar length; small
posteromedial occipital node weakly developed; posterior margin weakly convex posteriorly. SO transverse medially, directed anteriorly
laterally, wide (sag., exsag.), of moderate depth medially and becoming deeper abaxially. Preglabellar field absent. Preocular field subtriangular
in outline, strongly downsloping, weakly convex (exsag.), maximum length (exsag.) 10%-15% cranidial length (sag.). Anterior border strongly
convex, length (sag.) 15%-20% cranidial length, slightly expands (exsag.) abaxially to midway between sagittal line and lateral margin, then
tapers abaxially; anterior border furrow of moderate depth, lateral sections weakly convex anteriorly, medial section in front of glabella strongly
convex anteriorly. Palpebral lobe well developed, moderately convex (tr.), length (exsag.) 25%-30% cranidial length (sag.), width (tr.) 30%-35%
lobe length, anterior tip situated opposite L3, posterior tip situated opposite midlength of L1; palpebral furrow shallow, narrow (tr.). Eye ridge
well developed, forming a continuation of the palpebral lobe, moderately convex (exsag.), gently curved anteriorly, ridges diverge posteriorly at
120°-130°, proximal end situated adjacent to frontal lobe, separated from glabella by axial furrow, although anteroproximal edge of eye ridge
merges onto anterior margin of frontal lobe. Palpebral area moderately convex, width (tr.) at e approximately 50% adjacent glabellar width.
Postocular area short (exsag.), of subequal length (exsag.) to sagittal length of occipital ring. Posterolateral projection of fixigena very wide,
width (tr.) approximately 25% e-e, strongly downsloping, strongly tapers abaxially. Posterior border strongly convex (exsag.), slightly
expanding abaxially to fulcrum, then rapidly expands to distal end of border; small posterolaterally directed intergenal spine developed at distal
extremity of posterior border; posterior border furrow deep, expanding abaxially, connecting with axial furrow at the posteroproximal portion of
fixigena forming an L shape.
Rostral plate moderately curved, following same curvature as anterior margin of cranidium, strongly convex dorsoventrally; uniform length
(sag., exsag.), length approximately 25% width (tr.); rostral suture evenly curved; connective sutures concave abaxially; posterior margin
smooth. Ventral surface with ridge extending along midlength, representing change in slope.
Hypostome unknown.
Librigena small, up to 7 mm in length; width (tr.) approximately 40% length; lateral margin strongly curved; lateral margin and posterior
section of facial suture converge at genal angle. Genal field of low convexity, width (tr.) 55%-65% librigenal width. Lateral border strongly
convex, slightly tapers (tr.) posteriorly, anteriormost dorsal portion of border sharply tapers to a point; lateral border furrow shallow, narrow
(tr.). Posterior border and posterior border furrow absent. Genal spine absent. Librigenal doublure width (tr.) mimics dorsal surface of lateral
librigenal border and anterior cranidial border; sharp ridge developed on lateral margin of doublure (forming continuation of ridge on ventral
surface of rostral plate), representing a change in slope from rounded lateral margin to wide (tr.), slightly concave adaxial portion.
Thoracic segments with axis of strong convexity. Axial ring of uniform width (sag., exsag.), posterior margin slightly convex anteriorly; small
posteromedial axial node well developed; articulating half ring of subequal length (sag.) to axial ring, tapering abaxially; articulating furrow
straight, wide (sag., exsag.), equal depth across entire width. Pleura horizontal to fulcrum, then strongly downsloping to distal end; anterior
pleural band expands abaxially, anterior pleural margin with projection at anterolateral corner; posterior pleural band expands abaxially;
pleural furrow deep, terminating adjacent to projection at anterolateral corner of anterior pleural margin. Doublure of thoracic pleura developed
at pleural tip and tapers adaxially along posterior margin.
Pygidium unknown.
Entire dorsal surface of exoskeleton densely covered in tiny granules.
Morphogenesis.-Small cranidia of Yunnanocephalus macromelos (e.g., Fig. 12.15) show that there is considerable morphological variation during
ontogeny and display the following characteristics: anterior sections of facial suture slightly converge anteriorly; narrower (tr.) anterior
margin; glabella parallel-sided with rounded frontal lobe; S1 transglubellur, convex posteriorly; longer (exsag.) palpebral lobe; eye ridges more
widely divergent posteriorly, proximal ends continue around frontal lobe to form prominent parafrontal band.
Etymology.-Greek makros, long, and melon, limb; referring to the wide (tr.) posterolateral projections of the fixigenae.
Type material.-Holotype: cranidium, SAMP40660 (Fig. 12.1-12.5). Paratypes: four cranidia, SAMP40661-40664 (Fig. 12.12-12.15, 12.26);
three librigenae, SAMP40665-40667 (Fig. 12.6-12.11); one articulated librigena and rostral plate, SAMP40668 (Fig. 12.25); one rostral plate,
SAMP40669 (Fig. 12.16, 12.17); one thoracic axial ring, SAMP40670 (Fig. 12.20, 12.21); two thoracic pleurae, SAMP40671-40672 (Fig.
12.18, 12.19, 12.22-12.24).
Type locality.-Mernmerna Formation (MMF section), Hawker Group; base of section coordinates: 31°11°38.4''S, 138°52'28.7''E; map datum:
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WGS84, on the eastern side of The Bunkers, approximately 1 km south of Balcoracana Creek on Angorichina Station. Flinders Ranges, South
Australia (see Figs. 2, 3).
Type stratum.-Third Plain Creek Member of the Mernmerna Formation, MMF section, horizon MMF/64.5; 37 m (true thickness) above the base of
the section (Fig. 4).
Occurrence.-Pararaia bunyerooensis Zone, Third Plain Creek Member, Mernmerna Formation, MMF section horizons: MMF/ 44.5, MMF752.4,
MMF/58.3, MMF/64.5, MMF/68.7, MMF/75.2, MMF/81.5, MMF/82.6, MMF/103: 25.5-59.1 m (true thickness) above the base of the section
(Fig. 4).
Discussion.-There are currently three valid species of Yunnanocephalus: the type species Y. yunnanensis (Mansuy, 1912) from China; Y.
longioccipitalis Palmer (in Palmer and Rowell, 1995) from Antarctica; and Y. macromelos described herein. Several other species have also been
described from China, but they are considered to be junior subjective synonyms of Y. yunnanensis (see Zhang, 1987, p. 229 and Shu et al., 1995,
p. 222 for comprehensive synonymy lists). Comparison of diagnostic cranidial characters between species of Yunnsnocephalus are outlined in
Table 1. The librigenae of Y. macromelos (Fig. 12.6-12.11) and Y. longioccipitalis (Palmer in Palmer and Rowell, 1995, fig. 14.8, 14.9) are
indistinguishable, but can be differentiated from the librigena of K yunnanensis (e.g., Zhang, 1987, pl. 1, figs. 1, 2, 4; Shu et al., 1995, figs. 17a,
19a-e) in displaying a wider (tr.) genal field.
Genus ELICICOLA Jell in Bengtson et al., 1990
Type species.-Elicicola calva Jell in Bengtson et al., 1990, Early Cambrian, Parara Limestone, Kulpara, South Australia.
ELICICOLA CALVA Jell in Bengtson et al., 1990
Figure 13
Elicicola calva JELL IN BENGTSON, CONWAY MORRIS, COOPRR, JELL, AND RUNNRGAR, 1990, p. 300, figs. 192j-m, 193.
Figured material.-Six cranidia, SAMP40673-40678 (Fig. 13.1-13.8).
Occurrence.-Abadiella huoi Zone, Wilkawillina Limestone, MMF section horizon: MMF/0.0 (Fig. 4).
Discussion.-This monotypic species has been well described and illustrated by Jell (in Bengtson et al., 1990) from the Parara Limestone at
Kulpara (type locality) and the Wilkawillina Limestone at Wirrealpa Mine and west of Wirrealpa Springs in the Flinders Ranges, and requires no
further discussion herein.
ACKNOWLEDGMENTS
Sincere thanks to E. Alexander (PIRSA) for assistance and guidance in the field in September 2002 and for her continual support of our research
on the Cambrian faunas of South Australia. T. Wright (PIRSA) also provided technical and logistical support during the September 2002 field
season. We are indebted to MUCEP members P. Bell, H. Caldon, P. Cockle, B. Jonak, R. Morgan, B. Pyemont, D. Smith, L. Strotz, and N. Wilson for
their assistance during the 2003 and 2004 field seasons. L. Strotz and P. Bell assisted with acid processing and picking of residues. Thanks also to
the Fargher family (owners of Angorichina Station) for access to the field area. Discussions with J. Jago, J. Laurie, and P. Jell on various aspects
of this paper were extremely beneficial. G. Edgecombe kindly read and commented on an early draft of the manuscript. N. Hughes and A.
Rushton provided helpful reviews. D. Oliver drafted Figures 1-4. Funding for this study came from a Macquarie University Postgraduate
Research Fund grant to JRP and a Macquarie University Research Development Grant to GAB.
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Early Cambrian Trilobites From Angorichina, Flinders Ranges, South Australia, With A New Assemblage From The Pararaia Bunyerooensis Zone

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Early Cambrian Trilobites From Angorichina, Flinders Ranges, South Australia, With A New Assemblage From The Pararaia Bunyerooensis Zone

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Journal of Paleontology: EARLY CAMBRIAN TRILOBITES FRO... http://findarticles.com/p/articles/mi_qa3790/is_200701/ai_n171...

LOCALITY AND STRATIORAPHY

BIOSTRATIGRAPHY AND CORRELATION

Order REDLICHIIDA Richter, 1932

Suborder REDLICHIINA Richter, 1932

Superfamily REDLICHIOIDEA Poulsen, 1927

Family REDLICHIIDAE Poulsen, 1927

Genus REDLICHIA Cossmann, 1902

REDLICHIA GUIZHOUENSIS Zhou in Lu et al., 1974

Olenellus sp. ETHERIDGE, 1905, p. 247, pl. 25, fig. 1.

Olenellus? sp. ETHERIDGE, 1919, p. 382, pl. 39. fig. 1.

Genus EOREDLICHIA Chang in Lu and Dong, 1952

Genus WUTINGASPIS Kobayashi, 1944

WUTINGASPIS EURYOPTILOS new species

Figures 8.8-8.17, 9.1-9.7

Dolerolenus? sp. nov. ÖPIK, 1975B, p. 41, pl. 7, tig. 2.

Rostral plate and hypostome unknown.

Genus YORKELLA Kobayashi, 1942

YORKELLA aff. AUSTRALIS (Woodward, 1884)

Superfamily ELLIPSOCEPHALOIDEA Matthew, 1887

Family ESTAINGIIDAE Öpik, 1975a

Genus ESTAINGIA Pocock, 1964

Strenax ÖPIK, 1975b, p. 13.

ESTAINGIA OCCIPITOSPINA (Jell in Bengtson et al., 1990) new combination

Genus PARARAIA Kobayashi, 1942

PARARAIA BUNYEROOENSIS Jell in Bengtson et al., 1990

Family YUNNANOCEPHALIDAE Hupé, 1953

Genus YUNNANOCEPHALUS Kobayashi, 1936

YUNNANOCEPHALUS MACROMELOS new species

Diagnosis.-See Table 1 for summary of diagnostic characters.

Entire dorsal surface of exoskeleton densely covered in tiny granules.

Genus ELICICOLA Jell in Bengtson et al., 1990

ELICICOLA CALVA Jell in Bengtson et al., 1990

Figured material.-Six cranidia, SAMP40673-40678 (Fig. 13.1-13.8).

COSSMANN, M. 1902. Rectifications de la nomenclature. Revue Critique de Paléozoologie, 16:52.

GEYER, G. 1990. Die marokkanischen Ellipsocephalidae (Trilobita: Redlichiida). Beringeria, 3:3-363.

South Australia, 44:382.

McNAMARA, K. J. 1986. A guide to the nomenclature of heterochrony. Journal of Paleontology, 60:4-13.

ACCEPTED 29 JULY 2005

JOHN R. PATERSON and GLENN A. BROCK

Copyright Paleontological Society Jan 2007

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