Lower and Middle Devonian Radiolarian Biozonation of the Gamilaroi Terrane New England Orogen, Eastern Australia

Lower and Middle Devonian radiolarian biozonation of the Gamilaroi terrane New England Orogen, eastern Australia

Jonathan C. itch is on', Aileen M. ~avis ' , James M. C. stratford2, and Frances C. P. spiller3 'Department of Earth Sciences, University of Hong Kong, Pokfulam Road, Hong Kong, China

'school of Pure and Applied Sciences, University of the South Pac$c, P. 0.Box 1168, Suva, Fiji 3~epar tmentof Geology and Geophysics, University of New England, Armidale, NSW 2351, Australia

ABSTRACT: Seven distinctive radiolarian assemblages defined from the Gamilaroi terrane of eastern Australia form the basis of an uppermost Lower to Middle Devonian radiolarian biostratigraphy. In ascending order these are the: Stigrnosphaerostylus horrida, Helenifore laticlavium, Circulaforma admissarius, Heleniforepilosidiscus n. sp., Protoholoeciscus hindea, Ceratoikiscum regalinodus, and Trilonche minax assemblages. Gamilaroi terrane sedimentation occurred during the interval Early (Pragian) to Late (Frasnian) De- vonian in a volcanic island arc environment in which radiolarians were abundant but sedimentary facies were locally variable. Radiolarian assemblages can be used for terrane-wide correlation. Although assemblages are dominated by spumellarians they also include more distinctive ceratoikiscid forms. Two new ceratoikiscid taxa: Circulaforma davidi n. sp. and Helenifore pilosidiscus n. sp are described.

INTRODUCTION

Radiolarian biostratigraphy back to the uppermost (Famennian) Devonian is well established (Cheng 1986; Holdsworth and Jones 1980; Schwartzapfel and Holdsworth 1996). Recent work on the Lower Paleozoic marine strata has shown further potential for development of a useful biostratigraphic zonation amongst the earliest radiolarians (Noble and Aitchison 1995) and establishment of a Silurian biozonation is already well advanced (Noble 1994). Intervening radiolarian faunas of the Lower and Middle Devonian however, remain poorly known and their biostratigraphy has not been established. In this paper we report the results of investigations of Lower and Middle De- vonian radiolarian-bearing volcaniclastic sedimentary sections in eastern Australia and present the outline of a potential biozonation.

Well-preserved Lower to Middle Devonian radiolarians have been known from the Gamilaroi terrane in eastern Australia since the end of last century when G. J. Hinde (1899) described specimens from samples collected by David and Pittman (1899). For almost 100 years very little further attention was paid to radiolarians from the Gamilaroi terrane. At the cente- nary of the first report (David and Howchin 1896) of Gamilaroi terrane radiolarians it seems appropriate that our understanding of these fossils is updated. Recent work has confirmed the presence of radiolarians in the terrane (Aitchison 1988; Aitchison and Flood 1992; Aitchison, et al. 1992; Ishiga, et al. 1988; Spiller 1992, 1993; Stratford and Aitchison 1997; Metcalfe et al. 1997). Aitchison and Stratford (1997) revised and reassessed the original material described by Hinde and demonstrated (Stratford and Aitchison 1997) that a workable biostratigraphy could be established within this terrane. This paper builds on our previous work and we present results of a much wider sampling program throughout southern New England Orogen (NEO). This permits us to test the utility of the preliminary biozonation outlined from rocks in the Glenrock area.

Gamilaroi terrane radiolarian faunas are important for several reasons. The potential to elucidate a major-gap in radiolarian biostratigraphy in the Lower to Middle Devonian is of global importance. Few previous detailed reports exist in which radiolarians of this age are described (Hinde 1899) and the Gamilaroi terrane faunas offer an excellent indication of the nature of forms present at that time. Devonian strata likely to contain radiolarians have a wide global distribution thus it is important that a workable biostratigraphy be established for this period. An expanding knowledge of their biostratigraphy also provides an opportunity to interpret radiolarian evolution. Nazarov and Ormiston (1985, 1986, 1993) have suggested that the Early to Middle Devonian was an important time in radiolarian evolution. Data presented herein provide information which helps to elucidate these evolutionary changes. In a more local context, the various radiolarian assemblages enable us to make a better interpretation of the relative age of strata within the complex suite of different lithologies which make up the Gamilaroi terrane. Age control has traditionally been limited to isolated limestones which typically comprise less than five percent of all exposed lithologies. Therefore stratigraphic correlation constrained by radiolarian assemblages has the capacity to give a much firmer basis for understanding the tectonic evolution of this terrane as well as constraining links with adjacent terranes throughout the New England orogen.

REGIONAL STRATIGRAPHY

Radiolarians described in this study were collected from the De- vonian volcaniclastic sediments of the Gamilaroi terrane (Flood and Aitchison 1988 1992), the westemmost terrane in the NEO. This terrane comprises a complex association of volcaniclastic sediments, tuffs, volcanic rocks and minor carbonates formed in an intra-oceanic island-arc setting (Aitchison and Flood 1994, Stratford and Aitchison 1996) and accreted to the eastern margin of Gondwana sometime during the Late Devonian (Flood and Aitchison 1992).

micropaleontology, vol. 45, no. 2, pp. 138-162, text-figures 1-2, plates 1-6, 1999 138

Micmpaleontology, vol. 45, no. 2, 1999

TAMWORTH BELT HASTINGS BLOCK

Rundifferentiated rocks of the New England orogen

Overlap assemblage (predominantly Carboniferous rocks)

(Upper Devonian rocks) ----------

Gamilaroi terrane (Silurian-UpperDevonian)

TEXT-FIGURE 1 Simplifiedtectonostratigraphic map of the southern New England orogen, eastern Australia showing the areal distribution of rocks assigned to the Gamilaroitemane. Significant radiolarian localities are also indicated.

and overlap assemblage rocks (Devonian-Carboniferous)

J. C. Aitchison et al.: Lower and Middle Devonian radiolarian biozonation of the Gamilaroi terrane New England Orogen, eastern Australia

Gamilaroi terrane strata in the southern NEO have an arcuate distribution pattern throughout northeastern NSW over a distance of approximately 450 km (Figure 1). The lithostratigraphic succession within the terrane was first described from Nundle (Crook 1961) and formations within the Tamworth Group can be traced, with varying confidence, along strike for approximately 120 km from Attunga through Tamworth to Nundle. Radiolarians are abundant in fine-grained predominantly tuffaceous marine cherty sediments of the terrane. The best radiolarian-bearing lithologies are laminated black and white tuffs and mudstones of the Yarrirnie Formation at the top, and fine-grained tuffaceous horizons within the Silver Gully Formation in the middle of the Tamworth Group.

The terrane can be followed further SE from Nundle into the Upper Barnard River catchment. Detailed Tamworth Group lithostratigraphy cannot be correlated easily with strata in this area due to inherent variations in sedimentation patterns in the Gamilaroi terrane depositional setting and an independent local lithostratigraphic subdivision has been developed (Stratford and Aitchison 1998). The lithostratigraphic units in the Upper Barnard catchment southwest of Nundle can be traced from Barry through Glenrock to Pigna Barney (approx. 100 krn). The most productive radiolarian-bearing units are green or tan tuffaceous mudstones in the Frog Hollow Formation towards the base of the sequence and similar lithologies in the Curricabark Formation at the top of the sequence.

Other exposures of Gamilaroi terrane strata southeast of the Up- per Barnard occur around and to the south of Bundook in the Lower Manning River area. Potentially important radiolarian- bearing lithologies (Ishiga and Leitch 1988) also crop out in this area. The lithostratigraphic sequence in this area has never been described in any significant detail and awaits detailed geological mapping. Our reconnaissance surveys indicate that lithologies similar to those found elsewhere in the Gamilaroi terrane; felsic volcanic rocks, basalts (Benson 1918), and volcaniclastic sediments, are present.

The Hastings Block (for details see Roberts, et al. 1995) lies west of Port Macquarie. The oldest known strata are Middle to Upper Devonian sedimentary and volcanic rocks which can be correlated with the Gamilaroi terrane. Younger (Upper Devo- nian to Carboniferous) sedimentary rocks in this region are sim- ilarly correlated with their younger equivalents in the Tamworth Belt (Roberts, et al. 1993). Radiolarians have been extracted from predominantly tan coloured tuffaceous mudstones in the western portions of this block (Ishiga and Leitch 1988, Spiller 1992, 1993). Radiolarian faunas in the Hastings Block occur in both the intra-oceanic island arc sequence (Gamilaroi terrane) and the overlap assemblage as is seen in the Tamworth Belt. The overall lithological assemblage and abundance of radiolarians is strongly indicative of correlation with the Gamilaroi terrane although local facies variations make detailed stratigraphic correlation difficult.

RADIOLARIAN LOCALITIES

Several important localities from which Gamilaroi terrane radiolarians have been collected are briefly described below. Some additional localities from which radiolarians have been extracted lie within sediments deposited on top of rocks of the Gamilaroi terrane (sensu strict0 Aitchison and Flood 1992). These sedimentary rocks are referred to herein as an overlap assemblage and they place a minimum age constraint on the tim- ing of the development of the Gamilaroi terrane. Localities for

which grid references (GR) are given conform to the Australian Map Grid with eastings followed by northings. The names of relevant Central Mapping Authority NSW 1: 25 000 topo- graphic map sheets are also given. Four digit locality numbers prefixed with an L refer to sites registered on the University of New England index. Specimens referred to are either assigned Australian Museum catalogue numbers, prefixed AMF or Uni- versity of Hong Kong, Department of Earth Sciences catalogue numbers prefixed HKUDES.

With the exception of the Upper Barnard and Upper Manning catchments exposure of Gamilaroi terrane rocks is woeful and there are no areas where exposure is continuous. Nevertheless, as broadly comparable lithostratigraphies are widely recognisable and regional lithostratigraphic correlation is possible (Stratford and Aitchison 1998). Such correlation is greatly aided by examination of the radiolarian content of tuffaceous cherts and provides a useful test of the utility of our biozonation.

Attunga

Numerous exposures within the Yarrimie Formation around Yarramanbully and east of Attunga (30 krn north of Tamworth) were examined. Preservation of recovered material was typically poor such that definitive identification of taxa was not possible.

Tuffaceous cherts, volcaniclastic sandstones, and rare limestone lenses which together comprise the Yarrimie Formation crop out in the area around Tamworth City (David and Pittman 1899). We (Aitchison and Stratford 1997) have examined and redescribed the original material discussed by (Hinde 1899). We have also recollected along the original (David and Pittman 1899) line of section and various other localities around Tam- worth. In most instances although radiolarian 'ghosts' are visi- ble in hand specimen the original amorphous opaline silica has been replaced during diagenesis by zeolite minerals. Identifi- able radiolarians, of assemblage 7, were recovered from the original Hinde samples (Aitchison and Stratford 1997) and from Bournes Lane (Tamworth 9035-I-N GR 997660) slightly to the north of Tamworth. Conodonts which occur in limestone lenses immediately beneath the Hinde radiolarian locality are ascribed to the upper Givetian (hermanni zone) or younger. Lower Frasnian conodonts occur further upsection, just below the first Baldwin Formation (overlap assemblage) rocks (J. Talent pers. comm.).

Loomberah Heights

From a road-cut at housing lot 2 in Norman Close (Dungowan 9135-IV-S GR 128484), a new residential subdivision at Loomberah Heights to the south of Tamworth, we report moderately well preserved radiolarians of assemblage 4. Mawson, et al. (1995) reported an upper Emsian serotinus or lower patulus zone conodont fauna from limestones at this locality. The limestones occur as small blocks within a mass flow unit located stratigraphically below the radiolarian locality and their context suggests correlation with the Silver Gully Formation. Thus, they place a maximum age constraint on this fauna.

Silver Gully

Despite very poor and discontinuous exposure Silver Gully is location of the type sections of several of the constituent formations of the Tamworth Group (Crook 1961). Important

radiolarian faunas have also been collected from this area. They confirm that the radiolarians collected and described from Glenrock (Stratford and Aitchison 1997) are found elsewhere in the Gamilaroi terrane and that these assemblages occur in a similar succession.

From strata in a roadside cutting near the location where Silver Gully enters Chaffey Dam (Woolomin 9135-111-N GR 203300) we have collected radiolarians from green coloured tuffs mapped by Cawood (1982) within the Drik Drik Formation at the base of the Tamworth Group (Crook 1961). A low diversity (assemblage 1) radiolarian fauna which includes Stigmo-sphaerostylus horrida is present. We note that the radiolarians in this sample are accompanied by abundant sponge spicules and the possibility of some sort of facies control on the occurrence of this radiolarian assemblage cannot be excluded. Cono- dont data from limestones within the Drik Drik Formation (Furey-Greig 1995) suggest that these rocks may be Lower De- vonian (lower Emsian).

Conodont data (Furey-Greig 1995) constrain the Silver Gully Formation at the base of the section to the upper Emsian. Many samples were collected from the top of the Silver Gully Forma- tion through to the top of the Yarrimie Formation. Samples were collected from several different localities along Silver Gully yielding radiolarians of assemblages 4-7. Numerous entactiniids occur together with Helenifore laticlavium, Circulaforma davidi n. sp., Helenijore pilosidiscus n. sp. and Protoholoeciscus hindea in the lower Yarrimie Formation (Woolomin 9135- 11 1-N GR 1873 12). From slightly higher upsection we found Ceratoikiscum regalinodus as well as Palaeoscenidium cladophorum and numerous entactiniids (Woolomin 9135-1 11-N GR 187307). Continuing upsection, a sample from Woolomin 91 35- 11 1-N GR 184307 contained Helenifore laticlavium, and entactiniids, and a further sample (Woolomin 9135- 11 1 -N GR 1803 11) has produced a fauna similar to that of the Hinde locality at Tamworth.

Timor Caves

Samples containing radiolarians (assemblage 3) were collected from tuffaceous mudstones both above and below extensive outcrops of limestone at Timor Caves. On the basis of coral and conodont data this unit ranges from the mid-Eifelian through to uppermost Givetian (Pedder 1967, Philip and Pedder 1967). From below the limestones (Isis River 9134-IV-S GR 174963) in presumably lower Emsian strata of the Lilberne beds (Manser 1968) we obtained a moderately well-preserved fauna which includes poorly preserved specimens of Circulaforma admissarius, Circulaforma davidi n. sp. and various entactiniids.

Glenrock

Many samples were collected from sections through the Gamilaroi terrane at Glenrock and five radiolarian assemblages (2-6) were reported by Stratford and Aitchison (1997). The Helenifore laticlavium assemblage is the lowermost assem-blage in the Gamilaroi terrane (Stratford and Aitchison 1998) at Glenrock and includes the nominative species as well as Palaeoscenidium sp., Ceratoikiscum sp. A and various entactiniids. Aitchison (1993a) also reported an assemblage (2) containing Helenifore laticlavium from the same stratigraphic unit at Glenrock and this fauna is described in detail by Metcalfe et al. (1997). Four other assemblages (3-6) occur within younger strata and are, in ascending order: Circulaforma

Micropaleontology, vol. 45, no. 2, 1999

admissarius assemblage, Helenifore pilosidiscus n. sp. assemblage, Protoholoeciscus hindea assemblage, and the Ceratoikiscum regalinodus assemblage. A detailed discussion of the relative position and inferred stratigraphic ranges of these assemblages is presented by Stratford and Aitchison (1997).

Pigna Barney

The entire Gamilaroi terrane stratigraphic succession in the Pigna Barney district (upper Manning River catchment) was sampled for radiolarians. Although specimens were recovered from many samples the material present was generally poorly preserved and diagnosis is difficult. Nevertheless, tuffaceous mudstones located close to, and stratigraphically immediately above, limestones from which Emsian conodonts have been collected from Bennys Tops (Pigna Barney 9234-3-N GR 719802) and have yielded fragments of Helenifore laticlavium (Dongal 1995). Significant collections were made in the Pigna Barney River near Limestone Creek and along the Manning River on Glen Wards Road. Faunas characteristic of assemblage 2-3 were recovered from both of these localities and consist primarily of moderately well-preserved entactiniids. Helenijore laticlavium is common in most samples from low in the section (within the Frog Hollow Formation of Stratford and Aitchison 1998). Ceratoikiscids diagnostic of younger radiolarian assemblages were not recovered from any samples further upsection. Radiolarian ghosts are abundant in hand specimen but attempts at extraction of identifiable material were not successful.

Bundook

A strip of Gamilaroi terrane rocks occurs in the area south of the Khanghat Fault from Bundook SE to at least the Pacific High- way (Roberts, et al. 1991). Reconnaissance traverses in this area indicate the presence of a full range of Gamilaroi terrane lithologies including sodically-altered pillow basalts (Benson 1918), felsic extrusives, and accompanying volcaniclastic sediments. Traverses were made along several roads and forestry tracks which cross the area and tuffaceous mudstones were collected for radiolarian processing. Fragments of Helenifore laticlavium together with numerous entactiniids indicative of assemblage 2 were recovered (Bundook 9334-3-S GR 135695). Stratigraphy appears complex and it is unlikely the section is either homoclinal or unbroken by faulting.

Hustings Block

Radiolarian samples have been collected from a displaced block of Gamilaroi terrane rocks which crop out in the western part of the Hastings Block approximately 100 km west of Port Macquarie. Helenijore laticlavium (assemblage 2) occurs in tuffaceous sediments at Kennedy Farm (UNE L1932 locality reported by Spiller 1992, 1993 [Birdwood 9335-11-N GR 3353141) which are associated with a thick section of basaltic rocks similar to those which occur in the Glenrock area (Folly Volcanics; Stratford and Aitchison 1998). The only potential age constraint on these basaltic rocks is a U P b SHRIMP age of 377 + 8 Ma (Givetian) (Aitchison and Ireland 1995) for plagiogranite which may have formed in association with an arc-rifting episode during which the basalts were generated (Aitchison et al. 1994). However, we note that this correlation is tenuous and further work is required in this area. Pillow basalts occur at various levels within the Gamilaroi terrane and are not confined to the Folly Volcanics. Indeed we note the presence of pillow basalts from amongst H. laticlavium bearing strata at Glenrock. The presence of Middle Devonian corals (Pickett 1985) elsewhere in the western Hastings Block suggests that

J. C. Aircllison et al.: Loiter and Middle Devonian radiolarian biozonation of the Gamilaroi rerrane Nerr England Orogen, eastern A~rstralia

system Ma stages

Carboniferous Tournatslan

360

Famennian L

367 a 3

Frasnian 374

K Givetian .-a K

380

2 E

g Eifelian

387

Emsian 394

t Pragian

401

Lochkovian a 8

Stlurian Prtdollan

TEXT-FIGURE 2

a Q& W2 Holoec~scus foremanae (8)

Bournes Lane

Tr~lonchem~nax(7) (7) W5 Cerato~k~scumregabnodus(6) [r [r

Protoholec~scush~ndea(5)

5 Helen~forep~los~d~scus(4) III

rad~olar~an locat~on assemblages

Tamworth Sllver Gully Tlmor Glenrock P~gna Barney Bundook Hastlngs Block

&. 4-(7) ' (7)

'kb &% (6) (6)-

. (5)-+ . (5)Lwmberan

(4) (4) 'kb (4)

I @ UP@ SHRIMP

&% mnodonr

-2

Distribution of various Devonian radiolarian assemblages at locations throughout the Gamilaroi terrane. New England oropen. eastern Australia and associated paleontolopicaliradiometric control points.

further radiolarian studies may enhance correlations with the rest of the Gamilaroi terrane. The original stratigraphy of rocks along the western edge of the Hastings Block has been heavily disrupted by faulting and the original succession is yet to be de- termined.

Helerzijore laticlaviunl occurs together with many unidentified entactiniids at Yarras Road [UNE L1931 Yarras 9335-11-S GR 3542331. H. laticlavium was also reported from nearby at locality I41 of Ishiga and Leitch (1988). At these localities archocyrtids also occur in the radiolarian assemblages. To- gether with fragments of the conodont Polygrzathus asyrn- metricus from the Yarras Road locality they confirm lower Frasnian affinities for these rocks (Spiller 1992).

RADIOLARIAN CONSTRAINTS ON THE OVERLAP ASSEMBLAGE

Holoeciscus formanae assemblage

Holoeciscus forrnanae and other taxa, including archocyrtids, which indicate that the containing strata are Famennian (Cheng 1986; Holdsworth and Jones 1980; Schwartzapfel and Holds- worth 1996) occur in samples collected from the sequence of post intra-oceanic island arc rocks which overlie the Gamilaroi terrane. This assemblage is present at numerous localities detailed below.

Yanolee

Radiolarians have been collected from laminated tuffs south of Glenrock in the overlap assemblage along Branch Creek Road (Glenrock GR415885: Yanolee Farm) located between Gamila-

roi terrane exposures at Timor Caves and Glenrock. The radiolarian fauna is simple and contains Holoeciscus foremarzae in addition to rare archocyrtids and numerous entactiniids. The presence of H. forernanae indicates middle to upper Famennian affinity (Cheng 1986. Schwartzapfel and Holdsworth 1996) for this locality.

Bobo Creek

Ishiga and Leitch (1988) reported an uppermost Devonian radiolarian assemblage which includes archocyrtids from outcrops near the Khanghat Fault in the Bobo Creek district. These outcrops occur on the northeastern limb of an anticlinal structure in the Bundook area of which Gamilaroi terrane strata form the core. Tuffs at this locality are intercalated with pillow basalts. If these basalts are part of the Gamilaroi terrane then they are younger than any known from elsewhere in the terrane. Fur- ther detailed mapping work is required in this area along with geochemical studies to test the affinity of the basalts. On the basis of their age we would suspect that these rocks are likely to constitute part of the overlap assemblage.

Costigans Road, Yarras Quarry and Birdwood Quarry

Upper Devonian to Lower Carboniferous radiolarians are common in tuffaceous mudstones which overlie Gamilaroi terrane rocks in the western Hastings Block (Ishiga and Leitch 1988, Spiller 1992. Spiller 1993). In contrast to the western Hastings Block localities described above, these sites have yielded younger radiolarians thought to be associated with rocks correlative to the overlap assemblage. The fauna includes Holoeciscus forernarzae, Ceratoikiscurn hexastellaturn, Ceratoikiscilrn

quinquestellatum, Archocrytium spp., Kantollum sp.. Popofsky-ellum sp., Protoalbaillella sp., Robotium sp., Palaeoscenidium clacloyhorum and various entactiniids. The fauna is uppermost Devonian to lowermost Carboniferous.

RADIOLARIAN ASSEMBLAGES By combining all available data from many localities within a single terrane several important conclusions can be reached and a local biostratigraphy developed. Stratford and Aitchison (1997) recognised five radiolarian assemblages within the Gamilaroi terrane succession at Glenrock. Our investigations along the strike-length of the terrane suggest that these distinctive assemblages, along with two others described herein, form the basis of the uppermost Lower to Middle Devonian radiolarian biostratigraphy for this region. In ascending order the assemblages we recognise are: Stigmosphaerostylus horrida, Helenifore laticlavium, Circulaforma admissarius, Helenifore pilosidiscus n. sp., Protoholoeciscus hindea, Ceratoikiscurn regalinodus, and Trilonche minax assemblages. These assemblages define partial range or acme zones. Gamilaroi terrane sedimentation occurred during the interval Early (Pragian) to Late (Frasnian) Devonian. Conodonts, which are rare in limestones within the Gamilaroi terrane, provide valuable age constraints on these assemblages at some localities and younger radiolarians, which are interpreted to be from strata that constitute part of an overlap assemblage, provide a minimum age constraint. This overlap assemblage is considered to be Famennian or younger (Flood and Aitchison 1992).

The terrane-wide sampling program reported in this paper indicates that this biostratigraphy is robust. It can be used for correlation throughout the terrane despite poor rock exposure and the absence of other age diagnostic faunal elements. Our biostratigraphic zonation is not, however, considered definitive, rather it is presented as a local biozonation which can be used as a guide. It should be tested by, and further refined in the light of, future detailed work on Middle Devonian radiolarians worldwide.

Stigmosphaerostylus horrida assemblage ( 1 )

From strata in a roadside cutting near the location (Woolomin 9135-111-N GR 203300) where Silver Gully enters Chaffey Dam we have collected radiolarians from green coloured tuffs mapped by Cawood (1982) within the Drik Drik Formation at the base of the Tamworth Group (Crook 1961). A low diversity radiolarian fauna which includes Stigmosphaerostylus horrida is present. Radiolarians in this assemblage show a tendency to be very fragile. Although entactiniids of this assemblage possess three-bladed spines these are generally thin and less well developed than those seen in younger faunas. We note that the radiolarians in this sample are accompanied by abundant sponge spicules and the possibility of some sort of facies control on the occurrence of this radiolarian assemblage cannot be excluded. Conodont data from limestones within the Drik Drik Formation (Furey-Greig 1995) suggest that these rocks are Lower Devonian (lower Emsian).

Helenifore laticlavium assemblage (2 )

This assemblage is distinguished by an abundance, or acme, of specimens of Helenifore laticlavium with respect to other ceratoikiscids. Other radiolarians present are predominantly entactiniids with rare Palaeoscenidium cladophorum and rare Ceratoikiscum sp. The entactiniids are characteristically robust with well-developed three-bladed spines. This fauna is wide- spread but is confined to localities stratigraphically low in the


Gamilaroi terrane at Glenrock (Stratford and Aitchison 1997; Metcalfe et a1 1997), and along regional strike at Pigna Barney (Dongal 1995). It is also found at Kennedy Farm UNE L1932 in the Hastings Block in rocks of uncertain stratigraphic context. The blue grey tuffs and pillow basalts in which the assemblage occurs at Kennedy Farm are, however, similar to lithologies found at lower stratigraphic levels at Glenrock.

Lower Devonian (Emsian) conodonts occur in limestones intercalated with tuffs containing the Helenifore laticlavium assem-blage In our experience samples in which Helenifore laticlavium is the dominant ceratoikiscid are typically from Lower Devonian sections of the terrane. Helenifore laticlavium group is known to range at least as high as the Upper Devonian (Aitchison 1993b, Nazarov et al. 1982) but younger forms are not readily distinguishable from those seen earlier in the Devo- nian. However, where this taxon occurs within younger assemblages, at least within the Gamilaroi terrane, it occurs together with a wide range of other ceratoikiscids and entactiniids.

Circulaforma admissarius assemblage (3)

Together with the nominative species this assemblage includes; Circulaforma davidi n. sp., Ceratoikiscum calvum, Trilonche spp. and various other entactiniids. Compared with forms in younger Gamilaroi terrane assemblages entactiniids in the Circulafor~naadmissarius assemblage are typically simpler and much less robust. This assemblage first appears 40-50 m stratigraphically above lower Emsian limestone at Glenrock (Stratford 1995). It also occurs in red ribbon bedded cherts of another NEO terrane, the Djungati terrane (Aitchison 1993a). The presence, at the Timor Caves locality, of mid-Eifelian limestones above the tuffs that contain this assemblage provides an important minimum age constraint. This assemblage is thus most probably lower Eifelian (lowermost Middle Devonian).

Helenifore pilosidiscus n. sp. assemblage (4)

This assemblage occurs at Glenrock above strata containing the Circulaforma admissarius assemblage where Helenijore pilosidiscus n. sp. is found together with Circulaforma admissarius and very poorly preserved three-bladed entac-tiniids. Circulaforma admissarius is commonly present only as broken fragments. This assemblage also occurs in the lower part of the Yanimie Formation at Silver Gully. It is interpreted to be mid to upper Eifelian.

Protoholoeciscus hindea assemblage ( 5 )

The robust and distinctive taxon Protoholoeciscus hindea oc-curs at both Silver Gully and Glenrock where it is found together with rare specimens of Ceratoikiscum regalinodus, Ceratoikiscum paragyium, Circulaforma davidi n. sp. as well as indeterminate ceratoikiscid caveal rib fragments and Palaeoscenidium. Entactiniids are common and include Stigmosphaerostylus spp. and Trilonche spp. Faunas belonging to the Protoholoeciscus hindea assemblage are the most diverse known from the Gamilaroi terrane. This fauna is interpreted (Stratford and Aitchison 1997) to indicate that strata are Middle Devonian (uppermost Emsian to lower Givetian). Proto-holoeciscus is also known from Eifelian strata of the Kuro- segawa terrane in Japan (Umeda and Yamagiwa, 1997).

Ceratoikiscum regalinodus assemblage (6)

This is stratigraphically the highest assemblage present at Glenrock. It is also known from Silver Gully where the Gamilaroi terrane sequence continues further upsection. It includes Ceratoikiscum regalinodus, Palaeoscenidium spp. and

J. C. Aitchison et a/.: Lobcer and Middle Devonian radiolarlan blozonation of the Gamilarol termne New E~zglancl Orogen, emtern Au~tralra

numerous different entactiniids. Entactiniids within this assemblage are similar to those found in samples collected from the Hinde (1899) and Bournes Lane localities at Tamworth. The Ceratoikiscum regalinodus assemblage is interpreted to be upper Givetian and may be equivalent to (at least to the older part of) the Spongentactinella windjanensis - Trilonche nigra as-semblage (see systematic section for a discussion of synony- mies) of Nazarov (1988).

Trilonche minax assemblage (7)

This assemblage was collected from the Hinde (1899) localities at Tamworth is dominated by a diverse array of entactiniids (Aitchison and Stratford 1997). It also occurs in samples collected from the upper levels of the Yarrimie Formation at Silver Gully. Comparison with faunas known from elsewhere suggests correlation with parts of the upper Givetian portion of the Spongentactinella windjanensis - Heliosphaera nigra and Polyentactinia circumretia - Trilonche minax assemblages of Nazarov (1988) as all the nominative taxa are Dresent (Trilonche minax is regarded as the senior synonym of Entactinosphaera egindyensis see Aitchison and Stratford 1997). In this work we refer to this assemblage as the Trilonche minax assemblage. Conodonts occur in strata both above and below this fauna and provide uppermost Givetian to lower Frasnian constraints (Aitchison and Stratford 1997).

DISCUSSION

Radiolarians are common in tuffaceous lithologies of the Gamilaroi terrane but until recently their stratigraphic significance has been somewhat ambiguous. A combination of new radiolarian localities described herein and conodont (Furey-Greig 1995, Mawson et al. 1995), coral (Philip and Ped- der 1967, Stratford 1995, Stratford and Wright in prep) and UIPb SHRIMP (Aitchison et al. 1995) data are all in accord with our previous age estimates for the assemblages (Stratford and Aitchison 1997). Additional radiolarian work described by Aitchison and Stratford (1997) provides a link through to younger faunas known from Upper Devonian (lowest Frasnian) strata elsewhere (Aitchison 1993b, Nazarov et al. 1982) (Figure 2). Further detailed sampling is required in sections elsewhere to test the global applicability of the biostratigraphy. Neverthe- less we consider that we have now established the basic framework for a Lower to Middle Devonian radiolarian biozonation.

Our zonation is transitional into existing Upper Devonian radiolarian zonations (Holdsworth and Jones 1980, Cheng 1986, Schwartzapfel and Holdsworth 1996). Studies of Silurian radiolarian biozonation are already well advanced and constrained using conodont data (Noble 1994). However, a zonation which links existing Silurian biozonations and our Lower to Middle Devonian biozonation is not yet developed. Upper Silurian to Lower Devonian radiolarian bearing sections are known from Japan (Aitchison et al. 1991, 1996; Fumtani 1983, 1990; Ishiga 1992; Suzuki et al. 1996; Wakamatsu et al. 1990). Until recently, however, a lack of independent age constraints hindered understanding of the age significance of these faunas. Radiometric age constraints (Aitchison et al. 1996) indicate that the Japanese faunas provide an overlap with, and are most likely transitional between, the Silurian zonation from North America and the Devonian biozonation from the Gamilaroi terrane described herein. Recent work by M. Umeda (pers comm.) has revealed the presence in Japan of some taxa common to rocks of the Gamilaroi terrane in strata known to be Middle Devonian. We are hopeful that further work will soon

result in an integrated zonation for the entire Silurian through Devonian.

SYSTEMATIC PALEONTOLOGY

Suborder SPUMELLARIA Ehrenberg 1875 Family ENTACTINIIDAE Riedel 1967 emend. Nazarov and

Ormiston 1984 Subfamily ENTACTINIINAE Riedel 1967 emend. Nazarov

1975

Genus Trilonche Hinde 1899 emend. Foreman 1963 emend. Aitchison and Stratford 1997 Tjye Species: Trilonche vetusta Hinde

TriloncheHinde 1899, p. 48. -AITCHISON and STRATFORD 1997 p. 373.

Staurodruppa HINDE 1899, p. 51. Entactinosphaera FOREMAN 1963, p. 274.

Diagnosis: Two or more, well developed, spherical or sub-spherical shells, and an internal, six-rayed double spicule. When secondary spines are present, the six spines developed from the rays of the internal spicule are generally differentiated and recognisable as main spines. Up to six main spines may be present.

Remarks: The diagnosis of Hinde (1899) restricted the genus to specimens with three radial spines. In 1963 Foreman published a less restrictive, more general diagnosis of a synonymous genus Entactinosphaera which encompassed material described as Trilonche by Hinde. Aitchison and Stratford (1997) placed Entactinosphaera in synonymy and adopted the definition of Foreman (1963) as an appropriate emendation for the diagnosis of the senior synonym Trilonche. Although up to six (possibly more) main spines may be present it is common for not all spines to be developed to a similar degree. Where, in addition to the three bladed main spines, smaller "by-spines" are present these subordinate spines typically have a rod- or needle-like form.

Range and occurrence: Present in Upper Paleozoic radiolarian-bearing strata worldwide.

Trilonche davidi (Hinde 1899) emend. Nazarov 1975 Plate 3, figures K and L; plate 4, figures C, D, F, G, K, L; plate 5, figures M and N; plate 6, figures C and S

Staurolonche davidi HINDE 1899, p. 46, pl. 8, fig. 13. Staurolonche tenella HINDE 1899, p. 47, pl. 8, fig. 14. Staurolonche laterna HINDE 1899, p. 47, pl. 8, fig. 15. Staurodruppa nucula HINDE 1899, p. 52, pl. 9, fig. 7. Staurolonchidium obliquum HINDE 1899, p. 47, pl. 8, fig. 17. Entactinosphaera grandis NAZAROV 1975, p. 65, pl. 5, figs. 11, 12; pl.

7, figs. 1-4. Trilonche davidi (Hinde) 1899 - AITCHISON and STRATFORD 1997,

p. 380, pl. 1, fig. 8; pl. 2, fig. 4.

Diagnosis: The detailed diagnosis of the synonymised taxon Entactinosphaera grandis (Nazarov 1975, p. 65) was accepted (Aitchison and Stratford 1997) as an emendation of the diagnosis of this species.

Remarks: The test has two concentric spherical shells, robust main spines and rare, long, by-spines. The main spines are mas- sive, approximately equal in size, and three-bladed. They have the same width along their entire length. The outer shell is thick,

1997

with rounded or rounded-oval pores. By-spines are rare, quite thin, rod-like and terminate bluntly.

Range and occurrence: Present in samples from throughout the Gamilaroi terrane. Known from the Devonian of eastern NSW, Australia, North America and Russia.

Tn'lonche echinata (Hinde 1899) emend. Aitchison and Stratford

Plate 1,figures G-K, N; plate 2, figures D, K, L, Q; plate 3, figures 0and X; plate 4, figure S; plate 5, figures B-D, G, I; plate 6, figures A, I, L-N, P

Heliosoma echinatum HINDE 1899, p. 50, pl. 9, fig. 1, 2. Heliosomaparonae HINDE 1899, p. 50, pl. 9, fig. 3. Entactinosphaera palimbola -FOREMAN 1963, p. 277, pl. 2, fig. 7a-e;

pl. 3, fig: 3a-d. A

Entactinosohaera echinata? (Hinde)?- FOREMAN 1963. D. 279. ~ 1 .3. fig. 10; $4, fig. 12a-b. - NAZAROV 1975,p. 60,pl. 111,'figs. 1-3; pl. IV., figs. 1-4.

Entactinosphaera assidera NAZAROV 1975, p. 64, pl. V, figs. 6,7; pl. VI., figs. 6-8.

~ntactin&~haeracf. E. assidera NAZAROV 1975 - AITCHISON 1993b,p. 115, pl. 6., figs. 2, 6.

Trilonche echinata (Hinde).- AITCHISON and STRATFORD 1997, p. 377, pl. 1, figs. 5, 6, 10.

Diagnosis: Two concentric spherical lattice shells with approximately six three-bladed main spines of variable width and length. By-spines are common but are variable and typically thorn-like. One main spine is typically substantially longer and more robust than the others. An internal six-rayed spicule is present within the inner shell.

Remarks: See Aitchison and Stratford (1997) for a detailed discussion of this taxon.

Range and occurrence: Present in samples from throughout the Gamilaroi terrane. Known from the Devonian of eastern NSW, Australia, North America and Russia.

Tn'lonche elegans Hinde 1899 emend. Foreman (1963) Plate 1, figures C, D, F, P; plate 4, figures E, I and 0 ; plate 6, figures B and R

Trilonche elegans HINDE 1899, p. 48, pl. 8, fig. 22. - AITCHISON and STRATFORD 1997, p. 378, pl. 1, fig. 7; pl. 2, fig. 6.

Staurodruppa nana HINDE 1899, p. 52, pl. 9, fig. 8. Entactinosphaera symphypora FOREMAN 1963, p. 277, pl. 2, fig. 5a-c;

pl. 3, fig. 4.

Remarks: Aitchison and Stratford (1997) synonymized Entactinosphaera symphypora Foreman with the senior synonym Trilonche elegans but adopted the more detailed diagnosis of Foreman (1963, p. 277), who worked with matrix free material, as an appropriate emendation.

Range and occurrence: Present in samples from throughout the Gamilaroi terrane. Known from the Devonian of eastern NSW, Australia, and North America.

Trilonche hindea (Hinde) emend. Aitchison and Stratford 1997 Plate 6, figures D, E, T

Stylosphaera vetusta HINDE 1899, p. 46, pl. 8, fig. 10. Entactinosphaera vetusta HINDE 1899 - NAZAROV 1975, p. 64,pl. 5,

figs. 8-10; pl. 6, figs. 9-11. Trilonche hindea (Hinde) - AITCHISON and STRATFORD 1997, p.

376, pl. 1, fig. 9.


Diagnosis: Two concentric spherical shells. Two polar main spines somewhat larger than four other main spines which are arranged equatorially at 90' to one another. All spines are three-bladed.


Range and occurrence: Present in samples from upper levels of the Gamilaroi terrane at Bundook and Tamworth. Known from the Devonian of eastern NSW, Australia and Russia.

Tn'lonche minax (Hinde 1899) emend. Aitchison and Stratford 1997 Plate 2, figures H and I; plate 5 , figures J and K

Xiphosphaera minax HINDE 1899, p. 45, pl. 8, fig. 8. Stylosphaera obtusa HINDE 1899, p. 45, pl. 8, fig. 9. Staurodruppapraelonga HINDE 1899, p. 51, pl. 9, fig. 6. S t a ~ r o d r u ~ ~ a ? ~ r o l a t aFOREMAN 1963, p. 281, pl. 8, figs. la-b; pl. 9,

fig. 12. Entactinosphaera egindyensis NAZAROV 1975, p. 61, pl. 5, figs. 2,4;

pl. 6, figs. 1-5. Trilonche minax (Hinde) - AITCHISON and STRATFORD 1997, p.

377, pl. 1, figs. 3,4; pl. 2, figs. 3, 5.

Diagnosis: Spherical radiolarian with two concentric shells. Relatively thick cortical shell with small pores. Inner medullary shell small and coarsely porous. Two very large, long, robust, strongly grooved, polar main spines. The width of the base of the two main spines is approximately one radius of the outer shell or greater. Four other small, commonly rodded, main spines may be present on a plane perpendicular to the two major heavy three bladed spines.


Range and occurrence: Present in samples from the Gamilaroi terrane at Loomberah and Glenrock. Known from the Devonian of eastern NSW, Australia, North America and Russia.

Tn'lonche robusta (Aitchison) Plate 2 , figure F; plate 4, figure M

Entactinosphaera robusta AITCHISON 1993, p. 116, pl. 5, fig. 15.

Diagnosis: See Aitchison 1993b p. 116

Description: The figured specimens are very similar to the type specimen as described by Aitchison (1993). They have several, originally possibly six, three-bladed main spines. These spines are long and some exhibit torsion. Numerous short rod-like by-spines are also present. The main shells have numerous small sub-circular pores.

Range and occurrence: Present in samples from the Gamilaroi terrane at Glenrock Station. Known from the Devonian of eastern NSW, Australia, and Canning Basin, Western Australia.

Trilonche vetusta Hinde 1899 emend. Aitchison and Stratford 1997 Plate 2, figures J and R; plate 3, figure E; plate 5, figures F and H; plate 6, figures K, 0 , Q

Trilonche vetusta HINDE 1899, p. 48, pl. 8, figs. 18, 19. Trilonchepittmani HINDE 1899, p. 48, pl. 8, figs. 20, 21. Theodiscus hastatus HINDE 1899, p. 53, pl. 9, fig. 9. ?Triactiscus lanceola HINDE 1899, p. 53, pl. 9, fig. 11. Entactinosphaera aitpaiensis NAZAROV 1973, p. 697,pl. 1, figs. 4-5. -

NAZAROV 1975, p. 63, pl. 5, figs. 3, 5 ; pl. 6, figs. 2-4.

J. C. Aitchison et al.: Lower and Middle Devonian radiolarian biozonation of the Gatnilaroi terrane New England Orogelz, eastern Australia

Trilonche vetusta HINDE 1899 - AITCHISON and STRATFORD 1997, p. 374, pl. 1, figs. 1, 2; pl. 2, figs. 1, 2.

not Stylosphaera vetusta HINDE 1899, p. 46, pl. 8, fig. 10 Entactino~~hnern NAZAROV 1975, p. 64, pl. 5, figs.vetusta (Hinde) -

8-10; pl. 6, figs. 9-11

Diagnosis: Thin cortical test with small lattice pores and secondary spines. Inner shell has small circular to polygonal pores. Three main spines which are moderately stout, typically sub-equal in length although the third may be somewhat smaller than the other two. Main spines are strongly three-bladed and connect through to the inner shell and are disposed at approximately, 120" to each other.


Range and occurrence: Present in samples from throughout the Garnilaroi terrane. Known from the Devonian of eastern NSW and Russia.

Trilonche sp. A Plate 3, figure J

Description: Three-bladed thin main spines, tapered from the base with a series of two to four small rounded spinules spread from 113 to 213 the length down the ridge from the base of the spine. Apparently six main spines. Shell pores are not well preserved. Poorly preserved shell interiors indicate affinities with Trilonche.

Range and occurrence: Locality SW-5 of Stratford (1996) in the Gamilaroi terrane at Glenrock Station, Middle Devonian [?Eifelian] as far as is known.

Trilonche sp. B Plate 3, figures G and M; plate 4, figure R; plate 5, figures R-U

Description: Slender three-bladed main spines, number uncertain. By-spines are very prominent and between 113-213 the length of the main spines and often broken. By-spines appear to be circular in cross-section towards the tip and weakly

PLATE 1 All scale bars = 100p-n.

Figures A-E Helenifore laticlavium assemblage from stratigraphically low levels of the Gamilaroi terrane at Glenrock Station.

A Helenifore laticlavium Nazarov and Ormiston C Trilonche ?elegans (Hinde) HKUDES 9710102 HKUDES 9710101

D Trilonche ?elegans (Hinde) HKUDES 9710103 B Helenifore laticlavium Nazarov and Orrniston AMF

96648 E Entactiniid gen. et sp. indet. B HKUDES 9710104

Figures F-P Helenifore laticlavium assemblage from stratigraphically low levels of the Gamilaroi terrane (Frog Hollow Formation of Stratford and Aitchison 1998) at Pigna Barney.

F Trilonche e l e~ans ? (Hindel HKUDES 9710105 "

G Trilonche echinata ? (Hinde) HKUDES 9710106

H Trilonche echinata ? (Hinde) HKUDES 9710107

I Trilonche echinata ? (Hinde) HKUDES 9710108

J Trilonche echinata ? (Hinde) HKUDES 9710109

K Trilonche echinata ? (Hinde) HKUDES 9710110

L Helenifore laticlavium Nazarov and Ormiston

M Helenifore laticlavium Nazarov and Ormiston HKUDES 97101 12

N Trilonche echinata (Hinde) HKUDES 97/01 13

0 Stigmosphaerostylus ? sp. HKUDES 9710114

P Trilonche elegans ? (Hinde) HKUDES 97/01 15

Figures Q-X: Stigmosphaerostylus horrida assemblage from green colored tuffs mapped by Cawood (1980) within the Drik Drik For- mation at the base of the Tamworth Group (Crook 1961) where Silver Gully enters Chaffey Dam (Woolomin 9135-111-N GR 203 300)

Q Stigmosphaerostylus sp. A HKUDES 97/01 16

R Stigmosphaerostylus sp. A HKUDES 97101 17

S Sti~mosphaerostylus horrida (Furutani) HKUDES 9770118

T Stigmosphaerostylus sp. B HKUDES 97/01 19

U Stigmosphaerostylus horrida (Furutani) HKUDES 9710120

V Stigmosphaerostylus horrida (Furutani) HKUDES 9710 12 1

W Stigmosphaerostylus horrida (Furutani) HKUDES 97/01 22 . -

X Stigmosphaerostylus sp. B HKUDES 9710123

J. C. Aitchison et al. Plate 1

micropaleontology, vol. 45, no. 2, 1999 147

J. C.Aitchison et al.: Lower and Middle Devonian radiolariarz biozonatiorz of the Gamilaroi terrane New England Orogen, eastern Australia

three-bladed at the base. Pores are well developed. Some specimens have one large main spine longer than all the others.

Range and occurrence: Localities SW-5 and SW-7 of Stratford (1996) at Glenrock, Middle Devonian (?Eifelian) as far as is known.

Genus Stigmosphaerostylus Rust 1892, emend. Foreman 1963 Type species: Stigmosphaerostylus notabilis Riist 1892

Stigmosphaerost)lusRUST 1892,p. 142. Ellipsostigma HINDE 1899, p. 51. Entactinia FOREMAN 1963, p. 27 1. StigornosphaerosvlusRust - AITCHISON and STRATFORD 1997, p.

380.

Remarks: Foreman (1963) introduced a new genus named Entactinia which she described from matrix-free specimens. This genus is a junior synonym of Stigmosphaerostylus Riist

1892. However as Foreman's (1963) diagnosis is more precise it was adopted as an appropriate emendment by Aitchison and Stratford (1997).

Range and occurrence: Present in Upper Paleozoic radiolarian-bearing strata worldwide.

Stigmosphaerostylus gogoensis? (Aitchison) 1993 Plate 4, figure U

Entactinia gogoensis AITCHISON 1993, p. 112,pl. 5, fig. 3.

Diagnosis See Aitchison 1993b p. 113

Description Specimen has a spherical lattice shell with several delicate medium length three-bladed main spines. Several short spinules are developed at a node on the distal sections of the main spines. Similar to the specimen, figured by Aitchison

PLATE 2 Radiolarians representative of Circulaforma admissarius and Helenifore pilosidiscus assemblages. All scale bars = 100pm.

Figures A-F: Radiolarians of the Helenifore pilosidiscus assemblage collected from Glenrock Station.

A Helenifore pilosidiscus n. sp. Aitchison and Stratford D Trilonche echinata ? (Hinde) HKUDES 9710125 AMF 96672 holotype

E Entactiniid gen. et sp. indet. B HKUDES 9710126 B Helenifore pilosidiscus n. sp. Aitchison and Stratford

AMF 96674 paratype F Trilonche robusta (Aitchison) HKUDES 9710127.

C Entactiniid gen. et sp. indet. HKUDES 9710124

Figures G-M: Radiolarians of the Helenifore pilosidiscus assemblage collected from Loomberah area.

G Helenifore ?pilosidiscus n. sp. Aitchison and Stratford Figures N-S: Radiolarians of the Circulaforma HKUDES 97/01 28 admissarius assemblage collected from Timor dis-

trict. H Trilonche minax (Hinde) HKUDES 9710129

N Circulaforma admissarius Stratford and Aitchison I Trilonche minax(Hinde) HKUDES 9710130 HKUDES 9710135

J Trilonche vetusta Hinde HKUDES 97101 3 1 0 Circulaforma admissarius Stratford and Aitchison HKUDES 9710136

K Trilonche echinata (Hinde) HKUDES 9710132 P Circulaforma admissarius Stratford and Aitchison

L Trilonche echinata (Hinde) HKUDES 9710133 HKUDES 9710 137

M Entactiniid gen. et sp. indet. HKUDES 9710134. Q Trilonche echinata ? (Hinde) HKUDES 9710139

R Trilonche vetusta Hinde HKUDES 9710140.

Figures S-W: Radiolarians of the Circulaforma admissarius assemblage collected from Glenrock Station.

S Circulaforma admissarius Stratford and Aitchison V Ceratoikiscum calvum Stratford and Aitchison HKUDES 9710141 HKUDES 9710143

T Circulaforma admissarius Stratford and Aitchison W Stigmosphaerostylus sp. HKUDES 9710144 AMF 96665, U: Ceratoikiscum calvum Stratford and Aitchison HKUDES 9710142

J. C. Aitchison et

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J. C. Aitchison et al.: Lower and Middle Devonian radiolarian biozonation of the Garnilaroi terrane New England Orogen, eastern Australia

(1993b; Plate 5, figure 3) although the lattice shell appears to be more heavily constructed.


Stigmosphaerostylus horridQ (Furutani) Plate 1, figures S, U-W

Tlecerina horrida FURUTANI 1983, p. 110-111,pl. 23, fig. 7; pl. 24, figs. 1-3; pl.25, figs. 1-3. - WAKAMATSUet al. 1990,pl. 10, figs. 5-7.

Description: Specimens are closely similar to taxa described by Furutani (1983) as Tlecerina horrida. They have coarsely porous external shells. They also have several slender, grooved three-bladed spines which taper to a sharply pointed tip. One spine is commonly more prominent that the others.

Remarks: Specimens bear a superficial external resemblance to Trilonche elegans Hinde 1899 emend. Foreman (1963). How- ever, they posses a single rather than double shell. Their spines are also thinner and more deeply grooved along their entire length. Furutani (1983) named this taxon Tlecerina horrida and considered it to represent a form of Palaeosceniid based on the presence of an internal spicule which he regarded as closely similar to the spicular structure of "unshelled" Palaeosceniids. This internal spicule is homologous to the spicule present on Palaeosceniids but we disagree with the interpretation of Furutani (1983) of a genetic relationship to the Palaeo-

scenidiinae. Whilst it is possible to see relationships between the unshelled Palaeoscenidiurn, and shelled Parentactinia and Pactarentinia sensu Furutani (1983) it is not possible to conclu- sively demonstrate any evolutionary lineage between this taxon and the true Palaeosceniids. Although superficially similar, the spicule of Stigrnosphaerostylus horrida is markedly smaller (less than one-tenth the size). The mere possession of an homologous structure is not an adequate basis for demonstration of a phylogenetic relationship. Until a relationship can be unequivo- cally demonstrated we consider this taxon most appropriately assigned to family and genus (Stigmosphaerostylus)with whose gross characteristics it most closely fits.

Range and occurrence: Known from the Gamilaroi terrane (Drik Drik Formation) at Silver Gully. Also known from rocks of the Kurosegawa terrane at Yokokurayama on Shikoku, Japan (Furutani 1983,1990; Wakamatsu et al. 1990) but with poor age control. Lower Devonian (lower Emsian) so far as is known.

Stigmosphaerostylus hystricuosa (Aitchison) 1993 Plate 3, figure H

Entactinia hystricuosa AITCHISON 1993, p. 113,pl. 5, figs. 1, 2.


Description: Imperfectly preserved specimen which is externally similar to specimens of Stigmosphaerostylus hystricuosa (Aitchison), figured (as Entactinia hystricuosa) by Aitchison (1993b PI. 5, figs. 1,2). The external shell is small, and spheri-

PLATE 3 Radiolarians representative of the Protoholoeciscus hindea assemblage collected from Glenrock Station. All scale bars = 100ym.

A Circulaforrna davidi n. sp. Aitchison and Stratford N Stigrnosphaerostylus profundisculus (Aitchison) HKUDES 9710145 paratype HKUDES 9710 154

B Ceratoikiscurn paragyiurn Stratford and Aitchison 0 Trilonche echinata (Hinde) HKUDES 9710155 HKUDES 9710146

P Protoholoeciscus hindea Aitchison HKUDES C Ceratoikiscurn paragyium Stratford and Aitchison 9710156

AMF96687 holotype Q Protoholoeciscus hindea Aitchison HKUDES

D Ceratoikiscum para~yiurn Stratford and Aitchison 9710157

R Protoholoeciscus hindea Aitchison HKUDES E Trilonche vetusta Hinde AMF96693 9710158

F Entactiniid gen. et sp. indet. HKUDES 9710148 S Stigrnosphaerostylus ? sp. B HKUDES 9710159

G Trilonche sp. B HKUDES 9710148 T Helenifore pilosidiscus n. sp. Aitchison and Stratford HKUDES 9710 160

H Stigrnosphaerostylus hystricuosa (Aitchison) HKUDES 9710 150 U Circulaforma davidi n. sp. Aitchison and Stratford

I Entactiniid gen. et sp. indet. A HKUDES 9710151 HKUDES 9710161 holotype

J Trilonche sp. A AMF 96694 V Circulaforma davidi n. sp. Aitchison and Stratford

HKUDES 9710162 paratype

K Trilonche davidi (Hinde) AMF 96697 W Circulaforrna davidi n. sp. Aitchison and Stratford

L Trilonche davidi (Hinde) HKUDES 9710152 HKUDES 9710163 paratype

M Trilonche sp. B HKUDES 9710153 X Trilonche echinata (Hinde) HKUDES 9710164


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J. C. Aitchison et al.: Lower and Middle Devonian radiolarian hiozonation o f the Gamilaroi terrane New England Orogen, eastern Australla

cal. Main spines are long and deeply bladed. Several tiers of short spinules are evident on the distal sections of the main spines with each tier having three spinules, one developed on each blade of the main spine.


Stigmosphaerostylus profundisculus (Aitchison) 1993 Plate 3, figure N

Entactiniaprofundisculus AITCHISON 1993, p. 114. pl. 7, figs. 7, 8.


Description: Poorly preserved specimen which is externally similar to specimens of Entactinia profundisculus (Aitchison), figured (as Entactinia profundisculus) by Aitchison (1993b Pl;. 7, figures 7, 8). The specimen is robust with a spherical lattice shell. Those spines which are preserved are three-bladed and deeply grooved with a single tier of spinules developed on their distal portion.

Range and occurrence: Present in samples from the Gamilaroi terrane at Glenrock Station. Known from the Devonian of east- em NSW, Australia, and Canning Basin, Westem Australia.

Stigmosphaerostylus sp. A Plate 1, figures Q and R

Description: Specimens possess an ovoid shell (imperfect sphere) Approximately 8 short thin three-bladed spines are present. Pores are subcircular and irregular with a distinctive open lattice framework.

Range and occurrence: Known from the Gamilaroi terrane (Drik Drik Formation) at Silver Gully. Lower Devonian so far as is known.

Stigmosphaerostylus sp. B Plate 1, figures T and X; plate 4, figure T

Description Specimens are coarsely, or openly porous, with slender, fragile spines. These spines are proximally three-bladed and rodded at their distal extremities.

Range and occurrence: Known from the Gamilaroi terrane (Drik Drik Formation) at Silver Gully and from locality SW-7 of Stratford (1995) at Glenrock Station. Lower to Middle Devo- nian so far as is known.

Entactiniid gen. et sp. indet. A Plate 3, figure I; Plate 4, figures J and N

Entactinia sp., AITCHISON 1988, p. 54, fig. 3s. Entactinosphaera sp. B., ISHIGA 1988, p. 87, pl. 5 , fig. 10. Entactinosphaera sp. C., ISHIGA 1988, p. 87, pl. 5 , fig. 8.

Remarks: Specimens have up to six robust three-bladed spines. The first half of the spine from the base is straight. Spines then undergo torsion of between 1 to 1.25 rotations. Most commonly, but not exclusively, coiled sinistrally [looking towards the tip]. Spines taper to a fine point close to the tip. Pores are subcircular, with small nodes at the junction of the pores.

Range and occurrence: Localities SW-5, SW-6, and SW-7 of Stratford (1996) in the Gamilaroi terrane at Glenrock Station. It is also found in the Birpai subterrane and in the Djungati terrane. Probable age range Middle to Late Devonian.

Entactiniid gen. et sp. indet. B Plate 1, figure E; plate 2, figure E; plate 5, figures E and L

PLATE 4 Radiolarians typical of the Ceratoikiscum regalinodus assemblage collected from Glenrock Station. All scale bars = 100pm.

A Ceratoikiscum regalinodus Stratford and Aitchison AMF 96701 holotype

B Ceratoikiscum regalinodus Stratford and Aitchison HKUDES 9710165

C Trilonche davidi ? (Hinde) HKUDES 9710166

D Trilonche davidi ?(Hinde) HKUDES 9710167

E Trilonche elegans Hinde HKUDES 9710168

F Trilonche davidi ? (Hinde) HKUDES 9710169

G Trilonche sp. HKUDES 9710170

H Spongentactinia ? sp. AMF 96730

I Trilonche elegans ? (Hinde) HKUDES 9710171

J Entactiniid gen. et sp. indet. A AMF 96721

K Trilonche davidi ? (Hinde) HKUDES 9710172

L Trilonche davidi ? (Hinde) HKUDES 9710173

M Trilonche robusta ? (Aitchison) AMF 96726

N Entactiniid gen. et sp. indet. A HKUDES 9710174

0 Trilonche elegans ? (Hinde) HKUDES 9710175

P Entactiniid gen. et sp. indet. HKUDES 9710176

Q Entactiniid gen. et sp. indet. HKUDES 9710177

R Trilonche sp. B HKUDES 9710178

S Trilonche echinata (Hinde)HKUDES 9710179

T Stigmosphaerostylus ? sp. B AMF 96723

U Stigmosphaerostylus gogoensis ?? (Aitchison) AMF 96718

V Helioentactinia ? sp. AMF 96715


micropaleontology, vol. 45, no. 2, 1999 153

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Remarks: Two long robust three-bladed spines. The angle between the spines varies from 180 - 160" through the centre of the shell. One spine may be longer than the other with some main spines having minor torsion usually less than 114 rotation. Commonly has abundant small by-spines. Pores not well preserved.

Range and occurrence: Found at localities NE-1, NE-, SW-4, and SW-6 of Stratford (1996) at Glenrock. Age range is ?Lower-Middle Devonian as far as is known.

Suborder ALBAILLELLARIA Deflandre 1953, emend. Holds- worth 1969

Family CERATOIKISCIDAE Holdsworth 1969

Genus Ceratoikiscum Deflandre 19.53 Type species: Ceratoikiscum avimexpectans Deflandre 1953

Ceratoikiscum calvum Stratford and Aitchison 1996 Plate 2, figures U and V

? Ceratoikiscum spp. WAKAMATSU et al. 1990, pl. 9, fig.7. ? Ceratoikiscum sp. indet. FURUTANI 1990, pl. 13, fig. 3. Ceratoikiscum calvum STRATFORD and AITCHISON 1997, p. 224,

pl. 2 A-C.

Diagnosis: A species of Ceratoikiscurn with a simple, well-defined triangular frame and poorly developed caveal ribs. It has three long, slightly curved main rods with six well- developed extra-triangular rods. The extra-triangular rods, es- pecially the a.p., a.a., i.d., and b.d. rods are longer than the rods which form the triangular frame and they may also be more curved. All rods are thin with a circular cross-section. Four or five pairs of simple thin caveal ribs are developed on a.t. and all ribs curve a short distance straight downward. The largest rib originates from the a-b joint, the other ribs are spread out along

213 the length of the a.t. rod towards the posterior. The i-b joint commonly exhibits slight thickening.

Range and occurrence: Found in the Circulaforma admissarius assemblage from locality SW-1 of Stratford and Aitchison (1997) [GR 482 9371 in the Gamilaroi terrane at Glenrock. Up- permost Lower to lowermost Middle Devonian (Emsian-Eifelian) so far as is known.

Ceratoikiscum paragyium Stratford and Aitchison 1997 Plate 3, figures B-D

Ceratoikiscumparagyium Stratford and Aitchison 1997. p. 224, pls. 4E. I-K.

Diagnosis: This species has the basic triangular frame of Ceratoikiscum with an additional a-rod (aa-rod). A dense zone of ribs with associated patagial tissue originates from the a-b joint, forming a tent-like structure. Two a-rods, designated a and aa are developed. The aa-rod originates at the a-b joint and curves out in the plane of the a-rod to rejoin the frame at the a-i joint, where it may rarely form a small aa.p. rod. All seven extra-triangular rods tend to be long and slender with the exception of the aa.p. and a.p. sections which are rarely seen. A dense zone of patagial tissue originates from the a-b joint and forms a tent-like structure extending along the length of a.p. aa.p. and b.t. Fine caveal ribs oriented in the plane of the tent protrude from each free edge of the tent. The b.t. section is commonly bent past the free edge of the tent.

Range and occurrence: Locality SW-5, of Stratford and Aitchison (1997) [GR 469 9101, Protoholeciscus hindea assemblage at Glenrock. Middle Devonian (Eifelian) so far as is known.

Ceratoikiscum regalinodus Stratford and Aitchison 1997 Plate 4, figures A and B; Plate 5, figures P and Q

PLATE 5 Radiolarians representative of the Ceratoikiscurn regalinodus assemblage collected from Glenrock Station. All scale bars = 100pm.

A Entactiniid gen. et sp. indet. HKUDES 9710219 M Trilonche davidi(Hinde) HKUDES 9710189

B Trilonche echinata(Hinde) AMF96707 N Trilonche davidi (Hinde) HKUDES 9710190

C Trilonche echinata(Hinde) HKUDES 9710180 0 Palaeoscenidium cladophorurn Deflandre HKUDES 9710191

D Trilonche echinata (Hinde) HKUDES 9710 18 1 P Ceratoikiscurn regalinodus Stratford and Aitchison

E Entactiniid gen. et sp. indet. B AMF 96720 HKUDES 9710192

F Trilonche vetusta Hinde HKUDES 9710182 cf. vii/E Q Ceratoikiscurn regalinodus Stratford and Aitchison

G Trilonche echinata (Hinde) HKUDES 9710 183 HKUDES 9710193

R Trilonche sp. B HKUDES 9710194 H Trilonche vetusia ? Hinde HKUDES 9710184

I Trilonche echinata (Hinde) HKUDES 9710185 S Trilonche sp. B HKUDES 9710195

J Trilonche minux (Hinde) HKUDES 9710186 T Trilonche sp. B HKUDES 9710196

K Trilonche minax (Hinde) HKUDES 97101 87 U Trilonche sp. B HKUDES 9710197

L Entactiniid gen. et sp. indet. B HKUDES 9710188


micropaleontology, vol. 45, no. 2, 1999


rnicropaleontology, vol. 45, no. 2, 1999 157

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Remarks: Parallel grooves are observed along the length of the main rods on some specimens.

Range and occurrence: Found at Glenrock in the Circulafomza admissarius assemblage at localities SW-1 [GR 482 9371, SW-2 [GR 486 9321 of Stratford and Aitchison (1996) and the Helenifore pilosidiscus n. sp. assemblage at locality SW-4 [GR 465 9291. C. admissarius is relatively common in both the Gamilaroi and Djungati terranes. Fragments are also known from the Kurosegawa terrane in Japan (Umeda 1994). Lower to Middle Devonian (Emsian-Eifelian) so far as is known.

Circulaforma Aitchison and Stratford n. sp. Plate 3, figures A, U-W

Palaeoscenidium (?) sp. UMEDA 1994, p. 515, pl. 1, fig. 5. Deflantrica (?) sp. UMEDA 1994, p. 515, pl. 1, fig. 6. Circulaforma sp. ASTRATFORD and AITCHISON 1997, p. 246, pl. 2,

figs. D, E; pl. 4, figs. C, D, F-H.

Diagnosis: This taxon possesses two main skeletal rods, dorsal [d.r]and ventral [v .~] , of which the ventral rod is longer. The central ring is very small and often obscured by patagial tissue which covers the entire ring and is flattened normal to the direction in which the caveal ribs extend. Caveal ribs are prominent on the dorsal surface and occur evenly on either side of the central ring. Ribs are initially horizontal, curving ventrally towards the tip. The ribs are approximately parallel. Aventral subsidiary rod may also be seen on either side of the patagial ring.

Remarks: Specimens of this taxon are commonly found broken with only the protruding caveal ribs and the d.r remaining. There may be up to five caveal ribs per side. Because the caveal ribs and flattening of the patagial ring are oriented perpendicular to each other it is difficult to see both these features clearly on a single mounted specimen.

Dimensions: All measurements in (pn) i.d. 66 (34-93), i.v. 97 (47-124), length of body patagium 92 (48-114), width of body long direction 88 (74-106), width of body short direction 30 (23-42), length of caveal rib on one side, 103 (60-145), width of central ring 21 (13-33), length of subsidiary rod 28 (21-34). To- tal of 15 specimens measured.

Etymology: Named after Sir T. W. Edgeworth David the initia- tor of radiolarian studies in Australia.

Type Material: Holotype [HKUDES 9710161 Plate 3, figure U] and paratypes [HKUDES 9710162 Plate 3, figure V; HKUDES 9710163 Plate 3, figure W; HKUDES 9710145 Plate 3, figure A;] from Silver Gully.

Type Locality: Gamilaroi terrane at Silver Gully, NSW, Austra- lia.

Range: Circulaforma davidi n. sp. is found in assemblages 3-6 in the Gamilaroi terrane in the SW Block at Glenrock (Stratford and Aitchison 1998) and at Silver Gully, near Nundle. It also occurs in the Djungati terrane in the New England orogen. Ma- terial synonymous with Circulaforma davidi n. sp. has been reported from Devonian rocks in the Kurosegawa terrane in Japan (Umeda 1994). Lower to Middle Devonian (Emsian-Eifelian) so far as is known.

Genus Helenifore Nazarov and Ormiston 1983 Type species: Helenifore laticlavium Nazarov and Ormiston

Helenifore laticlavium Nazarov and Ormiston 1983 Plate 1, figures A, B, L, M; plate 6, figures G and H

Helenifore laticlavium NAZAROV and ORMISTON 1983, p. 464-466, pl. 2, figs. 8,11,13,14. - ISHIGA 1988,p. 82, pl. 1, figs. 1-4. - ISHIGA and LEITCH 1988, p. 46, pl. 1, fig. 6. - ISHIGA et al. 1988, p. 76, fig. 2g. - AITCHISON 1993a, p. 357, pl. 1, fig. 1. - AITCHISON 199313, p. 112, pl. 2, figs. 11,12. - STRATFORD and AITCHISON 1997, p. 246, pl. 1, figs. A-D. - AITCHISON and STRATFORD 1997, p. 385, pl. 1, fig. 15.

Helenifore? laticlavium Nazarov and Ormiston 1983 - SCHMIDT-EFFING 1988, p. 34, pl. 1, fig. 1.

Diagnosis: See Nazarov and Ormiston (1983)

Range and occurrence: Common throughout the NEO occur- ring in the Devonian of the Garnilaroi, Djungati, Yugambal and ~ i @ a iterranes. Also reported from the Gogo Formation (lowest Frasnian) of Western Australia (Nazarov and Ormiston 1983) and the Frankenwald (upper Famennian) in Germany (Schmidt-Effing 1988). Also known from North America, south west China and Japan (J. Aitchison, pers. observations). ?Lower to Upper Devonian (Frasnian). This taxon is found throughout the Gamilaroi terrane in strata of Emsian and younger ages (Stratford and Aitchison 1997) and is known from samples collected at Glenrock, Silver Gully, Bundook, Pigna Barney, Ken- nedy Farm and Yarras Road.

Helenifore pilosidiscus Aitchison and Stratford n. sp. Plate 2, figures A, B, G; plate 3, figure T

Helenifore sp. AITCHISON et al. 1992, p. 42, fig. 6C. Helenifore sp. A AITCHISON and STRATFORD 1997, p. 385-386, pl.

1, fig. 16.

Description: Helenifore pilosidiscus n. sp. is characterised by the extensive development of patagium around the entire outline with no significant indent below the b-rod as seen in Helenifore laticlavium. The central i-rod rod is separated into two main spines, apical ( i d . ) and basal (i.v.)joined across the small circular, central ring by a thin rod which is often incom- plete. The b-rod is mostly covered by patagial tissue, only the b.a. section of this rod appears to be developed. All spines have a similar thickness. Patagial tissue is dense and flattened with an ovate outline. As in H. laticlavium, the small central portion of the i-rod is often absent.

Remarks: H. pilosidiscus n. sp. is characterised by the extensive development of patagium around the entire outline with no significant indentation below the b-rod as seen in Helenifore laticlavium. This new species of Helenifore is significantly larger than H. laticlavium. Minor outgrowths of patagium may protrude from the ventral portion of the patagial ring, parallel to i.v.

Dimensions: All measurements in (pm). i.d. 104 (93-110), c.r 42 (31-53), i.v. 103 (97-109), b.a. 128 (115-141), body length, 179 (162-206), maximum body width, 173 (136-198). Total of 10 specimens measured.

Etymology: L. pilosus, shaggy; discus, a disk; refemng to the extensive development of patagial tissue.

Type Material: Holotype [AMF96672 Plate 2, figure A] and paratypes [AMF96674 Plate 2, figure B; HKUDES9710160 Plate 3, figure TI from Silver Gully.

Type Locality: Silver Gully, near Nundle, NSW, Australia. 1983

Range and occurrence: H.pilosidiscus n. sp. occurs in the SW Block at Glenrock and at Silver Gully. This species is also found in the Djungati terrane north of Gl~nrock (Aitchison, et al. 1992a) Middle Devonian, possibly mid to upper Eifelian so far as is known.

Genus Protoholoeciscus Aitchison 1993a Type Species: Protoholoeciscus hindea (Aitchison 1993a)

Remarks: In a major work on uppermost Devonian to Lower Carboniferous radiolarians Schwartzapfel and Holdsworth (1996) proposed inclusion of the genus Protoholoeciscus Aitchison within the genus Glanta which they emended. Nota- bly this was done without recourse to examination of the original type material. We disagree markedly with this approach and do not accept their taxonomic revision. The taxa, figured by Schwartzapfel and Holdsworth (1996) and placed within their emended ~ l a n t aare quite unlike any of t h e ~ o w e r to ?Middle Devonian material originally described by Wakamatsu et al. (1990). Protoholoeciscus Aitchison is distinctive and we consider that it warrants retention as a separate genus. It also differs from any of the taxa, figured by Schwartzapfel and Holdsworth (1996). We consider this classification best handled by establishment of a new genus for the material of Schwartzapfel and Holdsworth (1996) which, so far as is known, is restricted to the Lower Carboniferous and does not exhibit any known phylogenetic links to the older Middle Devonian taxa discussed herein. Furthermore, to emphasise major differences in the structure of Glanta Wakamatsu et al. (1990) and Protoholoeciscus Aitchison we suggest emendment of the diagnosis of Glanta to recognise that this genus is characterised by specimens with perforate or porous shells. We understand publication of such an emendment is currently in progress (M. Umeda pers. comm.)

Protoholoeciscus hindea Aitchison 1993 Plate 3, figures P-R

Albaillellaria AITCHISON et al. 1992, p. 42, figs. 60-R. Protoholoeciscus hindea AITCHISON 1993, p. 362, pl. I, fig. 3. -

STRATFORD and AITCHISON 1997, p. 246, pl. 4A, B.

Diagnosis: (see Aitchison 1993a, p. 362)

Remarks: May have a small series of nodes on the lower i.v. and b.v. sections of the main rods.

Range and occurrence: Protoholoeciscus hindea is found at Sil- ver Gully, at locality SW-5 [GR 469 9101 at Glenrock, and elsewhere in the Gamilaroi and Djungati terranes. The age range of this species appears to be Middle Devonian (Eifelian to lower Givetian) so far as is known. Protoholoeciscus is also known from Eifelian strata of the Kurosegawa terrane in Japan (Umeda and Yamagiwa, 1997).

Incertae sedis Family PALAEOSCENIDIIDAE Riedel 1967, emend. Holdsworth 1977, Goodbody 1982, Furutani 1983, Goodbody

Genus Palaeoscenidium Deflandre 1953 emend. Goodbody 1986 Type species: Palaeoscenidium cladophorum Deflandre 1953

Palaeoscenidium cladophorum Deflandre group Plate 5, figure 0 ; plate 6, figures F and J

Palaeoscenidium cladophorum DEFLANDRE 1953, p. 408, fig. 308. -DEFLANDRE 1960, p. 214, pl. 1, fig. 21. -FOREMAN 1963, p. 302,


pl. 8, fig. 10,pl. 9.,fig. 6. - HOLDSWORTH 1973,p. 128, pl. 1, fig. 19. - NAZAROV 1975, p. 96, pl. 13, figs. 4, 5; pl. 14, figs. 5, 6. -NAZAROV et al. 1982, p. 172, figs. 5 D-F. - NAZAROV and ORMIS- TON 1983, p. 465, 2, figs. 6.7. - ISHIGA et al. 1987,p, 299, PI. 1, figs. 1-10. - AITCHISON 1993, p. 121, pl. 1, figs. 15-17 19; pl. 2, figs. 17,20. - STRATFORD and AITCHISON 1997, p. 247, pl. 2, fig. 0 .

Remarks: There is a large degree of intraspecific variation in P. cladophorum with many minor variations on the same structural framework (Nazarov and Ormiston 1983). Work on extremely well-preserved Frasnian material from the Gogo Formation (Aitchison 1993b) has seen eight new species of Palaeo-scenidium described. It is likely that in less well-preserved material only a few of the simpler of these new species will be recognisable. Specimens ascribed to Palaeoscenidium cladophorum Deflandre group were observed in many residues of samples collected from throughout the Gamilaroi terrane.

Range and occurrence: This is a cosmopolitan species group reported from Australia, Europe, Asia minor, North America, the former Soviet Union and Japan. Middle Devonian to Lower Carboniferous so far as is known. The biostratigraphic potential of Palaeosceniids remains to be established.

ACKNOWLEDGMENTS We gratefully acknowledge the financial assistance of the Aus- tralian Research Council. We would also like to thank Bruce McNaughton for access on Glenrock Station, and NSW Water Management at Chaffey Dam for allowing access to Silver Gully localities. Thanks also to Masaki Umeda for help with de- termining the age ranges for some species and Paula Noble for discussions and her helpful review of the manuscript. We thank our Garnilaroi terrane colleagues (John Talent, Ruth Mawson, Terry Furey-Greig and others at Macquarie University and Evan Leitch at UTS) for assistance with conodontology and discussions on the evolution of the Garnilaroi terrane.

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