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Paterson JR and Jago, JB (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.

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Further information regarding the copyright applicable to this article can be obtained from Association of Australasian Palaeontologists, http://www.gsa.org.au/specialgroups/aap.html. New trilobites from the Lower Cambrian Emu Bay Shale Lagerstätte at Big Gully, Kangaroo Island, South Australia

JOHN R. PATERSON & JAMES B. JAGO

PATERSON, J.R. & JAGO, J.B., 2006:07:29. 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. ISSN 0810-8889.

Two new trilobites from the Lower Cambrian Emu Bay Shale Lagerstätte at Big Gully, Kangaroo Island, are described. Megapharanaspis nedini gen. et sp. nov. is placed in the Megapharanaspidae, a new family of the Emuelloidea. Holyoakia simpsoni sp. nov. represents the first occurrence of the genus outside Antarctica, providing further confirmation of the close faunal affinity between Cambrian trilobite faunas from Australia and East Antarctica. New specimens of Redlichia takooensis are documented that provide new morphological information, including a description of the rostral plate and hypostome.

J.R. Paterson ([email protected]), South Australian Museum, Division of Natural Science, North Terrace, Adelaide, SA 5000 and Department of Earth & Planetary Sciences, Macquarie University, NSW 2109, Australia; J.B. Jago ([email protected]), School of Natural and Built Environments, University of South Australia, Mawson Lakes, SA 5095, Australia. Received 21 March 2006.

Keywords: Early Cambrian, Australia, East Gondwana, Trilobita, Redlichia, Megapharanaspis, Holyoakia.

TRILOBITES, brachiopods and hyoliths were the these stratigraphic horizons, the known biota ﬁrst reported fossils from the Emu Bay Shale type is of relatively low diversity compared to other section in Emu Bay (Sprigg et al. 1954; Sprigg Cambrian Lagerstätten such as the Chengjiang 1955; Daily 1956) thus establishing an Early and Burgess Shale faunas (Briggs et al. 1994; Cambrian age. Numerous studies on the Emu Bay Hou et al. 2004). We describe two new trilobites, Shale fauna (Pocock 1964, 1970; Glaessner 1979; Megapharanaspis nedini gen. et sp. nov. and Conway Morris & Jenkins 1985; Conway Morris Holyoakia simpsoni sp. nov., and provide new 1989; Bengtson et al. 1990; McHenry & Yates information about Redlichia takooensis from the 1993; Nedin 1995a, 1997, 1999; Briggs & Nedin Emu Bay Shale at Big Gully. 1997; Nedin & Jenkins 1999; Dzik 2004; Paterson & Edgecombe 2006) have recorded the trilobites STRATIGRAPHIC SETTING AND AGE Redlichia takooensis Lu, 1950, Estaingia bilobata Big Gully is situated to the east of Emu Bay and Pocock, 1964, Emuella dalgarnoi Pocock, 1970 approximately 1.5 km west of White Point on the and Balcoracania dailyi Pocock, 1970, the northern coast of Kangaroo Island (Fig. 1). The arachnomorphs Naraoia sp. and Xandarella Emu Bay Shale crops out to the east of the mouth sp., the bivalved arthropods Isoxys communis of Big Gully and is approximately 78 m thick Glaessner, 1979 and Tuzoia australis Glaessner, (Daily et al. 1979). Here the unit unconformably 1979, the anomalocaridids Anomalocaris overlies the White Point Conglomerate and briggsi Nedin, 1995a and Anomalocaris sp., the conformably underlies the Boxing Bay Formation palaeoscolecid priapulan Palaeoscolex antiquus (Daily et al. 1979; Briggs & Nedin 1997). Most Glaessner, 1979, the problematic forms Myoscolex fossils occur in the dark grey silty shales within ateles Glaessner, 1979, Vetustovermis planus the basal 8 m of the unit (Briggs & Nedin 1997), Glaessner, 1979 and Wiwaxia sp., in addition to although Daily et al. (1979) recorded fossils up to undescribed phosphatic brachiopods, hyoliths, 23.5 m above the base of the formation. chancelloriids and complete sponges. Fossils Trilobites are by far the most common occur at several stratigraphic horizons throughout constituent of the Emu Bay Shale Lagerstätte at the unit at Emu Bay and Big Gully (Fig. 1); Big Gully, representing approximately 60% of however, the Lagerstätte is only preserved at the the total assemblage (Nedin 1995b). The majority latter locality (Nedin 1995a, b, 1997; Briggs & of specimens are partially or fully articulated, Nedin 1997). Although fossils are abundant in but a high proportion represent moults, which 44 AAP Memoir 32 (2006)

Fig. 1. Location of Big Gully on the northern coast of Kangaroo Island, South Australia and the Cambrian geology along the coast between Cape D’Estaing and Point Marsden; modified from Nedin (1995b). may exaggerate the apparent dominance of also supports correlation with the Tsanglangpuan trilobites in the fauna (Briggs & Nedin 1997). The Stage, based on the occurrence of this species in most abundant taxa at Big Gully are Redlichia China (see Zhang et al. 1980) and in the Lolab takooensis and Estaingia bilobata (Nedin 1995b) Formation in Kashmir (Jell & Hughes 1997; of which several hundred specimens are available Hughes & Jell 1999). in the South Australian Museum collections. The only previously described species of Rarer species include Balcoracania dailyi and Holyoakia is the type species, H. granulosa, from the new taxa described herein, Megapharanaspis a small fauna within the Shackleton Limestone of nedini gen. et sp. nov. and Holyoakia simpsoni the Central Transantarctic Mountains. It occurs sp. nov., each of which is known from only five in association with the trilobites described in specimens. Palmer & Rowell (1995) as Calodiscidae gen. et The occurrence of Estaingia bilobata and sp. undetermined A, Berabichia cf. B. subditus Balcoracania dailyi allows correlation with the (Palmer & Gatehouse, 1972) and Pensacola? sp. Pararaia janeae Zone of South Australia (Jell 2. The fauna was assigned a late Atdabanian age in Bengtson et al. 1990; Paterson & Edgecombe on the basis of Berabichia being closely related to 2006; Paterson & Brock, in press). Paterson Chorbusulina from the late Atdabanian of Siberia & Brock (in press, fig. 5) have correlated this (Palmer & Rowell 1995). If this age determination zone with the early-mid Tsanglangpuan Stage of is indeed correct, then Holyoakia simpsoni sp. China and the mid-late Botoman of Siberia. The nov. is younger than H. granulosa. However, the occurrence of Redlichia takooensis at Big Gully paucity of age diagnostic taxa from the Shackleton AAP Memoir 32 (2006) 45

Limestone assemblage and its weak correlation 1997 Redlichia takooensis Lu; Jell & Hughes, with Siberia based on generic affinities casts doubt p. 22, pl. 2, figs 4-12 (see for additional on a late Atdabanian age. Other occurrences of synonymy). Berabichia appear to indicate a Botoman age. For 1999 Redlichia takooensis Lu; Nedin, p. 988, example, it occurs in the uppermost Banian Stage fig. 2B. of Morocco (Geyer 1990a, b, 2005), equivalent 2005 Redlichia takooensis Lu; Vannier & Chen, to the Botoman Stage of Siberia (Geyer & p. 14, fig. 9. Shergold 2000; Geyer 2005); while its occurrence in Avalonia indicates a Botoman (or perhaps Material. Over 100 internal and external moulds younger) age (Westrop & Landing 2000). of complete specimens, isolated cranidia and The occurrence of Holyoakia in South librigenae are available as well as several Australia provides further confirmation of the separate pygidia and hypostomes; registered close faunal affinity between Cambrian trilobite and unregistered specimens are housed in faunas from Australia and Antarctica (Brock the palaeontological collection of the South et al. 2000; Lieberman 2003, 2004; Meert & Australian Museum. There are two specimens Lieberman 2004). In addition to Holyoakia, in which the hypostome is associated with the there are five other Early Cambrian congeneric rostral plate. Figured specimens: SAMP35566, occurrences between Australia and East Antarctica, 40181, 41173-41182. including Balcoracania, Estaingia, Pagetides (Discomesites), Redlichia and Yunnanocephalus Remarks. This species has been well documented (Palmer & Gatehouse 1972; Bengtson et al. 1990; (see synonymy). However, new specimens from Palmer & Rowell 1995; Paterson 2005; Paterson the Emu Bay Shale at Big Gully, (illustrated & Brock, in press). Paterson (2005) demonstrated herein and others housed in the South Australian that the respective species of Estaingia and Museum), warrant further discussion. In light Pagetides (Discomesites) from the Cymbric Vale of this new material, Jell’s (in Bengtson et al. Formation in western New South Wales, Australia 1990) description of the Big Gully population and the Shackleton Limestone in the Central of Redlichia takooensis requires minor revision. Transantarctic Mountains are actually conspecific, Firstly, Jell noted that the tenth thoracic segment and suggested that a continuous carbonate-detrital possesses a long, slender medial spine, but the shelf along the East Gondwanan margin allowed new specimens clearly indicate its position on faunal exchange between the two regions. the eleventh segment. This was, in fact, suggested by Jell (p. 273) in his discussion of the specimen SYSTEMATIC PALAEONTOLOGY illustrated by Zhang et al. (1980, pl. 23, fig. 13). The specimens described herein occur as internal Moreover, new material reveals that the thoracic and external moulds in weathered siltstone. Latex axial nodes that are apparent in many specimens casts were prepared from the external moulds (e.g., Jell in Bengtson et al. 1990, fig. 181B-D; prior to photography. Terminology follows Conway Morris & Jenkins 1985, figs 1B, 2A, B; Whittington & Kelly (1997). Specimen numbers Figs 2, 3A, D, 4A) appear to represent the bases refer to the palaeontological collection of the of axial spines (Fig. 3C, D). Secondly, Jell’s South Australian Museum, Adelaide (SAMP). description of the pygidium indicates the presence of only one axial ring and a bilobed terminal piece, Order REDLICHIIDA Richter, 1932 but well preserved material illustrated herein (Figs Suborder REDLICHIINA Richter, 1932 2, 4A, E, G) shows two axial rings. Superfamily REDLICHIOIDEA Poulsen, 1927 New specimens of Redlichia takooensis from Family REDLICHIIDAE Poulsen, 1927 Big Gully also reveal considerable ontogenetic variation in the librigenae, in particular, the width Redlichia Cossmann, 1902 (tr.) of the posterior margin, pitch of the intergenal angle (sensu Chang 1966, fig. 1) and length Type species. Hoeferia noetlingi Redlich, 1899. (exsag.) of the genal spine. In general, librigenae of smaller individuals have a wider (tr.) posterior Redlichia takooensis Lu, 1950 (Figs 2-4) margin, distinct intergenal angle (115-135°) – creating a more advanced (anterior) position of the 1979 Redlichia sp.; Daily et al., p. 14, fig. 15. genal angle – and longer genal spines (e.g., Fig. 1990 Redlichia takooensis Lu; Jell in Bengtson 4B). Librigenae of larger individuals (exoskeletal et al., p. 271, fig. 181 (see for additional length >12 cm) display a narrower (tr.) posterior synonymy). margin, more obtuse intergenal angle (150-180°) 1993 Redlichia takooensis Lu; McHenry & Yates, and shorter genal spines (Fig. 2). p. 81, figs 10-12. The hypostomes of most specimens are crushed 46 AAP Memoir 32 (2006)

Fig. 2. Redlichia takooensis Lu, 1950. Internal mould of complete exoskeleton (SAMP40181), x1.5. AAP Memoir 32 (2006) 47

Fig. 3. Redlichia takooensis Lu, 1950. A, latex cast of external mould of almost complete exoskeleton, SAMP41173, x1.5. B, internal mould of partial thorax with very long axial spine, SAMP35566, x2. C, oblique view of internal mould of partial thorax and pygidium, arrow indicating posteriorly directed thoracic axial spine, SAMP41180, x1.5. D, latex cast of external mould of pygidium and posterior part of thorax, SAMP41177, x1. beneath the glabella (Jell in Bengtson et al. 1990, middle body; the maculae are present as slits. p. 271); Jell had no hypostome suitable for The posterior border furrow is poorly deﬁned description. We have available two hypostomes in the smaller specimens but well deﬁned in that appear to be attached to the rostral plate (Fig. the large specimen. The posterior border in the 4C, D) and a much larger partial hypostome, the large specimen is wide and gently convex with anterior part of which is missing (Fig. 4F). The two pairs of posterolateral spines; the anterior length of the hypostome is about 45% that of the pair of spines are situated opposite the maculae cranidium; it has a moderately convex, ovate and project ventrally, the posterior pair of spines 48 AAP Memoir 32 (2006)

Fig. 4. Redlichia takooensis Lu, 1950. A, internal mould of complete exoskeleton, SAMP41174, x1.5. B, internal mould of small complete exoskeleton, SAMP41176, x3. C, internal mould of rostral plate and hypostome, SAMP41181, x3. D, internal mould of rostral plate and hypostome in place beneath cephalon, SAMP41175, x2. E, latex cast of external mould of pygidium, SAMP41179, x1.5. F, latex cast of external mould of hypostome, SAMP41182, x2. G, internal mould of partial pygidium and posterior of thorax, SAMP41178, x1.5. AAP Memoir 32 (2006) 49 at posterolateral extremities of the border and hypostomal suture in R. takooensis. separated by a gently convex posterior margin. Guo & Zhao (1998) erected the subspecies In the larger specimen, the posterior lobe of the Redlichia takooensis longispina. Guo et al. middle body is much more clearly defined; it is (1999, p. 163) differentiated the subspecies from separated from the anterior lobe by a shallow, R. takooensis on the basis that longispina has but distinct furrow. Entire surface covered in fine no occipital spine, but a very long librigenal reticulate ornament; posterior border with well spine. However, the specimens illustrated by developed terrace ridges. Guo et al. (1999) and reillustrated by Yuan et al. The hypostomes of Redlichia takooensis (Fig. (2002) are quite small and it is difficult to make 4C, D, F) are less elongate than those of R. idonea a meaningful comparison. The smallest specimen (Öpik 1958, pl. 4, fig. 2, pl. 5, fig. 1, pl. 6, figs 4, of R. takooensis from Big Gully illustrated herein 5), Redlichia sp. of Whittington (1988, text-fig. (SAMP41176; Fig. 4B) is about the same size as 6), R. nobilis (Zhang & Jell 1987, pl. 7, fig. 8) and the largest illustrated specimen of R. t. longispina R. forresti (Kruse et al. 2004, fig. 20R, T-V, X, (see Guo et al. 1999, pl. 1, fig. 1a, b). In specimen Y). The posterior border of the hypostome in both SAMP41176 (Fig. 4B), the genal spine extends R. forresti and R. idonea is more clearly defined to about the sixth thoracic segment, whereas in than that of R. takooensis. The rostral plate of R. R. t. longispina the genal spine extends much takooensis has well developed terrace lines; it has further to the posterior, possibly to about level a similar shape to that of R. nobilis. The rostral with the tenth thoracic segment in the specimen plates of Redlichia sp. of Whittington (1988), illustrated by Guo et al. (1999, pl. 1, fig. 2a, b). R. idonea (Öpik 1958, pl. 5, fig. 1, pl. 6, figs 4, In R. takooensis from Big Gully, the longest genal 5), R. longtangensis (see Zhang et al. 1980, pl. spines extend to the level of the eighth thoracic 20, fig. 9; Fortey 1990, text-fig. 6), R. mansuyi segment (Figs 3A, 4A). (Luo et al. 1994, pl. 8, fig. 5) and R. murakamii (Luo et al. 1994, pl. 11, fig. 5) have a row of pits; Superfamily EMUELLOIDEA Pocock, 1970 such pits appear to be absent in the available Family MEGAPHARANASPIDAE fam. nov. specimens of R. takooensis. The rostral plates of R. longtangensis, R. mansuyi, R. murakamii and Diagnosis. Cranidium subquadrate; glabella Redlichia sp. of Whittington (1988) also differ gently tapered anteriorly with three transglabellar from that of R. takooensis in having a central furrows; preglabellar field very short (sag.) with posterior extension towards the anterior of the weakly developed plectrum; palpebral lobe hypostome. long (exsag.), crescentic; eye ridge short (tr.); Fortey (1990) noted that the nature of the posterolateral projection of fixigena extremely attachment of the rostral plate to the hypostome short (exsag., tr.). Thorax of 18 segments, with in Redlichia is conterminant. It is uncertain as prothorax of three segments and opisthothorax to whether Redlichia possessed a hypostomal of 15 segments; third prothoracic segment suture. Öpik (1958, p. 28) suggested that the macropleural. Pygidium very small, subelliptical; hypostome and rostral plate in R. idonea formed posterior margin smooth; axis strongly tapered, a ‘fused unit’. However, Whittington (1988) and with two rings and a terminal piece; pleural region Fortey (1990) noted that while there are some effaced; border absent. illustrated examples of Redlichia with rostral plate and hypostome attached, others are preserved Remarks. The diagnostic characters of as isolated sclerites, hence suggesting that the Megapharanaspis gen. nov. are similar to the latter could be the result of breakage or that a genera of the Emuellidae in many respects, but hypostomal suture was present. An alternative differ sufficiently as to warrant erection of a suggestion is that perhaps the hypostomal suture new family, the Megapharanaspidae, within the in Redlichia is species-specific. Nedin (1995b, Superfamily Emuelloidea; discussed below under pl. 6, figs 1, 3) illustrated a poorly preserved the remarks for Megapharanaspis. Therefore, the hypostome, with no attached rostral plate, of R. diagnosis of the Superfamily Emuelloidea by takooensis from Big Gully. On the basis of this Zhang (1997) should now become the familial single specimen he concluded that R. takooensis diagnosis for the Emuellidae. had a hypostomal suture. Most hypostomes of R. takooensis from Big Gully appear to be in position Megapharanaspis gen. nov. beneath the glabella, and in at least two specimens the hypostome appears to be attached to the rostral Type species. Megapharanaspis nedini gen. et plate (Fig. 4C, D). Unfortunately, the preservation sp. nov. of these specimens, which appear to be crushed from compaction, precludes confirmation of a Etymology. Greek mega = big or large, pharangos 50 AAP Memoir 32 (2006)

= gully, and aspis = shield; in reference to its differences in the thorax of Megapharanaspis occurrence at Big Gully. include the position of the macropleural segment (the third segment in Megapharanaspis versus Diagnosis. Small trilobite. Cranidium subquadrate; the sixth in emuellids) and the number of glabella moderately convex, gently tapered thoracic segments (three prothoracic and 15 anteriorly, sagittal length 80-85% cranidial length. opisthothoracic in Megapharanaspis versus Preglabellar field very short (sag.); plectrum six prothoracic and up to 97 opisthothoracic weakly developed. Anterior sections of facial in emuellids) (Paterson & Edgecombe 2006). suture diverging anteriorly at 50-55° between γ The pygidium of Megapharanaspis is similar and β, then weakly converging between β and in size and shape to that of Emuella dalgarnoi α; posterior sections of facial suture very short (Paterson & Edgecombe 2006, fig. 4.3), and and strongly diverging posteriorly. S1, S2 and both have an effaced pleural region and smooth S3 transglabellar. Palpebral lobe well developed, posterior margin. However, Megapharanaspis crescentic, extends from SO to S3. Eye ridge well has clearly defined axial furrows and only two developed. Posterolateral projection of fixigena axial rings; Emuella dalgarnoi has poorly defined extremely short (exsag., tr.). Librigena with almost axial furrows and four axial rings (Paterson & flat genal field; genal spine long, with distal tip Edgecombe 2006). Furthermore, the width (tr.) of opposite fourth or fifth thoracic segment. Thorax the posteriormost thoracic segments in emuellids of 18 segments; prothorax of three segments, appears to be comparable to the lateral margins of opisthothorax of 15 segments; prothoracic the pygidium (Paterson & Edgecombe 2006, figs segments longer (sag., exsag.) and wider (tr.) 3.4, 4.3, 7.4), whereas those in Megapharanaspis than opisthothoracic segments. Width (tr.) of axis are considerably wider (tr.) than the lateral approximately 50% width of segment; axial rings margins of the pygidium (Fig. 5A-D). with small medial nodes. T3 macropleural, pleura Megapharanaspis does not appear to be closely strongly directed posterolaterally, spine extending related to any member of the Redlichioidea to level of pygidium. Pygidium very small, (sensu Zhang 1997), though it does have subelliptical, posterior margin smooth. Axis superficial similarities to Lemdadella (see Sdzuy strongly tapered, two axial rings and a terminal 1978; Liñán & Sdzuy 1978; Liñán Guijarro piece; pleural region effaced; border absent. 1978; Palmer & Rowell 1995; Liñán et al. 2005). Cranidial similarities include: gently Remarks. Megapharanaspis gen. nov. appears to tapered glabella, variably developed plectrum, have affinities with members of the Emuellidae. length (sag., exsag.) of the anterior cranidial The cephalon and thorax of this genus are border, long (exsag.) crescentic palpebral lobes, characteristic of the Redlichiina (sensu Zhang width (tr.) of the palpebral area, and short (tr.) 1997), but the pygidium is quite different from posterolateral projections of the fixigenae. The any other member of the group. thorax of Lemdadella (Liñán & Sdzuy 1978, pl. The similarities between Megapharanaspis and 1, fig. 1; Liñán Guijarro 1978, pl. 3, figs 3-8, pl. the Emuellidae are quite striking. Shared cranidial 4, figs 10-12, pl. 6, fig. 7, pl. 7, fig. 4; Liñán et characteristics include three transglabellar furrows al. 2005, fig. 7.2) is easily distinguished from and very short (tr.) posterolateral projections of Megapharanaspis in that it is not subdivided the fixigenae (see Pocock 1970; Paterson & into a pro- and opisthothorax, nor does it Edgecombe 2006). However, the cranidium of possess a macropleural segment. The pygidium Megapharanaspis is easily distinguished by of Megapharanaspis differs considerably its anteriorly tapered glabella, longer (exsag.) from most species of Lemdadella, for e.g., L. palpebral lobes and shorter (tr.) eye ridges. The linaresae (Liñán Guijarro 1978, pl. 4, figs 5-9) librigena of Megapharanaspis is nearly identical and L. antarcticae (Palmer in Palmer & Rowell to that of Balcoracania except that the latter has 1995, figs 8.10-8.12, 8.17). However, Zhang a shorter (exsag.) palpebral suture and a shorter (1997, fig. 279.3b) illustrated a pygidium of (tr.) posterior margin (Paterson & Edgecombe the type species of Lemdadella, L. spectabilis, 2006, figs 5-8). Thoracic similarities include that is similar to that of Megapharanaspis in the division into pro- and opisthothorax, with having a subelliptical outline, smooth posterior the post-macropleural segments representing margin and two axial rings and terminal piece. the opisthothorax, width (tr.) of the axis and an Nonetheless, the pygidium of Megapharanaspis identical macropleural segment. The macropleural can be differentiated from that of L. spectabilis in segment in Megapharanaspis is identical in every having a narrower (tr.) axis that does not reach the detail to that in emuellids, with the exception that posterior margin and effaced pleural regions. in the latter, the macropleural segment is fused From a phylogenetic standpoint, Mega- to the adjacent anterior (fifth) segment. Major pharanaspis appears to represent a sister taxon AAP Memoir 32 (2006) 51 to the Emuellidae (Emuella + Balcoracania). The margin (excluding occipital ring) straight, directed inclusion of Megapharanaspis in the Emuelloidea slightly posterolaterally. Anterior sections of is strongly supported by the character evidence facial suture diverging anteriorly at 50-55° presented above, but its systematic position is between γ and β, then weakly converging between better illustrated in terms of thoracic tagmosis. β and α. Posterior sections of facial suture very Emuellids display prothoracic/opisthothoracic short and strongly diverging posteriorly. Glabella tagmosis, but Paterson & Edgecombe (2006) moderately convex, gently tapered anteriorly, clearly demonstrate that this thoracic division sagittal length 80-85% cranidial length, anterior evolved independently in several clades within the width (tr.) at midlength of frontal lobe 75-85% Redlichiida. However, emuellid thoracic tagmosis occipital ring width; frontal lobe rounded, sagittal is unique in that the prothoracic/opisthothoracic length 25-30% cranidial length. Axial furrow boundary is marked by the macropleural segment, shallow, narrow (tr.) and straight; preglabellar as is the case in Megapharanaspis, whereas in other furrow shallow and narrow (sag., exsag.). S1 and phylogenetically disparate clades, e.g. Olenellina, S2 transglabellar, moderately impressed, deepest the prothoracic/opisthothoracic boundary is abaxially, abaxial portions directed slightly decoupled from the position of the macropleural posteromedially, becoming transverse sagittally; segment (Lieberman 1998, 2001). It is therefore S3 transglabellar, moderately impressed, deepest possible that the position of the macropleural abaxially, transverse. Occipital ring length (sag.) segment at the prothoracic/opisthothoracic approximately 10-15% cranidial length; anterior boundary represents an autapomorphy for the margin straight; posterior margin strongly Emuelloidea. Megapharanaspis can be excluded convex posteriorly. SO straight, narrow (sag., from the Emuellidae in that the macropleural exsag.), of moderate depth across its entire width. segment is not fused to the adjacent anterior Preglabellar field very short, length (sag.) 5-10% segment; fusion of the fifth thoracic segment to cranidial length; plectrum weakly developed, the sixth macropleural segment is considered an width (tr.) approximately 50% width of frontal autapomorphy for the Emuellidae (Paterson & glabellar lobe. Preocular field gently downsloping Edgecombe 2006). anteriorly, flat to gently convex, maximum length (exsag.) approximately 20-25% cranidial length Megapharanaspis nedini gen. et sp. nov. (Fig. (sag.). Anterior border flat to weakly convex, 5) length (sag., exsag.) approximately 10% cranidial length; anterior border furrow very narrow (sag., Etymology. After Chris Nedin, for his work on the exsag.), well developed laterally, shallow medially Emu Bay Shale Lagerstätte at Big Gully. where posterior margin of border extends back to form plectrum. Palpebral lobe well developed, Diagnosis. As for genus. crescentic, strongly convex, length (exsag.) 40-45% cranidial length (sag.), width 25-35% Type material. Holotype: internal and external lobe length, anterior tip situated opposite S3, mould of partial exoskeleton (SAMP41183, Fig. posterior tip situated opposite SO; palpebral 5A-D). Paratypes: internal mould of cephalon furrow well-developed, shallow. Eye ridge well and first four thoracic segments plus external developed, forming continuation of palpebral mould of most of thorax and pygidium of the lobe, strongly convex, short, width (tr.) 15-20% same individual (SAMP41184); external mould cranidial length (sag.), straight, ridges diverge of partial cranidium and part of first five thoracic posteriorly at 120-140°, meets axial furrow just segments (SAMP41185); internal mould of forwards of S3 at posterior portion of frontal cephalon and first seven thoracic segments glabellar lobe. Palpebral area flat, width at ε (tr.) (SAMP41186); external mould of almost approximately 45% adjacent glabellar width. complete thorax (SAMP41187). Postocular area absent. Posterolateral projection of fixigena extremely short (exsag., tr.). Posterior Description. Small opisthoparian trilobite, border moderately convex (exsag.), very narrow length of largest complete specimen, 8.5 mm; (exsag.), slightly expanding abaxially; posterior other available specimens suggest a maximum border furrow of moderate depth, wide (exsag.), length of about 13 mm. Cephalon semicircular expanding abaxially. in outline, sagittal length approximately 50-60% Hypostome and rostral plate unknown. maximum width (tr.) and 40% total length of Librigena small, up to 8 mm in length exoskeleton; low convexity (sag., tr.); surface (including genal spine); width (tr.) approximately smooth. Cranidium subquadrate in outline; 40-45% length (excluding genal spine); lateral sagittal length 90-95% maximum width (tr.); margin moderately curved, continuing evenly onto anterior margin moderately curved; posterior genal spine; posterior margin straight, width (tr.) 52 AAP Memoir 32 (2006)

Fig. 5. Megapharanaspis nedini gen. et sp. nov. A-D, holotype, SAMP41183. A, internal mould of almost complete exoskeleton, x7; B, internal mould of pygidium, x15; C, latex cast of external mould of almost complete exoskeleton, x6; D, latex cast of external mould of pygidium, x13. E, internal mould of cephalon and anterior of thorax, SAMP41186, x4. F, internal mould of cephalon and anterior of thorax, SAMP41184, x5. G, latex cast of external mould of partial cranidium and anterior of thorax, SAMP41185, x7.5. H, latex cast of external mould of partial thorax, SAMP41187, x4.

45-50% librigenal length (excluding genal spine). segments; prothoracic segments longer (sag., Genal field almost flat, width (tr.) at midlength exsag.) and wider (tr.) than opisthothoracic 55-65% librigenal width. Lateral border flat to segments. Axis moderately convex (tr.), gently weakly convex, width (tr.) at midlength 35-45% and evenly tapering posteriorly, width (tr.) librigenal width; lateral border furrow shallow, approximately 50% width of each segment; axial narrow (tr.). Posterior border weakly convex, rings with small medial nodes. Axial furrows length (exsag.) 5-10% librigenal length (excluding well defined. T1 and T2 pleurae directed laterally genal spine); posterior border furrow shallow and (perpendicular to axial line), width (tr.) of narrow (exsag.). Genal spine long, length 45-50% pleura approximately 25% width of prothoracic librigenal length (including spine), with distal tip segment, projected horizontally to fulcrum, then opposite fourth or fifth thoracic segment; base slightly downsloping to pleural spine; anterior narrow, moderately tapering posteriorly, slight pleural band of moderate convexity, rapidly adaxial curvature. Faint caeca on genal field, expanding abaxially to fulcrum, then tapers to radiating out from palpebral lobe. pleural spine; posterior pleural band of moderate Thorax of 18 segments, divided into prothorax convexity, expanding abaxially; pleural spine of three segments and opisthothorax of 15 short, nonfalcate (‘thorn-like’); pleural furrow AAP Memoir 32 (2006) 53 moderately deep, very wide (exsag.) proximally, cranidium subtrapezoidal in outline; glabella rapidly tapering abaxially. T3 macropleural, subrectangular with slight median waist, extends pleura strongly directed posterolaterally; anterior to anterior border furrow; lateral glabellar furrows pleural band rapidly expanding abaxially to poorly defined to absent. Occipital ring smooth. base of macropleural spine; posterior pleural Palpebral lobes situated at or slightly posterior to band considerably narrower (exsag.) than midlength. Eye ridge poorly developed, low relief, anterior pleural band, slightly tapering abaxially very narrow, directed posterolaterally, meets axial to base of macropleural spine; macropleural furrow at anterior quarter of glabella. Librigena spine extending almost to level of pygidium; with narrow field; genal spine long. Thorax of T3 pleural furrow deep, very wide (exsag.) eight segments. Pygidial axis extends to posterior proximally, rapidly tapering abaxially, extending margin; axis with five to eight axial rings plus onto base of macropleural spine, then becoming terminal piece; four to six pairs of pleural ribs; shallow and narrow along length of spine. pygidial margin spinose or smooth. Opisthothoracic pleurae become progressively more posterolaterally directed posteriorly, sharply Remarks. Palmer (in Palmer & Rowell 1995, tapering abaxially; pleural furrows moderately p. 15) originally placed Holyoakia in the deep, very wide (exsag.) proximally, rapidly Emuellidae “with serious reservation”. Paterson tapering abaxially; opisthothoracic pleural spines & Edgecombe (2006) suggested placement relatively longer than T1 and T2 pleural spines. within the Dorypygidae based on the subquadrate Pygidium very small, subelliptical in outline, glabella, absence of a preglabellar field, short sagittal length approximately 45% maximum (exsag.) preocular field, well differentiated eye width (tr.); weakly convex (sag., tr.); posterior ridge and palpebral lobe, wide (tr.) posterolateral margin smooth. Axis strongly tapered, anterior projection of the fixigena, pygidium of subequal width (tr.) 30-35% pygidial width; length (sag.) size with the cranidium, pygidial axis well defined 80-85% pygidial length; low convexity (tr.); two with eight axial rings and a terminal piece, well axial rings and a terminal piece; axial furrow developed pleural ribs and furrows, and a spinose shallow; anterior inter-ring furrow shallow, but margin. The specimens of Holyoakia from Big better impressed than posterior inter-ring furrow. Gully support this assignment. Pleural region effaced. Border absent. Holyoakia simpsoni sp. nov. (Fig. 6) Remarks. The small size of Megapharanaspis nedini gen. et sp. nov., with a maximum Etymology. For David Simpson who discovered exoskeletal length (sag.) of 13 mm, raises the and collected the first specimens. question of whether the type specimens represent holaspides. Unfortunately, its rarity in the Emu Diagnosis. Holyoakia with occipital ring Bay Shale prohibits a census on its size range in length (sag.) 10-15% glabellar length and a statistical sample, but the pygidial morphology posterior margin of occipital ring slightly convex suggests the type specimens do indeed represent posteriorly; palpebral lobe small, length (exsag.) holaspides as protothoracic segments are absent. 20-25% cranidial length (sag.), situated opposite Furthermore, the close phylogenetic relationship midlength of glabella; palpebral area width (tr.) of Megapharanaspis to the Emuellidae suggests at ε 90-100% adjacent glabellar width; pygidium that small holaspides are characteristic of the transversely elliptical in outline, sagittal length Emuelloidea, since the largest emuellid, B. dailyi, 45% maximum width (tr.), anterior margin reaches a maximum exoskeletal length (sag.) of strongly curved; pygidial axis comprises five about 30 mm (Paterson & Edgecombe 2006). rings plus terminal piece; four pairs of pleural ribs with shallow interpleural furrows; border narrow, Order CORYNEXOCHIDA Kobayashi, 1935 with distinct nodes on margin of border adjacent Suborder CORYNEXOCHINA Kobayashi, to distal ends of pleural furrows. 1935 Family DORYPYGIDAE Kobayashi, 1935 Type material. Holotype: internal and external mould of almost complete exoskeleton Holyoakia Palmer in Palmer & Rowell, 1995 (SAMP41188, Fig. 6A, B). Paratypes: internal mould of almost complete exoskeleton Type species. Holyoakia granulosa Palmer in (SAMP41189); internal moulds of two partial Palmer & Rowell, 1995, p. 15, figs 9.2, 11.6- exoskeletons (SAMP41190, 41191); and an 11.10. internal mould comprising partial pygidium and partially attached thorax (SAMP41192). Diagnosis. Small opisthoparian trilobite; 54 AAP Memoir 32 (2006)

Fig. 6. Holyoakia simpsoni sp. nov. A, B, holotype, complete exoskeleton, SAMP41188; A, internal mould, x6; B, latex cast of external mould, x6.5. C, internal mould of complete exoskeleton, SAMP41189, x6. D, internal mould of partially complete exoskeleton, SAMP41190, x7. E, internal mould of partial pygidium and posterior part of thorax, SAMP41192, x5. F, internal mould of almost complete exoskeleton, SAMP41191, x7.

Description. Small opisthoparian trilobite; length (exsag.) 20-25% cranidial length (sag.), reniform, of largest available complete specimen, 7.5 mm. opposite midlength of glabella; palpebral furrow Cephalon semicircular in outline, sagittal length shallow. Eye ridge poorly developed, low relief, approximately 50% width (tr.) along posterior very short (exsag.), moderately curved, directed margin and 40% total length of exoskeleton; low posterolaterally, meets axial furrow at anterior convexity (sag., tr.); surface slightly granulose. quarter of glabella. Palpebral area gently convex, Cranidium subtrapezoidal in outline; anterior width (tr.) at ε 90-100% adjacent glabellar width. margin gently curved; posterior margin gently Postocular area moderately convex, length curved with lateral portions directed slightly (exsag.) 20-25% cranidial length. Posterolateral posterolaterally. Anterior sections of facial projection of fixigena gently downsloping, sutures slightly convergent anteriorly; posterior directed slightly posterolaterally, width (tr.) 15- sections of facial sutures gently curved and widely 20% width of posterior cranidial margin. Posterior divergent posteriorly. Glabella subrectangular border convex (exsag.), width (exsag.) expanding with slight median waist, reaching anterior border abaxially; posterior border furrow well developed, furrow; width (tr.) at LO approximately 30% of moderate depth, width (exsag.) expanding width of posterior margin; frontal lobe truncate; abaxially. S1-S3 represented by indentations on the side of Hypostome not preserved separately, but the glabella rather than distinct furrows. Axial on at least two specimens (Fig. 6A, B, D) the and preglabellar furrows shallow. Occipital ring impressions of a small hypostome, approximately short, length (sag.) 10-15% glabellar length, 50% the length of the glabella, can be seen. posterior margin slightly convex posteriorly. SO Librigena with narrow (tr.) field; genal spine well-developed, of even depth across entire width. extends along the margin of the thorax to about Preglabellar field absent. Preocular field very short third thoracic segment; border furrow narrow, (exsag.), gently downsloping. Anterior border shallow; border convex. convex, extremely short, length (sag., exsag.) less Thorax of eight segments; width (tr.) of than 5% cranidial length; anterior border furrow segments slightly decreases posteriorly. Axis shallow and narrow. Palpebral lobe small, length moderately convex, tapers evenly to posterior; AAP Memoir 32 (2006) 55 width (tr.) of axis 25-30% width of thorax. Axial shallow interpleural furrows. The pygidial axis of furrows well defined. Axial ring gently curved H. granulosa has eight axial rings and a terminal anteriorly. Thoracic pleurae wide, width (tr.) 35- piece; that of H. simpsoni has five axial rings and 40% width of thoracic segment; inner portion a terminal piece. of pleura projecting laterally and horizonally to fulcrum, then outer portion downsloping ACKNOWLEDGEMENTS posterolaterally to pleural spine; anterior pleural The two new species described herein were brought band slightly narrower (exsag.) than posterior to our attention by David Simpson (Adelaide). pleural band; pleural spine very short, falcate; JBJ thanks David Simpson and family for their pleural furrow shallow, deepening abaxially, hospitality during field work on Kangaroo Island. extending almost to distal tip of pleural spine. Paul Buck kindly granted access to Big Gully Pygidium transversely elliptical in outline, through his farming property. Sarah Laurence sagittal length 45% maximum width (tr.), low (South Australian Department of Environment convexity (sag., tr.); anterior margin strongly and Heritage) is thanked for organising permission curved. Axis strongly convex, moderately to collect from the Big Gully site. Thanks to Jim tapered, extends to posterior margin; anterior Gehling (South Australian Museum) for valuable width (tr.) 25% maximum width of pygidium; advice, and Dennis Rice (South Australian axial furrows narrow (tr.), shallow; axis of five Museum), for arranging the cataloguing of the rings, each bearing a faint medial node, and specimens described and figured herein. Peter terminal piece; ring furrows well developed Jell constructively commented on an earlier draft and short (sag., exsag.). Pleural regions well of the manuscript. 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