late emsian rugose corals of the mount podge area, burdekin basin, north queensland

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Late Emsian rugose corals of theMount Podge area, Burdekin Basin,north QueenslandYong-Yi Zhen a ba Department of Earth Sciences , University of Queensland ,Queensland, 4072b Centre for Ecostratigraphy and Palaeobiology, School of EarthSciences , Macquarie University , N.S.W., 2109Published online: 27 Nov 2008.

To cite this article: Yong-Yi Zhen (1995) Late Emsian rugose corals of the Mount Podge area,Burdekin Basin, north Queensland, Alcheringa: An Australasian Journal of Palaeontology, 19:3,193-234, DOI: 10.1080/03115519508619506

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Late Emsian rugose corals of the Mount Podge area, Burdekin Basin, north Queensland

YONG-YI ZHEN

zu~q, Y.-Y., 1995:10:31. Late Emslan mgos¢ corals of the Mount Podge area, Burdekin Basin, north Queensland. Alcheringa 19, 193-234. ISSN 0311-5518. The Laronna Formation and Mount Podge Limestone ate defined for the lower conglomexates

and m i ~ sandstones, and the upper ¢oralline limestones exposed in the Mount Podge area, north Queensland. Acanthophyllum (Acanthophyilum) clermontense -Protomacgeea fauna from these two units (mainly from the Mount Podge Limestone) in the area is of late Emsian age, and comparable with other Emsian to early Effefian coral faunas from Queensland and New South Wales. FiReen species belonging to 14 genera (one genus and 7 species new) are described from the Mount Podge Limestone, including Microplasma ronense (Mansuy 1913), Lekanophyllum laroonaense sp. nov., Sanidophyilum sp., Tabulophyllum carinatum sp. nov., Carlinastraea callosa sp. nov., Australophyllum sp., Xystriphyllum of. dunstani (Etheridge 1911), X. of. magnum Hill 1942a, Tatmyrophyllum pedderi sp. nov., Loroonaphyllum jacki gen. et sp. nov., Acanthophyllum (Acanthophyllum) ciermontense (Etheridge 1911), Disphyllum (Disphyilum) paracouvinense sp. nov., Phill~osastrea sp. of. macuiosa Hill 1942a, Protomacgeea minor sp. nov. and Thamnophyllum sp., and one new species Gaynaphyllum runningense sp. nov. from the very top of the underlying Latoona Fomuttio~.

Yong-yi Zhen, Department of Earth Sciences, University of QueenMand, QueenMand 4072;present address Centre for Ecostratigraphy and Palaeobiology, School of Earth Sciences, Macquarie Uni- versity, N.S.W. 2109; received 18 May 1993.

Keywofds: late Fsn~ian, Rugnsa, new tsxa, distribution, Mount Podge Limestone, Laroona Focmation, biostraligraphy, north Queensland.

RECENT rugose coral biostratigraphic and biofacies studies of the Devonian Fanning River Group of north Queensland (Zhen, 1991) dem- 0infrared that the two units exposed in the Mount Podge area, which Wyatt et al. (1970) mapped as the Big Bend Arkose and the Burdekin For- marion respectively, were deposited in an earlier transgression of late Emsian age. However, the major part of the Burdekin Basin sedimentation did not commence until a later transgression of the latest Eifelian to middle Givetian age, which is represented by the Big Bend Arkose, the Burdekin Formation and the Cultivation Gully Formation. Hence the Laroona Formation and the Mount Podge Limestone were proposed for the lower conglomerate and sandstone unit and the upper limestone unit in the Mount Podge area

0311/5518/95/020193-43 $3.00 ©AAP

with a total thickness up to 450 m (Zhen, 1991; Zhen et al., 1993).

Stratigraphically, the Laroona Formation lies nonconformably on the Proterozoic Argentine Metamorphics (Fig. 1). At the type locality (Run- ning Creek section, UQL5447, Fig. 1), it con- sists of 146.5 m of conglomerate, micaceous sandstone and shale with calcareous sandstone and sandy, coral-bearing limestone nodules or interlayers near the top. The basal conglomerate abuts a Carboniferous rhyolite sill, with the Argentine Metamorphics cropping out about 200 m further down the creek, but the direct contact relationship is not clear. On the Mount Podge section (UQL5445), the upper exposed part con- sists of 60 m of conglomerate and medium to very coarse-grained micaceous sandstone. Fur- filer southeast (UQL5543), about 150 m of conglomerate and free to coarse-grained micaceous

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194 Y.-Y. ZHEN ALCHERINGA

sandstone of the Laroona Formation is generally poorly exposed in a gully, but the gradational boundary with the Mount Podge Limestone is well exposed. The Mount Podge Limestone con- sists of dominantly packstones and calcirudites 156 m thick on the type section (UQL5445) and up to 300 m thick of packstones, bindstones, grainstones and calcirudites with coral and stromatoporoid build-ups near the top of the Run- ning Creek section (UQL5447), and contains abundant corals and stromatoporoids. It is overlain by the Keelbottom Group of the Late Devonian to Early Carboniferous (Fig. 1).

Rugose corals, brachiopods and conodonts from the two formations indicate that this late

Emsian transgression extended to the Mount Podge area from the north via the Broken River Province. The Mount Podge area, now fault bounded, may have been a southern extension of the Broken River Province 'left behind' following left-lateral movement along the Clarke River Fault Zone, during the Middle to Late Devonian (Lang et al., 1990). This transgression is also represented by sedimentation of the Conjuboy Formation of the Georgetown Inlier and the lower part of the Broken River Group (the Lomandra Limestone or equivalent) of the Broken River Province, north Queensland, the Douglas Creek Limestone of central Queensland, and the Mount Frome Limestone of New South

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Fig. 1. Locality map with collecting siles oflhe Mount Podge area, north Queensland.

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ALCHERINGA DEVONIAN RUGOSE CORALS 195

Wales, but it did not extend to the major part of the Burdekin Basin (Fig. 2).

Age of the fauna

Sixteen species of rugose corals, including one new genus and 8 new species, are described herein from the top part of the Laroona Forma- tion and the lower and middle parts of the Mount Podge Limestone (Fig. 3). This fauna from the Mount Podge area, here referred to as the Acanthophyllum (Acanthophyllum) clermontense-Protomacgeea fauna, is readily comparable with other Emsian to early Eifelian coral faunas of Queensland and New South Wales (Fig. 2).

Conodonts, including the important index conodont species Polygnathus serotinus, have dated the Mount Podge Acanthophyllum (Acanthophyllum) clermontense-Protomacgeea coral fauna as late Emsian in age. In the type section, the whole limestone formation might lie within the serotinus conodont Zone (R. Mawson, 1991, pers. comm.). One sample from the Carlinastraea callosa beds yielded lcriodus cf. corniger, Polygnathus linguiformis linguiformis and P linguiformis bul(yncla', indicating an age of latest Emsian or early Eifelian (R. Mawson, 1991, pers. comm.). In the Running Creek section (UQL5447), P serotinus first occurs at the top of the Laroona Formation and extends into the lower part of the Mount Podge Limestone, but conodonts from the very top of the limestone sequence exposed in the Running Creek indicate an early Givetian age which may represent a separate transgression (Talent & Mawson, 1994). Alternatively, the late Emsian transgression in the Mount Podge area might extend continuously to the early Givetian. However, further detailed biostratigraphical studies are needed before this can be certain. Corals from the upper part of the Mount Podge Limestone on this section indicate that the formation exposed there may extend into the Eifelian.

Zdimir, which is abundant in the Mount Podge Limestone, is a distinctive late Emsian to early Eifelian brachiopod, widely distributed in the

former Soviet Union, south China, Europe and eastern Australia.

The type material of Acanthophyllum (A.) clermontense is from the Douglas Creek Lime- stone (Emsian), Queensland (Jell & Hill, 1970). This species has also been recorded from the Conjuboy Formation (late Emsian) and Broken River Group (late Emsian part) of north Queens- land; Dunstable Beds (Emsian) of central Queensland (Fordham, 1976); and doubtfully from the Garra Formation (Pragian to early Emsian) of New South Wales (Strusz, 1966). Overseas, it has been reported from the Yangmaba Formation (late Emsian part) of Sichuan, southwestern China (He, 1978).

Protomacgeea occurs in the Phillipsastrea fauna (late Emsian to early Eifelian) of the Bro- ken River Group, in the Conjuboy Formation (Emsian), and in limestone pebbles at the base of the Drummond Group (Jell & Hill, 1969), Queensland, and in the Mount Frome Limestone (late Emsian to early Eifelian) of New South Wales (Wright, 1966).

Taimyrophyllum pedderi is close to T. grande (Dun, 1918) from the Loomberah Limestone (latest Emsian to early Eifelian) of New South Wales, a fauna (Pedder, 1967) which closely re- sembles the fauna of the Mount Podge Lime- stone. Zdimir has also been reported from the Loomberah Limestone. The type of Xystriphyl- lure magnum Hill 1942a is from the middle of the Sulcor Limestone (late Emsian to early Eifelian), where it is associated with Trapezophyllum? coulteri, Sulcorphyllum brownae, Phillipsastrea linearis, P. maculosa, Thamnophyllum sp., Tipheophyllum sp., Pseudamplexus sp. and Calceola (Pedder, 1967). Xystriphyllum magnum shows a close resemblance to X. densum Yu & Kuang 1982 from the top of the Guitang Member (early Eifelian) Beiliu Formation of Guangxi where it is associated with Zdimir, Tabulophyllum beiliuense, T. undulatum, Dohmophyilum guitangense and Calceola sandalina.

Carlinastraea has a wide geographical distribution and has been reported Horn Europe, North America, China and eastern Australia with an age

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/ ~ . 2. Correlation of the AeanthophyUum (Aoanthophyllum) clermontense-Protomacgeea fauna in Queens- land and Now South Wales. * bearingA. (.4.) elermontense; + bearing Protomacgeea.

ranging from Late Silurian to early Emsian. It is very common in the Lochkovian and Pragian in eastern Australia Oru & Jell, 1990). Conodonts indicating a latest Emsian age (R.Mawson, 1991, pers. comm.) have been recovered from the crinoidal packstones of the Carlinastraea callosa beds of the Mount Podge Limestone. This has considerably extended the range of Carlinas- traea.

Elements of Endophyllidae, like Tabulophyl- lure Fenton & Fenton 1924, made their first occurrence in the Pragian of eastern Australia and became common in the Emsian. The fasciculate forms tentatively referred here to Tabulophyllum and the cerioid forms of Endophyllum Milne- Edwards & Haime 1851, are very common in the Emsian ofeastemAustralia; elsewhere these forms mainly occur in the Middle or even Late Devonian. Sanidophyllum Etheridge 1899 is endemic to eastern Australia, being previously reported only from the Middle Devonian of New South Wales and Queensland. It is found in as- sociation with Carlinastraea in the Mount Podge Limestone.

Lekanophyllum laroonaense shows a close

resemblance to L. cornubovis (Etheridge 1899) from the Eifelian Moore Creek Limestone of New South Wales. Microplasma ronense (Mansuy 1913) also recorded from the slightly older Garra Formation and the late Emsian Sutchers Creek Formation, New South Wales.

Systematic palaeontology The suprageneric classification used in this paper follows the classification of Hill (198 I). Only the basionyms are given for the described genera because of limited space. The terminology follows that of Hill (1981 ) except the few terms discussed previously by the author (Zhen, 1994). All specimens and thin sections used in this study are housed in the Geology Museum, University of Queensland. They are catalogued in the fossil catalogues and carry F numbers, and all material from a given locality carries the same L numbers referred to in the Fossil Locality Cata- logue (see Appendix).

The following abbreviations are used in the systematic descriptions: Dc, corallite diameter;

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Dr, tabularium diameter; Dt/Dc, ratio of the tabularial diameter to corallite diameter; Nd, number of rows of dissepiments and/or presepiments; Ns, total number of septa; TS, Iransverse section; LS, longitudinal section; OD, original designation; and SD, subsequent designation.'

Repositories of type and other specimens are indicated by the following prefixes to catalogue numbers: GSQ, Geological Survey of Queens- land, Brisbane; UQ, The University of Queens- land, Brisbane; and SU, University of Sydney, Sydney.

Subclass RUGOSA Milne-Edwards & Haime 1850

Order CYSTIPHYLLIDA Nicholson 1889

Family CYSTIPHYLLIDAE Milne-Edwards & Haime 1850

Mieroplasma Dybowski 1874

1874 Microplasma Dybowski, p.508, partim. 1974 Nardoplasma Spasskiy & Kravtsov in Spasskiy et

al., p.171. 1978 Paeudomicroplasma (Phaceiloplasma) Kong in Kong

& Hmmg, p. 142.

Type species (SD; Wedekind, 1927). Micro- plasma gotlandicum Dybowski 1874, p.508, pl.5, fig.5a-d; ?Slite Beds, (Silurian) Stora Karlso, Isle of Gotland, Sweden.

Diagnosis. Fasciculate corallum with peripheral increase; septa represented by small, trabecular spines projecting from narrow peripheral stereozone; septal crusts absent; septal crests rare or nonexistent.

Remarks. McLean (1976) included in Micro- plasma both colonial and solitary forms. He suggested that Microplasma differs from Silurian Cystiphyllum and Devonian Cystiphylloides in having more pronounced septal development at the corallite periphery instead of random distribution as in Cystiphyllum and the greater development of septal crusts on the dissepiments of Cystiphylloides. Hill (1981) restricted Micro- plasma to fasciculate forms and placed the

Devonian solitary species, previously included in Microplasma by McLean (1976), in Zonophyl- lum Wedekind, with which she merged Pseudomicroplasma. The present author follows the concept of Hill (1981) in retaining Micro- plasma for those Silurian and Devonian fasciculate forms with short septal spines confined to a thin peripheral stereozone, and including those solitary species previously referred to Microplasma by McLean (1976) in Pseudo- microplasma.

Birenheide (1974) confined Microplasma only to Silurian forms and included Devonian forms in Cystiphylloides Chapman 1893, which he regarded as a subgenus of Mesophyllum Schliiter 1889. Based on the revisions by McLean (1976) and Hill (1981), Cystiphylioides can be distinguished ~om Microplasma by its weakly developed septal crusts.

Kong (in Kong & Huang, 1978) proposed Pseudomicroplasma ( Phacelloplasma ) to include those Devonian fasciculate forms with the internal structure of Pseudomicroplasma, and suggested that P. (Phacelloplasma) could be distinguished from Silurian Microplasma by its monacanthine trabeculae rather than the holacanths of Microplasma. As holacanths are considered as the diagenetic product of

ALITIES

Micropla~ma ronense Lekanophyllum laroonaense sp. nov. Sanidophyllum sp. Tabulophyllum carinatum sp. nov. Carlinastraea callosa sp. nov. Australophyllum sp. Gaynaphyllum runningense sp. nov. Xystriphyllum cf. dunstani Xystriphyllum cf. magnum Taimyrophyllumpedderi sp. nov. Laroonaphyllum jacki gen. et sp. nov. Acanthophyllum (A.) clermontense Disphyllum (D.) paracouvinense sp. nov. Phillipsastrea sp. cf. maculosa Protomacgeea minor sp. nov. Thamnophyllum sp. *

F/g. 3. Distribution of the described rugose corals from the Mount Podge area, north Queensland.

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recrystallization of both monacanths and rhabdacanths, and since all material available to the present author is with obscured microstmc- tare, it is then regarded as synonymous with Microplasma.

Mieroplasma ronense 0Vlansuy 1913) (Fig. 4A- F)

1913 Michelinia ronensis Mansuy, p.10, pl.l , fig.12a-¢. ?1942b 'Cystiphyilum' conjunc~an Hill, p. 185, pl.5, figs

3a, 6. 1961 Microplasma roncn.ff.¢; Fontaine, p.198, pi.32,

fig. 1, pl.33, fig.9; pl.34, fig.6. 71962 M/crop/asma sp. el. M. devonica; Philip, p.192,

pl.29, figs 2, 6. 1963 Microplasma conjuncture; Strusz, p.224, p[.49, figs

1-3. 1966 Plasmophyll~ (Plasmophyllwn) ronens/s; Wright,

p.333, pi.55, figs 1, 2.

Holotype. Specimen figured by Mansuy, 1913, pl.1, fig.12a-c; Emsian or early Eifelian, Ron Region, Vietnam.

Material. UQF78594 and UQF78576-80 from UQL5444; UQF78667, UQF78670 and UQF78675 from UQL5447-2.

Description. Coralla phaceloid. Sinuous cylindrical corallites are slender and rather widely spaced with a diameter ranging from 6 to 15 ram, and distorted when in contact; adult corallites rarely seen in contact with each other. Periph- eral stereozone thin, 0.2 to 0.6 mm thick; increase lateral; calice shallow with flat bottom. Septal spines well developed of wedge-like holacanths, projecting inwards to form denticulate inner surface of peripheral stereozone, or most oRen extending deep into dissepimentarium, up to 3 mm long with fine grains on the surface. Septal spines thin and sinuous in dissepimentarium; in a corallite of 10.5 mm diameter, 56 septal spines counted; in some corallites, septal spines differ- entiated into two orders, minor about half to two- thirds as long as the major. Dissepiments small and concentric in transverse sections.

Dissepimentarium relatively narrow; dissepiments in two to six rows, hemispherical to greatly elongated, variable in size, typically smaller and hemispherical near wall, larger and

elongated near inner margin of dissepimentarium, and steeply inclined adaxially. Tabularium about half width ofcorallite; tabellae larger, flat-floored, with peripheral series slightly adaxiaUy inclined; boundary between dissepimentarium and tabularium rather distinctive.

Remarks. Wright (1966) regarded Microplasma conjuncture (Hill 1942b) as a junior synonym of M. ronense. The type material of M. conjuncture from the Garra Formation (Pragian to early Emsian) of Wellington, New South Wales resem- bles the present material with respect to corallite size and features seen in longitudinal section, but has thinner walls and rarer occurrence of septal spines. Strusz (1963) discussed variation in M. conjuncture from various localities in New South Wales. He noted that material from the Catombal Bioherm differs from the topotypes in having a wider peripheral stereozone, up to 1 mm thick. The figured specimen (Strusz, 1963, pl. 49, fig. 2) from locality Gg-1 shows well-developed short, but prominent, septal spines. The type material of M. ronense is from the Emsian or lowermost Eifelian of the Ron region, Vietnam. It is also reported from ?Eifelian beds of Laos (Fontaine, 1961). The material from Laos has more closely spaced corallitcs, which are often subpolygonal in cross section.

This species is very common in locality UQL5A.A.s., and preserved as in situ colonies up to 40 cm tall and 50 cm diameter. More than 20 specimens have been collected and they all consistently show well-developed, long septal spines in adult corallites. A few single corallites from UQL5447-2, have the same internal features but thinner outer walls and rarely occuring, much shorter, septal spines.

The specimen from the Sutchers Creek For- marion (late Emsian) of New South Wales referred to Microplasma ronense by Wright (1966) is very close to the present material, except for its more arched upper surface of the tabulae, as compared with the type material. However, those arched and more or less horizontally floored tabulae are common too in the Mount Podge material.

The specimen figured as Cystiphylioides

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F

F/g. 4. A-F, Microplasma ronense (Mansuy 1913). A, 13, UQF78576 from UQL5444, x2. A, TS; B, LS. C, D, UQF78580 from UQL5444, x2. C, 'IS; D, LS. E, F, UQF78578 from UQL5444, x2. E, TS; F, LS. G, H, 8anidophyllum sp. UQF78726 from UQL5459, xl.5. G, TS; H, I~.

fruticosum (Nicholson 1875) by McLean (1976) possibly from the Bois Blanc Formation (late Emsian) of Ontario has similar internal features to Microplasma ronense, but the Canadian material has slightly smaller mean corallite diameters, thinner outer walls and more widely spaced tabulae.

Family DIGONOPHYLLIDAE Wedekind 1923

Lekanophyllum Wedekind 1923

1922b Mesophyilloides Wedekind, p.5 l,partt'm. 1923 Lekanophyllum Wedekind, p.29. 1925 Hemicystiphyllum Wedvkind, p.28. 1925 Atelophyllum Wedekind, p.37.

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1925 Diatytophyllum Amanshauser in Wedekind, p.40, 42.

1960 Pseudodigonophyilum Spasskiy, p.39. 1960 Pseudodigoaophyilum Spasskiy/n Bulvanker et al.,

p.236. 71963 Neozonophyllum Ulitina, p.5, nora. imperf. 1964 Asperophyilum Spasskiy i , Dubatolov & Spasskiy,

p.132, 133. 71972.4culeatophyllum Zhavoronkova in Strehaikov &

Zhavoronlmva, p.94. 1974 Scissoplasma Spasskiy & Kravtsov in Spasskiy et

a/., p.171. 1978 Dialithophyllum (Protodialithophyilum) Kong in

KoGg & Huang, p.158, 159.

Type species (SD; Lang, Smith & Thomas 1940). Lekanophyllum punctatum Wedekind 1924; p.30, 34, figs 36-38; Junkerberg Formation, Eifelian, Germany.

Diagnosis: Solitary or fasciculate genus of digonophyllinid corals; septal crests typically well developed throughout, especially in ephebic stages where they commonly form complete septa, some dilation of septa occurring in ncanic stages, but never to the extent of producing fused crests that entirely fill the corallite; in ephebic stages, axial dilation of the trabeculae insignifi- cant, discrete peripheral carinae either very rare or absent (after Pedder & McLean, 1982, p.75).

Remarks. Mcl_~.an (1976) and Pedder & McLean (1982) regarded Dialythophyllum as synonymous with Lekanophyllum and suggested that the only difference between Dialythophyllum and Lekanophyllum was in the former having more dilated major septa, which may extend to the corallite wall. However, this is also the case in some species of Lekanophyllum, where presepiments are rare. Hill (1981) took Dialytophyllum as a separate subgenus of Mesophyllum, and characterized it by its thick and laminar major septa present peripherally and buttressed by lateral dissepiments.

Hill (1981) considered Zonodigonophyllum Vollbrecht 1926 and Lekanophyllum as separate subgenera of Mesophyllum. She suggested that Zonodigonophyllum could be distinguished from Lekanophyllum by its weak occurrence of key- hole outline in the calical pit and prominent septal crusts, especially in the early stage. McLean

(1976) and Pedder & McLean (1982) doubtfully regarded Zonodigonophyllum as a junior synonym of Lekanophyllum. They regarded the type species as a doubtful species of Lekanophyllum and referred all other species originally described by Vollhrecht (1926) to Digonophyllum.

Kong (in Kong & Huang, 1978) proposed Protodialithophyllum as a subgenus of Dialitho- phyllum to include those intermediate forms between Atelophyllum, Zonodigonophyllura and Dialythophyllum. He defined it as being like Dialythophyllum but containning thin septa only, with well-developed lateral dissepiments. This is well within the defmition of Lekanophyllum.

Mesophyllum Schliiter 1889 differs from Lekanophyllum in its extensive development of peripherally discrete carinae, typically forming crossbars.

Lekanophyllum laroonaense sp. nov. (Fig. 5A- F)

Etymology. After Laroona Homestead, to the south of the type locality.

Material. Holotype UQF78678 from UQL5447- 3, lower part of Mount Podge Limestone, Mount Podge area. Paratypes: UQF78659 from UQL54.47-1, top part of Laroona Formation; UQF78661-3 from UQL5447-2; UQF78679-80, UQF78685 and UQF78687 from UQL5447-3; UQF78692-3 from UQL5447-6; and UQF78589 from UQL5445-3, Mount Podge Limestone.

Diagnosis. Large Lekanophyllum with diameter to 65 ram; major septa long, discontinuous peripherally, and often carinate; minor septa present as fragmentary low septal crests or absent; dissepimentarium wide, consisting of outer series of presepiments and inner series of dissepiments.

Description. Large solitary trochoid or ceratoid corallum, subcylindrical in late growth stage; calice inverted conical, with a depth half the coraUite diameter, maximum diameter up to 65 mm. Outer wall with smooth surface, without septal grooves and interseptal ridges; it is 0.7 to 1 mm thick and composed of dilated septal bases

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t~g. 5. Lekanophyllum laroonaense sp. nov. A-C, holotype, UQF78678 from UQL5447-3. A, TS, xl.5; B, I~, xl.5; C, TS, x2.5. D, E, paratype, UQF78687 from UQL5447-3, xl.5. D, TS; E, LS. F, paratype, UQF78659 from UQL5447-1, I~, xl.5.

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and sclerenchyme fibres perpendicular to the epitheca. Sepia in two orders, mouacanths, peripherally discrete forming zigzag cariuae and more or less retiform, cross-bar carinae absent or only sporadically occurring in some larger specimens; sepia (39-65)x2 in number as shown in Table 1. Major sepia nearly reaching axis, unequal in length, leaving a small space at the cenC'e, about 3 to 8 mm wide; cardinal fossula indistinct. Minor sepia half to three-fifths as long as major, much more degenerate, represented by trabecular fragments or low sepial crests, sometimes absent. Lateral dissepiments occasionally occur.

Dissepimentarium wide, in the holotype consisting of 17 to 19 rows of steeply inclined, hemispherical to moderately elongate dissepiments; outer dissepimentarium consisting of four to eight rows of larger, hemispherical, adaxially less steeply inclined presepiments. The inner series of dissepiments are smaller, often moderately elongated, and more steeply inclined. Tabularium narrow, 1.5 mm wide in the holotype (De 47 ram), without distinctive boundary between Iabularium and dissepimentarium; iabellae small, adaxially inclined.

Remarks. Compared with the type material of Lekanophyllum cornubovis (Etheridge 1899) from Moore Creek Limestone, Tamworth, N.S.W. (also see Hill, 1942a), the new species from Mount Podge shows a slightly wider Iabularium, less interrupted major sepia, especially peripherally. Neither Hill's specimen (1942a) nor the type material figured by Etheridge (1899) shows any cross-bar carinae.

Table I. Lekanophyllum laroonaense sp. nov. nw, uuremeats.

Ns Nd Dc (ram)

UQF78678 UQF78679 UQF78662 UQF78663 UQF78661 UQF78680 UQF78659 UQF78692 UQF78693

56x2 17-19 47 54x2 20 52 45x2 15 45 51x2 12 42 65x2 20-22 50 40x2 - 31 39x2 17-18 32 54x2 17-22 51

17-19 65

Cross-bar carinae are commonly absent from L. laroonaense, but very rare, ill-defmed cross-bar carinae occur in some larger specimens, although zigzag carinae are common in most specimens.

Lekanophyllum alhambraens e Merriam 1973b from the upper part (coral zone F and G) of the Nevada Formation has numerous thickened sepia and better developed minor sepia. The major sepia typically reach the axis or are slightly ro- Iated in the centre of the corallite.

Order STAURIIDA Verrill 1865

Family ENDOPHYLLIDAE Torley 1933

Sanidophyllum Etheridge 1899

1899 Sanidophyllum Etheridge, p.154.

Type species (OD). Sanidophyllum davidi Etheridge, 1899, p.154, pl. 16; pl.17, fig. 1; pl. 19, fig.7; pl.20, figs 4, 5; pl.38, fig.2; Timor Lime- stone, late Eifelian part, New South Wales.

Diagnosis. Phaceloid and briefly cerioid when periodically developed calicular expansions of each eorallite meeting the similar expansions from its neighbours; sepia in two orders, radially arranged; minor sepia commonly short, not extending far beyond narrow peripheral stereozone; tabulae flattened domes that may sag axially with edges curved into a peripheral trough; no dissepiments.

Remarks. Sanidophyllum is characterized by its periodically developed calicular expansions. Rims ofcalicular expansions regularly join their neighbouring expansions at the same level to form briefly a cerioid form. This speciacular fea- ture of the genus would have made it a success- ful competitor in very shallow muddy environ- ments with unstable substrates, such as mud fiats. Calicular expansions would have counteracted the effect of sedimentation with a high ratio of terrigenous material, and these were probably also a rather effective way to protect the young from the unfavourable muddy conditions.

Sanidophyllum, endemic to eastern Australia, was previously recorded only from the Middle Devonian.

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Sanidophyllum sp. (Fig. ziG, H)

Material. UQF78726 and UQF78728 from UQL5459.

Description. Large subcerioid coraUa with diameters exceeding 30 cm.Corallites closely spaced, rounded or subpolygonal, often in contact with each other laterally; lateral, non-parricidal increase, new offsets arising just inside the calicular expansions; adult coraUites with a diameter ranging from 14 to 24 mm. Calicular expansions generally less than 8 mm broad, extending outwards and upwards with a high angle to horizontal, and the margin strongly upwardly deflected to form walls up to 8 mm high; in cross section, corallites often subpolygonal when connecting with calicular expansions of neighbouring corallites; calicular expansions 11 to 13 mm apart vertically. Septa in two orders, (24-28)x2 in number, strongly dilated peripherally, forming a prominent peripheral stereozone of 1 to 1.8 mm wide, thin and smooth beyond peripheral stereozone; major septa nearly reaching axis or somewhat withdrawn from the cenae, leaving an open space 4 to 5 nun wide; minor septa one-third as long as radius when fully developed, 2 to 4 mm in length, unequal, some reduced to short ridges at periphery. Tabulae complete or incomplete, domed with flat tops, slightly concave or convex at centre, and with sharply down-turned margins, vertically 10-14 tabulae per 10 nun.

Remarks. Sanidophyllum etheridgei Pedder (in Pedder, Jackson & Ellenor, 1970) has very short (less than 0.5 mm) or even entirely suppressed minor septa, and a narrower peripheral stereozone. S. colligatum (Etheridge 1920) shows some resemblance to the present species, but has larger corallites (De up to 30 ram), numerous scpta (Ns up to 40x2) and nearly horizontally extended calicular expansions (as figured by Pedder in Pedder, Jackson & Ellenor, 1970, pl.21, ~gs l, 2).

Tabulophyllum Fenton & Fenton 1924

1924 Tabulophyilum Fenton & Fenton, p.30,partim. 1928 Apolythophyllum Walthex, p.135.

71937 $inospongophyllum Yoh, p.56. 1939 Diversophyllum Slogs, p.66.

Type species (OD). Tabulophyllum rectum Fenton & Fenton, 1924, p.31, pl.6, figs 8-12; Late Devonian, Hackberry Group, Cerro Gordo County, Iowa.

Diagnosis. Solitary, major septa long, smooth or weakly carinate, axial ends with unequal cur- vature or somewhat withdrawn from axis; mi- nor septa reduced to low ridges on the dissepiments and presepiments; presepiments forming a peripheral zone of variable width; neither a distinctive fossula, nor cardinal or counter septa of distinctive length developed in adult stages; tabularium wide, tabularial floors low to flat-topped domes with edges curved into somewhat asymmetrical peripheral trough.

Remarks. All the loosely fasciculate species with inverted conical corallites, previously referred to Tabulophyllum by various authors, might be better excluded from this genus. Sinospongophyllum might prove to be a valid name for those fasciculate species. Wang (1948) described the type species of Sinospongophyllum as mostly simple, sometimes weakly compound. Jia (in Jia et al., 1977) also remarked that '...it is still ques- tionable whether those forms of endophyUoids from the Middle Devonian of south China are solitary or segments of loosely colonial forms'. Fontaine (1966) described and figured specimens ofS. planotabulatum Yoh 1937, from the Mid- dle Devonian of eastern Yunnan, which are quite large phaceloid coralla, with ceratoid to trochoid corallites. These corallites are associated with frequent peripheral, non-parricidal budding. They show a close resemblance to the material from the Emsian and Eifelian of eastern Aus- tralia. Unfortunately, the type material of S. planotabulatum is missing and most species included in this genus by Chinese authors are described as solitary. Until the topotypes of S. planotabulatum are re-examined this name is unusable, and tentatively regarded as a junior synonym of Tabulophyllum. McLean & Pedder (1984) recognized that their genus Tarphy-

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phyllum, with T. besti from the Frasnian of western Canada as the type species, could be distinguished from Tabulophyllum by its fasciculate growth form, narrower and imperfectly developed dissepimentarium and well-developed and normally wider peripheral stereozone. Smithphyllum Birenheide 1962 has fasciculate or cerioid growth form of typically small, cylindrical corallites and narrow dissepimentarium of typically one row or even an incomplete row of presepiments and dissepiments.

Kong (in Kong & Huang, 1978) defined Tabulophyllum as small, solitary coralla with stringophylloid trabeculae distinguishable from Sinospongophyllum by smaller corallite size, discontinuous septa, and normally with non-carinate septa. Thus the Tabulophyllum species described by Kong (in Kong & Huang, 1978) should be referred to the stringophylloid Neospongo- phylloides Jia (in Jia et al., 1977), which can be distinguished from Tabulophyllum by its coarse monacanthine trabeculae ofstfingophyUoid type as stated by the original author.

Merriam (1973b) suggested that the Silurian Dokophyllum Wedekind I927 and the Early Devonian Papiliophyllum Stumm 1937 and other genera of Halliidae differ from Tabulophyllum in having a well-developed cardinal fossula. Ac- cording to Stumm's (1937) original description, Papiliophyllum elegantulum Stumm, the type species of Papiliophyllum, could be well distinguished from Tabulophyllum by having a prominent cardinal fossula, strongly dilated septa in cardinal quadrants and without differentiation into the major and minor series. Eurekaphyllum proposed by Stumm (1937) in the same paper has the septa dilated in cardinal quadrants not as strongly as in Papiliophyllum and only weakly developed cardinal fossula. Hill (1956, 1981), Birenheide (1978) and many other authors regarded Eurekaphyllum a junior synonym of Papiliophyllum. Birenheide (1978) expanded the definition and included Tabulophyllum magnum Fenton & Fenton 1924, T. multiseptatum Tsien 1969 and T. mcconnelli (Whiteaves 1898) within Papiliophyllum. None of the three species has septa more dilated in cardinal quadrants, and the

cardinal fossula is only indistinctly developed in T. mcconnelli. According to the original description, only one of the type specimens of T. magnum shows a slight trace of fossula. The type material of Z multiseptatum shows no trace of fossula. The present author follows Stumm (1937) and Hill's (1981) interpretation of Papiliophyllum, retaining only those species with a distinct cardinal fossula and commonly more strongly dilated septa in cardinal quadrants. Those species with only inconstant occurrence of weak cardinal fossula remain here in Tabulophyllum.

Diversophyllum Sloss 1939 was distinguished from Tabulophyllum by its definite and persist- ent minor septa. According to Sloss, presepiments might occur in the adult stage of Diversophyllum, but the holotype and hypotypes figured by Sloss show rare presepiments. Watkins (1959) regarded the genus as synonymous with Tabulophyllum as do Stumm (1962), Tsien (1969), Pitrat (1962), Birenheide (1978) and doubtfully Hill (1981). Merriam (1973b) suggested that Diversophyllum differs from Tabulophyllum in having longer septa reaching the axis, more arched and irregular tabulae, and less tendency to produce presepiments. The hypotypes of the type species were well-figured by Stumm (1962). They show frequent presepiments in late stages of growth and a prominent peripheral stereozone, and appear congeneric with Tabulophyllum rectum.

Some species included in Tabulophyllum by former Soviet Union authors, such as T. curtoseptatum Bulvanker 1958, and T. omulevskiense Spasskiy & Kravtsov (in Besprozvannykh et al., 1975) have complete, flat tabulae, short septa and a narrow dissepimentarium without presepiments and should be excluded from Tabulophyllum. They may be closer to Breviphyllum Stumm 1949.

Tabulophyllum bifurcatum Soshkina 1939 and T. complicatum Soshkina 1939 from the Upper Devonian (Frasnian) of the Urals have wide dissepimentarium of numerous globose or subglobose dissepiments and incomplete, axially convex tabulae, and should be excluded from

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Tabulophyllum. They are probably closer to Dohmophyllum or other related genera of Ptenophyllidae. Soshkina (1952) described and figured eight species of Tabulophyllum, but only two of them, T. grandivesiculosum Soshkina, 1952 and T. normale (Walther 1928) from the lower Upper Devonian, are referred here to Tabulophyllum. T. delicatum Soshkina 1952 is a fasciculate form with narrow dissepimentarium of small subglobose dissepiments and complete or incomplete, flat tabulae and is a species closely related to Planetophyilum. T. heckeri Bulvanker (in Soshkina, 1952), and T. pseudosociale Soshkina 1952, both from lower Upper Devonian, show no relationship with Tabulophyllum and are probably more closely related to Mictophyllum Lang & Smith 1939.

The three species described by Sun (1958) as Tabulophyllum cylindricum, T.curvatum and T. gigantum, all from the Shetianqiao Formation (Frasnian) of Hunan, are disphylloid species related to Sinodisphyllum Sun 1958.

Tabulophyllum contortum Stumm 1940, from the Nevada Limestone is a ceratoid species with strong growth constrictions, numerous radially arranged septa and incomplete, axially concave tabulae. No presepiments are observed in the type material figured by Stumm (pl.8, fig.5a-b). It is excluded here from Tabulophyllum.

Sorauf (1988) indicated that Tabulophyllum with a range from Pragian to Frasnian did not survive the Frasnian-Famennian extinction event, and none of the post-Frasnian species previously referred to Tabulophyllum could be included in this genus with certainty.

Tabulophyilum carinatum sp. nov. (Fig. 6A-H)

Etymology. From the Latin carinatum - keel- shaped structure; for the well presented carinae.

Material. Holotype UQF78596 from UQL5445- 4, Mount Podge Limestone, Mount Podge area. Paratypes: UQF78593, UQF78595, UQF78597- 8, UQF78600, UQF78611 UQF78626 and UQF79539 from UQL5445-4; and UQF78701 from UQL5450-1.

Diagnosis. Medium to large (Dc 30 to 70 mm)

trochoid or subturbinate species of Tabulophyl- lure, with strong rejuvenescence;septa numerous, moderately thickened in" dissepimentarium, carinate, frequently withdrawn from periphery.

Description. Solitary corallum, trochoid or subturbinate, medium to large, diameter ranging from 30 to 70 mm measured at calices; corallites normally with strong rejuvenescence. Calice moderately deep, with steep sides and domed floor, and with a circular depression over peripheral tabellae of about 3.5 mm wide and 3-4 mm deep; in a corallite of 34 mm calical diameter, the calice is 18 mm deep and 21 mm wide at the bottom and 30 mm wide near the calicular rim; no prominent septal grooves and interseptal ridges; corallite wall often poorly preserved. Septa in two orders, numerous, 134 septa counted at a diameter of 57 mm in the holotype, and in the largest corallite (70 mm in diameter) 77 septa were counted for each order. Septa rather strongly dilated peripherally, forming a stereozone of 1- 2 mm thick, carinate or with trabecular spines or flanges, and typically sinuous. Carinae zigzag or ill-developed yard-arm. Septa frequently interrupted peripherally by one row of large presepiments, which are commonly well developed in late growth stages. Major septa unequal, reaching or nearly reaching axis. Most septa reach axis and swirl in central region; in some corallites, major septa somewhat withdrawn from axis. Minor septa one-third to half as long as radius, commonly shorter in young stage in which presepiments are rare; cardinal fossula indistinct, but in a few sections, cardinal fossula weakly developed with cardinal septum slightly shortened and with more or less sym- metrically arranged septa in cardinal quadrants.

Dissepimentarium composed of one to three rows of large, steeply adaxially inclined presepiments, often strongly elongated. Tabularium wide, endophylloid, two-thirds as wide as the corallite; axial tabulae also wide, flat or axially slightly concave or convex, with sharply down-turned edges to meet the next one, vertically 10 to 15 tabulae per 10 mm, peripheral tabellae flat or slightly concave, about 4 nun wide in the holotype, developed in the pefiph-

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eral depression of the tabularium; trabeculae poorly preserved, coarse, 0.5-1.0 mm wide in longitudinal section, extending inwards and upwards 500 to the horizontal at the periphery; small short septal trabeculae frequently developed on upper surface of tabulae.

Remarks. Tabulophyllum carinatum sp. nov. can be distinguished from other species of this genus by having carinate septa, wide, flat axial tabulae with sharply downturned edges, and large presepiments.

Tabulophyllum shuimogouense file 1978), from the Middle Devonian of Sichuan, shows some resemblance to the present species but with major septa strongly rotated at centre, sparsely carinate septa, and narrower axial tabulae which are often incomplete and domed.

Family SPONGOPHYLLIDAE Dybowski 1873

Carllnastraea Merriam in Merriam & McKee 1976

1976 Carlinastraea Merriam in Merriam & McKe¢, p.32, 33.

Type species (OD). Carlinastraea tuscaroraensis Merriam in Merriam & McKee, 1976, pp.33, 34, pl.6, figs 5-7. pl.7, figs 1-4; Roberts Mountains Formation, late Lochkovian, southern Tuscarora Mountains, northern Nevada.

Diagnosis. Cerioid corallum; corallites slender with moderately thick walls; septa much reduced in dissepimentarium and discontinuous or irregular length in tabularium; dissepimentarium wide, consisting of one or two series of large, steeply inclined presepiments; tabularium narrow, with complete, rather widely spaced, fiat to concave tabulae.

Remarks. Carlinastraea differs from Spongophyllum (Beugniesastraea) Coen-Aubert 1988 in having a much thicker wall with well developed septal crests and all corallites with

larger presepiments. Hill (1981) doubtfully considered Carlinastraea congeneric with Spongophyllum. Pedder (1985) suggested that Carlinastraea differs from Spongophyllum in having a marginarium dominated by presepiments rather than dissepiments. HoweverYu & Jell (1990) showed that this can be variable within one colony.

Huang & Kong (in Kong & Huang, 1978) proposed Heterospongophyllum with H. simplex He ( in Kong & Huang, 1978) as the type species to include those forms like Spongophyllum but with presepiments. They believed that the massive form of Sociophyllum was probably derived from Silurian Heterospongophyllum. They originally included three species, Heterospongophyllum simplex He (in Kong & Huang, 1978), Spongophyllum kunthi Schliiter 1880, and Neomphymapseudofritchi Soshkina 1962 in their Heterospongophyllum. Neomphyma pseudo- fritchi is a species of Carlinastraea (see Pedder, 1985). According to the description and figure of the type species from the Lower Silurian of Guizhou (Kong & Huang, 1978),/-/. simplex differs from Carlinastraea in having relatively larger diameter corallites, very thin intercorallite walls and wider tabularium, and is possibly a species of Yassia Jones 1930. Spongophyllum kunthi from the Givetian of the Eifel is now the type species ofS. ('Beugniesastraea).

Carlinastraea callosa sp. nov. (Fig. 7A-F)

Etymology. From the Latin callosus - a hard skin; referring to the wide intercorallite walls.

Material. Holotype UQF78718 from UQL5459, Mount Podge Limestone, Mount Podge area. Paratypes UQF78717, UQF78722-3 and UQF78731 all from UQL5459.

Diagnosis. Large cerioid Carlinastraea with slender corallites 3-5 mm diameter; 16-22 septa confined to tabularium or absent, septal ridges

Fig. 6. Tabulophyllum carinatum sp. nov. A-C, holotype, UQF78596 from UQL5445-4, xl.5. A, TS; B, TS; C, I.~. D, E, UQF78701 from UQL5450-1, xl. D, TS; E, LS. F, H, paratype, UQF79539 from UQL5445-4. F, TS, x2; H, TS, x6, showing carinate septa. G, paratype, UQF78600 from UQL5445-4, LS, x3.

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Fig. 7. Carlinastraea callosa sp. nov. A-D, holotype, UQF78718 from UQL5459, x5, A, D, TS; B, C, LS. E, F, paratype, UQF78722 from UQL5459, E, TS, x4; F, LS, x5.

seldom seen on thickened intercorallite walls; tabularium narrow (Dt 0.9-1.8 mm).

Description. Large domed or massive, cerioid coralla up to 75 cm across, with several hundreds

of very narrow prismatic corallites; corallites slender, firmly amalgamated laterally and mostly pentagonal or hexagonal in cross section; mar- ginal, non-parricidal increase; adult corallites 3 to 5 mm in diameter. Well-defined, thickened

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walls 0.4 to 0.8 mm thick; intercorallite walls straight with prominent median dark lamellae, stereozone probably composed oft-me fibres perpendicular to median dark lamellae. Septa restricted to tabularium or even totally absent in some corallites, commonly 16-22 septa in two orders, major septa long, reaching axis, moderately thickened, thinning towards distal ends, minor septa slightly less than half as long as the major, commonly poorly presented; septal crests or ridges seldom seen on intercorallite walls.

Dissepimentarium wide, with one to three rows of large, elongated, steeply inclined presepiments or dissepiments, the outer series larger than the inner series ff more than one series are developed. Tabularium narrow, one-third of the corallite diameter or less; tabulae complete, flat or slightly concave and regularly distributed, vertically 23 to 32 tabulae per 10 mm.

Remarks. The material described above differs from Carlinastraea halysitoides halysitoides (Etheridge 1918), in lacking septal ridges on intercorallite walls which are irregularly thickened. The specimens from the Garra Formation of New South Wales described by Strusz (1966) as Spongophyllum halysitoides halysitoides and by Pedder (1985) as Carlinastraea sp. cf. C. halysitoides from the limestone lenses or megaclasts in the Coopers Creek Limestone, Coopers Creek, Victoria are similar to the material described here from Mount Podge.

Family PTENOPHYLLIDAE Wedekind 1923

Australophyllum Stumm 1949

1949 Australophyllum Stumm, p.34. 1973a Au~traiophyllum (Toquimaphyllum) Merriam, p.54.

Type species (OD). Spongophyllum cyathophylloides Etheridge, 1911, p.7, pl.A, fig.3, pl.C, figs 1, 2; Emsian, Douglas Creek Limestone, Clermont, Queensland.

Australophyllum sp. (Fig. 8A, B)

Material. UQF78720 and UQF78732 from UQL5459.

Description. Massive cerioid coralla, with polygonal corallites, four- to six-sided, with diameter of adult corallites ranging from 8 to 15 mm; intercorallite walls straight or slightly curved, 0.5 to 0.8 mm wide with prominent, straight and thin, median dark lamellae. Septa straight or slightly wavy, thin to slightly thickened, often weakly flanged in tabularium, more or less radially arranged at centre, 30 to 40 septa counted in two orders; major septa long, reaching axis and interdigitating at centre; minor septa two-thirds to three-quarters as long as radius.

Dissepimentarium composed of four to seven rows of dissepiments and presepiments with prominent boundary between tabularium and dissepimentarium; presepiments common, out- ermost series of presepiments normally larger, less steeply inclined than inner rows. Tabularium narrow, about one-third as wide as corallites, 5 mm wide in corallite of 15 mm diameter; tabulae complete and incomplete, concave with central depression, 15-18 tabulae per 10 mm vertically.

Remarks. The present specimens have corallite diameters slightly larger than those of A. cyathophylloides yohi Zhen 1994 from the Burdekin Formation of the Kirkland Downs area. They can be further distinguished from the latter in having a less well-developed peripheral zone ofpresepiments, well-developed minor septa and less closely spaced tabulae, which arc occasionally complete.

Australophyllum qinlingense Cao in Cao et al., 1983, from the Early Devonian of Sichuan has more septa (46 to 48) in corallites of similar size, and axial ends of major septa are dilated.

Gaynaphyllum Pedder 1980

1980 Gaynaphyllum Pedder, p.608.

I~pe species (OD). Xystriphyllum hyperbolicus Crickmay, 1960, p. l l , pl.7, figs 1, 2; lower part o f the Hume Formation, Eifelian, western Canada.

Diagnosis. Like Xystriphyllum but with axially domed tabulae.

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l~g. 8. A, B, Australophyllum sp. UQF78732 from UQL5459, x2.5. A, TS; B, LS. C-F, Xystriphyllum of. dunstani (Etheridge 1911). C, D, UQF78727 from UQL5459, x3. C, TS; D, LS. E, F, UQF78719 from UQL5459, E, TS, x3; F, LS, x4.

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Remarks. According to the original definition, Gaynaphyllum can be dist inguished from Xystriphyllum mainly in having domed tabularial floors and inner dissepiments not well-differen- tiated from outer tabellae, but the new species from the Laroona Formation does not show the gradation between the inner dissepiments and outer tabellae as in the type species.

Gaynaphyllum runningense sp. nov. (Fig. 9A- E)

Etymology. After Running Creek, the type locality.

Material. Holotype UQF78657 from UQL5447- 1, top of the Laroona Formation, Mount Podge area. Paratypes UQF78653-4, UQF78656, UQF78658, UQF78968, UQF78971 and UQF78981, all from UQL5447-1.

Diagnosis. Large, cerioid coralla, intercorallite walls 0.2 to 0.5 mm wide; adult corallites ranging from 6 to 10 nun in diameter; septa 16 to 21 in each order, with thickened bases; tabulae incomplete, closely spaced, with axially domed tabellae.

Description. Large cerioid coralla, domed or hemispherical with diameter up to 40 cm. Corallites polygonal in transverse sections, three- to seven-sided, with diameter of adult corallites ranging from 6 to 12 ram, commonly 7 to 8 ram; peripheral non-parricidal increase. Intercorallite walls moderately thick formed by wedge-like expansions of septal bases, straight or slightly wavy; median dark lamellae obvious, slightly wavy, between narrow stereozones of O.2 to 0.5 mm width. Septa in two orders, 16 to 21 in each, slightly wavy; relatively thin except near the intercorallite walls where they expand sharply, forming wedge-shaped peripheral ends; typically radially arranged; major septa long, unequal in length, reaching or nearly reaching axis, interdigitating at the axis; minor septa variable in length, two-thirds to three-quarters as long as radius, just extending into tabulariurn; occasionally with a few as short as one-third of radius and with distal ends attached to neighbouring

major septa. Septa rarely withdrawn from periphery.

Dissepimentarium composed of two to eight rows of small subglobose to moderately elongated dissepiments, typically four to seven rows in adult corallites; dissepiments adaxially steeply inclined, inner series somewhat elongated and almost vertically arranged; in juvenile stage, dissepimentarium narrower, consisting of only two to three rows of dissepiments. Tabularium one-third to half as wide as corallites, typically one-third of corallite in adults; tabulae incomplete, periaxial series gently adaxially inclined or flat, the axial series of domed tabellae forming a central elevation, vertically closely spaced, 22 to 30 tabulae per 10 ram.

Remarks. In the transverse section, the new species seems close to Xystriphyllum mitchelli Etheridge in Jack & Etheridge, 1892, except for its fewer septa. The longitudinal sections through the centre of the tabularium show the well developed, small, domed axial series of tabellae distinguishing it from Xystriphyllum. Gay- naphyllum hyperbolicum (Crickmay 1960) from the lower part of the Hume Formation, Macken- zie, Canada has smaller, less steeply inclined dissepiments, and gradation between inner dissepiments and outer tabellae.

Xystriphyllum Hill 1939

71938 Entelophyllum (Entelophylloides) Rukhin, p.23. 71938 Kozlowiaphyllum Rukhin, p.34. 1939 Xystriphyllum Hill, p.62. '/1972 Shastaphyllum schucherti Merriam, p.38.

Type species (OD). Cyathophyllum dunstani Etheridge, 1911, p.3, pl.A, figs 1, 2; Douglas Creek Limestone, Emsian, 8 km south-southwest of Clermont, central Queensland.

Diagnosis. See Hill (1981, p.F244).

Xystriphyllum cf. dunstani (Etheridge 1911) (Fig. 8C-F)

1911 Cyathophyllum dunstani ELheridge, p.3, pl.A, figsl,2. 1939 Xystriphyllum dunstani; Hill, p.62, pl.5, figs 5-8. 1940b Xystriphyllum dunstani; Hill, p.163, pl.3, fig~4a-b. 1942b Xystriphyllum dunstani; Hill, p.183, pl.6, fig.2a-b.

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~CHERINGA DEVONIAN RUGOSE CORALS 213

1966 Xystriphyllum dunstani; Scusz, p.578, pl.91,fig.2; 7p1.92, figs 1, 2.

1967 Xystriphyllum dunstani; Hill, Playford & Woods, pl.D8, fig.3.

1970 Xystriphyllum dunstani; Jell & Hill, p.100, pl.4, ~gs i-3.

1970 Xystriphyllum dunstani; Pedder in Pedder, Jackson & Philip, p.227, text-fig.7.

71977 Xy$~iphyllum dunstani; Jia in Jia et al., p.161, pl.57, fig. 1.

71985 Xystriphyllum dunstani; Scrutton, p.23, pl.3.3.42.

Lectotype (chosen by Hill, 1939): GSQ2425 (= C16), from the Douglas Creek Limestone, 8 krn south-southwest of Clermont, Emsian, central Queensland.

Material. UQF78727, UQF78719, UQF78721 and UQF78724 all from UQL5459.

Description. Large, massive cerioid coralla, with polygonal corallites, commonly four- to six- sided, 6 to 12 mm in diameter in adult stage. Intercorallite walls thick, with prominent gently waved median dark lamella, often irregular in thickness, ranging from 0.3 to 1.0 mm wide on each side of median dark lamellae; in one specimen (UQF78721), intercorallite walls up to 1.8 mm wide on each side of median dark lamella. Septa in two orders, 17 to 20 in each, radial to weakly pinnate in arrangement; septal bases wedge-like, thinning towards axis, wavy, weakly carinate in tabularium; major septa reaching axis and interdigitating at centre; minor septa long, two-thirds to four-fifths as long as corallite radius, extending well into tabularium; septa rarely interrupted by presepiments peripherally.

Dissepiments in four to seven rows, adaxially inclined, subglobose to moderately elongated, almost vertically arranged near the well-marked tabularial margin. Tabularium diameter 3 to 4.3 ram, slightly over one-third ofcorallite diameter. Tabulae incomplete, concave, closely spaced, with a deep central depression, 25 to 32 tabulae per 10 mm vertically.

Remarks. The type material of X. dunstani from the Douglas Creek Limestone, Clermont, ten-

tral Queensland has thinner intercorallite walls, and smooth, less dilated septa. The Mount Podge specimens are characterized by their wider intercorallite walls, the thickness of which var- ies considerably from specimen to specimen or even within one specimen. The material described as Xystriphyllum mitchelli by Philip (1965) from the Coopers Creek Limestone is also close to the present specimens, except for its smaller coraUite diameters.

Xystriphyllum ef. magnum Hill 1942a (Fig. 10E, F)

1942a Xystriphyllum magnum Hill, p.147, pl.3, fig.2a, b. 1942b Xystriphyllum magnum; Hill, p.183, pl.6, fig.3. 1966 Xystriphyllum magnum; Slrusz, p.584, pl.93, rigA. 71982 Xystriphyllum densum Yu & Kuango p.66, pl.6, fig.7.

Holotype. SUP7270, Hill 1942a, p. 147, pl.3, fig.2 a,b; Sulcor Limestone, Attunga, Late Emsian to early Eifelian, New South Wales.

Material. UQF78640 from UQL5445-12.

Description. Massive cerioid corallum; corallites polygonal, four- to seven-sided, with a mean diameter of 14 mm, the largest corallite observed with diameter of 15x20 mm. Intercorallite wails 0.3 mm wide, straight or slightly curved, with thin, median dark lamellae, which are straight or gently waved. Septa in two orders, with wedge- shaped bases, thin in dissepimentarium and tabularium, 36 to 46 in number in adult corallites, smooth, straight or slightly wavy in dissepimentarium. Major septa long, reaching axis, where they interdigitate at centre, with their axial ends curved or occasionally with zigzag-type, thin carinae; minor septa also long, just extend- hag into tabularium, three-quarters to four-fifths as long as radius. Dissepiments mainly concentric in transverse section, occasionally with some geniculate dissepiments near periphery; presepiments absent or very rare.

Dissepiments eight to eleven rows, subglobose to slightly elongated, regular size, adaxially inclined and steeper near tabularium. Tabularium

Fig. 9. Gaynaphyllum runningense sp. nov. A-E, holotypo, UQF78657 from UQL5447-1. A, LS, x6; B, LS, x6; C, TS, x6; D, TS, x2; E, TS, x6.

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narrow, one-quarter as wide as corallites, 4 nun wide in corallite of 17 mm De; tabulae incomplete, concave, with central depression, closely spaced, 28 to 32 tabulae per 10 mm vertically.

Remarks. This single specimen is closely comparable with the holotype of X. magnum from the Sulcor Limestone in all respects except its smaller corallite diameters. Hill (1942a) described the maximum diameter of this species as 20 mm. SU'usz (1966) mentioned that the type material of this species could reach a diameter of 28 mm. The holotype figured by Hill (1942a) has a corallite diameter of 23 mm. Only one transverse section of the Garra material is available (Hill, 1942b; Strnsz, 1966), the coraUite diameters ranging from 10 to 17 mm (measured from the figure illustrated by Strusz, 1966).

XystriphyUum densum from the Guitang Mem- ber, Beiliu Formation of Guangxi can only be distinguished from Xystriphyllum magnum Hill 1942a by its smaller mean corallite diameter (mean Dc 14 nun), and slightly wider tabularium. Further study of variation within each species might indicate the need to merge X. densura with the latter.

Taimyrophyllum Chemyshev 1941

1941 Taimyrophyllum Clmmyshev, pp.12, 53. 1942a Eddastraea Hill, pp. 147, 148. 1942b F_.ddastraea Hill, p.lM.

Type species (OD). Taimyrophyllum speciosum Chernyshev, 1941, pp.12, 13, 53, 54, text-fig.l, pl.1, figs 1-3; pl.2, figs 1-3; pl.5, fig.5; Lower Devonian, Tareya River, southwest Taymyr, formec Soviet Union.

Remarks. Taimyrophyllum has a wide distribution in the Lower and Middle Devonian in east- em Australia (Pragian to Givetian). It is also widely distributed in northwestern Canada (Pragian to Eifelian), central Great Basin (Mid- dle Devonian) and the former Soviet Union

(Early to Middle Devonian, Taimyr-Kolymian, Salair, Rudney Altay, Kuznetsk Basin and the Urals). However, it has not been recorded from China, southeast Asia and western Europe. This tripodal distributional pattem of Taimyrophyllum seems unusual compared with the distributional patterns of many other Old World Realm metro- politan genera in the Early and Middle Devonian.

Taimyrophyllum pedderi sp. nov. (Fig. 10A-D)

Etymology. In honour of Dr A. E. H. Pedder, who has made important contributions to knowl- edge of the Devonian rugose corals of eastern Australia.

Material. Holotype UQF78638 from UQL5445- 12, Mount Podge Limestone, Mount Podge area. Paratypes UQF78639 and UQF78641 from UQL5445-12; and UQF78729 from UQL5460.

Diagnosis. Thanmasterioid Taimyrophyllum of up to 60 cm diameter; septa 16 to 25 in each or- dec, thin and smooth in dissepimentarium and weakly carinate in tabularium; calicular platform wide, slightly everted.

Description. Thamnastecioid coralla, large massive form, domed and hemispherical with diameter up to 60 cm. Distance between axis of adult corallites ranging from 8 to 16 nun. Septa in two orders, thin and smooth in outer part of dissepimentarium, weakly carinate in tabularium, mainly zigzag carinae, axial ends dilated, and 16 to 25 in each order; major septa long, unequal in length, radially arranged, most reaching axis, typically with distal ends twisted or convoluted; minor septa long, extending to inner margin of dissepimentarium or just projecting into tabularium, indistinguishable from major septa in dissepimentarium.

Dissepiments in eight to thirteen rows, arched, small to medium sized. Calicular platform wide, slightly everted. The dissepimentarium is reflexed, thus dissepiments dip adaxially in the in-

Fig. 10. A-D, TaimyrophyUumpedderi sp. nov. A, B, holotype, UQF78638 from UQL5445-12. A, LS, x8; B, 'IS, x6. C, D, paratype, UQF78639 from UQL5445-12. C, 'IS, x3; D, I~, x4. E, F, Xystriphyllum el. magnum Hill 1942a, UQF78640 from UQL5445-12. E, I~, x3; F, 'IS, x3.

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ner part of, and abaxially in the outer part of, dissepimentarium; smaller and steeply adaxially inclined or even vertically arranged near inner margin of the dissepimentarium; near periphery ofdissepimentarium, dissepiments flat-lying, and larger dissepiments often occur periodically. Trabeculae more or less vertical in outer part of dissepimentarium and inclined adaxially upwards in inner part. Tabularium narrow, 3 to 5 mm diameter; tabulae incomplete, closely spaced, and concave, with a median depression, and vertically 35 to 42 per 10 mm.

Remarks. The specimen figured by Hill, Playford & Woods (1967) as Taimyrophyllum sp. from the Broken River Group shows a close resemblance to the present material, but has thinner septa near the inner margin of the dissepimentarium and in the tabularium, and with narrower calicular platform not evened.

Taimyrophyllum grande (Dun 1918) differs from the present species in having larger corallites (the distance between two corallites about 20 mm compared with 13 mm in T. pedderi), and more strongly convoluted distal ends of major septa, and commonly more heavily carinate septa in tabularium. Taimyrophyllum speciosum is not as consistently thamnasterioid as the present species and has a shallow inverted conical calice with a gently inclined calicular platform.

The new species shows a close resemblance to Taimyrophyllum nolani Merriam 1973b from the Eifelian of the central Great Basin, but the latter has slightly larger corallite diameters, ab- sence of zigzag carinae in tabularium, and more common occurrences o f large, f lat- lying dissepiments near the periphery of the corallites.

The specimen referred to Taimyrophyllum sp. by Pedder (in Pedder, Jackson & Ellenor, 1970) from the Timor Limestone of New South Wales has more rows of smaller dissepiments near the inner margin of dissepimentarium and a slightly larger distance between corallite axes.

Laroonaphyllum gen. nov.

Etymology. After the Laroona Homestead, south of the type locality.

Type species. Laroonaphyilum jacki gen. et sp. nov.

Diagnosis. Coralla cerioid with large polygonal corallites; intercorallite walls thin; septa in two orders, long, typically bearing zigzag carinae, and occasional ly discontinuous peripherally; dissepimentarium wide, comprising numerous small subglobose dissepiments; inner series typically smaller and more steeply inclined than outer ones; tabularium narrow, consisting of incomplete, closely spaced, adaxially inclined tabellae, typically with a central notch, but more often disturbed by convoluted, axial ends of septa.

Remarks. The new genus is similar to Acanthophyllum except for its cerioid growth form. Exilifrons Cfickmay 1968 is a disphyllid genus with domal tabulae which are not as closely spaced as in Laroonaphyllum. Taimyro- phyllum can be distinguished from Laroona- phyllum by lack of intercorall i te walls. Psydracophyllum Pedder 1971 is typically dendroid with a wide peripheral zone of presepiments and domed tabular floors.

LaroonaphyUum jacki gen. et sp. nov. (Figs l lA, B, 12, 13A-C)

Etymology. In honour of the pioneer geologist, who worked in this region, Robert L. Jack.

Material. Holotype UQF78592 from UQL5445- 4, lower part of the Mount Podge Limestone, late Emsian, Mount Podge area. Paratypes UQF78616 and UQF78590 from UQL5445-4.

Diagnosis. Cerioid coralla with large corallites of 11 to 38 mm diameter; septa numerous, thin; major septa convoluted at axis, with zigzag carinae in tabularium; dissepimentarium wide,

Fig. II. Laroonaphyllumjachigen. et sp. nov. A, B, holotype, UQF78592 from UQL5445-4, x4. A, TS; B, I~.

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0 ALCHERINGA DEVONIAN RUGOSE CORALS 217

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of small subglobose dissepiments; narrow tabularia consisting of incomplete, closely spaced, slightly concave Iabulae.

Descr~tion. Cerioid coralla with large polygonal corallites, four to seven-sided, unequal in size, diameter of adult corallites ranging from 11 to 38 mm. Calice shallow with wide platform and moderately deep central pit. Intercorallite walls thin, straight, O. 1-0.8 mm thick. Sepia in two orders, sinuous, consistently thin, 16-41 for each order, often dilated at base forming wedge-like peripheral ends; major septa long, reaching axis, with axial ends strongly convoluted in corallite centre and with zigzag carinae in tabularium; minor sepia also long, thinner than major, extending to or just into margin of Iabulariurn, three-quarters as long as radius, or longer, and peripherally often discontinuous. Presepiments occasionally developed near intercorallite walls, larger than dissepiments, and more often in cor- ners of polygonal cross sections ofcorallites.

Dissepimentarium wide, consisting of ten to fn°teen series of subglobose, adaxlally inclined dissepiments; inner series relatively steeper and smaller in size, some moderately elongated. Tabularium narrow, one-fourth as wide as corallites; tabellae incomplete, closely spaced, 30 to 40 tabulae per 10 mm vertically; tatular floors flat or slightly concave, typically with a central notch, and disturbed by convoluted distal ends of major septa.

Remarks. Laroonaphyllumjacki is the only species assigned to the new genus. It shows some resemblance to Exilifrons varians Yu (in Yu & Birenheide, 1987) from the Sipai Formation (late Emsian, serotinus Zone) of Guangxi with regard to its thin, carinate and axially rotated septa, but the latter is a disphyllid species with smaller corallite diameters (Dc 11-20 mm), fewer sepia and subcerioid growth form.

Aeanthophyllum Dybowski I873

1873 Acanthophyllum Dybowski, p.339.

Fig. 12. Laroonaphyllum jacki gen. et sp. nov., paratype, UQF78590 from UQL5445-4, TS, xl.5.

1922a Neostringophyllum Wedekind, p.16. 1922b Mesophflloides Wedekind, p.51,part/m. 1923 Ptenophyllum Wedekind, p.28. 1924 Astrophyilum Wedekind, p.46. 1924 Rhopalophyllum Wedekind, p.52. 1925 Stenophyllum Wedekind, p.9 , partim. 1925 Leptoinophyllum Wedekind, p.4.

Type species (SD; by Schliiter, 1889). Cyathophyllum heterophyllum Milne-Edwards & Haime, 1851, p.367, pl. 10, fig. 1-1b; Junkerberg Formation, Eifelian, Eifel, Germany.

Acanthophyilum (Acanthophyllum) dermontense (Etheridge 1911) (Fig. 14A-F)

1911 Cyathophyllum (?) clermontensis Eth~'idge, p.5, pl.B, figs 1, 2, pl.D, fig.3.

1939 Acanthophyllum? clermontensis; Hill, p.57, p|.4, figs 1-5.

1940a Acanthophyllum? clermontense, Hill, p.252, pl.9, fig.5.

1950 Acanthophyllum aft. clermontense; Hill, p.139, pl.5, fig.3a-b.

Fig. 13. LaroonaphyUum jaclti gen. et sp. nov. A, paratype, UQF78590 from UQL5445-4, L~q, x5. B, C, paratype, UQF78616 from UQL5445-4, B, TS, x3; C, LS, x5.

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71966 Acanthophyllum (Acanthophyllw/n) sp. cf. clermontense, Scu~z, p.552, pL85,figs 3-6; pl.86, fig.l.

1970 Acanthophyllum (Acanthophyllum) clermontense; Jell & HilL, p.97, pl.3, ~ 2-6.

1976 Acanthophyll~m (Acanthophyllum) clermontense; Fordham, p.76, pl.3, figs 5, 6.

1978 Acanthophylhmf clermontense; He, p.148, pi.72, rigA.

Lectotype (chosen by Hill, 1939, p.57). GSQF2414 (=2c), figured by Etheridge, 1911, pl.D, fig.3; and by Jell & Hill, 1970, pl.3, rigA; Douglas Creek Limestone, Emsian, from Doug- las Creek, 11 km south-southwest of Clermont, central Queensland.

Material. UQF78591, UQF78594, UQF78614- 5 and UQF78617 from UQL5445-4; UQF78634 from UQL5445-10; UQF78643-4 from UQL5445-12; UQF78586 from UQL5444; UQF78668 from UQL5447-2; and UQF78730 from UQL5460.

Description. Large solitary or weakly compound coraUum, turbinate to patellate form, with adult diameter ranging from 12.2 to 70 mm. Calice slightly everted with flared rim, wide platform, and very deep calicular pit about width oftabul- arium; in one specimen calical pit is 10 mm wide at the bottom and 22 mm deep; calicular floors flat or slightly concave. Epitheca very thin, often poorly preserved, with faintly developed septal grooves and interseptal ridges. Septa in two orders, numerous, (29-40)x2; expanded sharply at periphery to form wedge-shaped septal bases which are laterally contiguous to comprise a narrow but prominent peripheral stereozone of rather irregular thickness ranging from 0.14 to 1.2 ram. Triangular septal bases typically retiform, with small rounded voids of 0.05 to 0.1 mm diameter. Beyond the stereozone, septa are thin to moderately thickened, thickest in middle part ofdissepimentarium; in outer part ofdissepimentarium, somewhat spindle-shaped, typically thin, smooth and wavy; moderately thickened and flanged or carinate in inner part of dissepimentarium. Major septa long, and reaching axis; often irregularly dilated and carinate in tabularium, typically convoluted at centre;

carinae mainly zigzag type. Minor septa just extending into tabularium, three-quarters as long as radius. Lateral dissepiments common in outer part of dissepimentarium, especially in larger specimens; in one large corallite of 70 mm diameter, lateral dissepiments very well developed, sometimes as many as 3 rows of lateral dissepiments flanking one side of septum. Pre- sepiments rarely occur only in larger specimens.

Dissepiments numerous, subglobose, more arched and somewhat horizontally floored near periphery or abaxially inclined when calice is inverted, steepening inwards, and almost vertically arranged near tabularium. Tabularium narrow, the ratio of Dt/Dc varying considerably from specimen to specimen, ranging from 0.14 to 0.36; tabulae incomplete, slightly concave, with a cen- teal notch, closely spaced, commonly strongly interrupted by convoluted axial ends of major septa; vertically 18-20 tabulae per 10 mm.

Remarks. Compared with the type material from the Douglas Creek Limestone, the present specimens have larger diameters, up to 70 mm in one patellate specimen that is doubtfully referred to this species. The Mount Podge specimens also show distinctive retiform septal bases, which are contiguous laterally forming a narrow but prominent peripheral stereozone along with fibrous sclerenchymal tissue normal to the epitheca.

Family DISPHYLLIDAE Hill 1939

Disphyllum de Fromentel 1861

1861 Disphyllum de Fromentel, p.302. 1866 Astrocalamocyathus Ludwig, p.188, partim. 1866 Toeniodendrolopas Ludwig, p.188, partim. 1893 Cannophyllum Chapman, p.45. 1922a Schlueteria Wedekind, p.3, parlim. 1939 Megaphyllum Soshkina, pp.14, 46. 1952 Solominella Ivaniya, p.141. 1978 Pseudodsphyllum Kong in Kong & Huang, p.79. 197g Temnophylloides Luo (MS) in Kong & Huang, p.76.

Type species (SD; by Lang & Smith, 1934). Cyathophyllum caespitosum Goldfuss, 1826, p.60, pl.19, fig.2b; = Disphyllum goldfussi (Geinitz, 1846); Givetian, Eifel, Germany.


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Remarks. Disphyllum is one of the Devonian's most cosmopolitan genera with about 70 species recognized. The present author is inclined to subdivide the genus into three subgenera: D. (D.) de Frornentel, 1861, D. (Pseudodisphyllum)

Kong in Kong & Huang, 1978, and D. (Solom- inella) Ivaniya, 1952.

Kong (in Kong & Huang, 1978) proposed the name of Pseudodisphyllum to include those lran- sitional forms between Disphyllum and

B

Fig. 14. Acanthophyllum (Acanthophyllum) c[ermontense (ECheridge 1911). A, B, UQF78634 from UQL5445- 10, xl.5. A, TS; B, LS. C-F, UQF78614 from UQL5445-4. C, TS, xl.5; D, I~, xl.5; E, TS, x3, showing the septa| structure; F, TS, xl.5, showing compound growth form.

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Cylindrophyllum Simpson 1900. He originally indicated that Pseudodisphyllum could be distinguished from Disphyllum by having flanged septa. Kong originally included five new species from the Dushan Formation (Givetian) of Guizhou into Pseudodisphyllum. The type species, Pseudodisphyllum fiangzhaiense Kong in Kong & Huang, 1978, is a phaceloid species with fusiform septa, which are thickest near the inner margin of dissepimentarium and attenuating towards axis. However, Pseudodisphyllum guizhouense Kong in Kong & Huang, 1978 is a cerioid species and is more closely related to Argutastrea and Cystohexagonaria Yoh in Yoh & Bai, 1978. Herein Pseudodisphyllum is regarded as a subgenus of Disphyllum to include those species with flanged, weakly carinate septa previously referred to Disphyllum. The following additional species should be included in Disphyllum (Pseudodisphyllum) because of the occurrence of prominent flanges and zigzag carinae: Disphyllum breviseptatum Yoh 1937, Middle Devonian, Guangxi, south China; and Disphyllum eurekaense Merriam 1974, late Emsian to early Eifelian, Nevada Formation, central Great Basin. Disphyllum (Pseudo- disphyllum) bears a close resemblance to Paradisphyllum SUusz 1965 and lipheophyllum Hill 1956, in respect to the flanged or carinate septa. However, both Paradisphyllum and lipheophyllum have wide, arched dissepimental floors formed by the reflexed calicular rim. Ad- ditionally, septa are often discontinuous near the periphery in the type species of Paradisphyllurn and in lipheophyllum carinae are strongly developed and cross-bar carinae are common.

Temnophylloides was described by Luo (1965) in his postgraduate thesis at Beijing University and published in Kong & Huang (1978). In Luo's original description, he listed Temnophyllum devonicum SoshkJna 1949, as the type species, but remarked that the new genus was better represented by the material from Hunan, which he referred to a new subspecies. The specimens described by Luo (1965) would be referable to Temnophyllum devonicum except for their prominent fasciculate growth form with long,

subcylindrical corallites. According to Luo (1965), Temnophylloides could be distinguished from Disphyllum by its prominent peripheral stereozone formed by the strongly dilated septal bases and from Temnophyllum by its fasciculate growth form. Temnophylloides seems very dose to Alaiophyllum Goryanov 1961, except that its septa are not as withdrawn from the axis as in Alaiophyllum. In terms of the present author's understanding, Temnophylloides becomes a junior synonym ofSolorainella Ivaniya 1952. How- ever it seems practicable to separate those forms with a rather wide and consistent peripheral stereozone from Diskhyllum, and Solomineila Ivaniya is retained to include this group as a subgenus of Disphyllum.

Disphyllum (Disphyllum) de Fromentel 1861

Type species. Cyathophyllum caespitosurn Goldfuss 1826.

Diagnosis. Fasciculate to subcerioid, with lateral or peripheral increase; septa complete, rarely withdrawn from periphery, carinae absent; trabeculae monacanths; dissepiments in several series, small, globose or subglobose; tabularium typically biseriate, of fiat, concave or slightly mesa-shaped axial plates and large, adaxially inclined periaxial tabellae.

Disphyllum (Disphyllum) paracouvinense sp. nov. (Fig. 15A-F)

Etymology. Referring to the similarity to D. (D.) couvinense Tsien 1969.

Material. Holotype UQF78690 from UQL5447- 3, lower part of the Mount Podge Limestone, Running Creek. Paratypes UQF78681-2, UQF78689, UQF78984-9 and UQF78992-3, all from UQL5447-3.

Diagnosis. A species of Disphyllum (Disphyllum) with a wide range of coraUite diameters (6-22 mm), 18-33 septa of unequal length, and widely spaced tabulae.

Description. Phaceloid coralla with cylindrical corallites, increasing peripherally. Corallites

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Fig. 15. Disphyllum (DisphyUum)paracouvinease ap. nov. A-D, holotype, UQF78690 from UQL5447-3. A, TS, x2; B, TS, x2; C, IS, x3; D, LS, x3. E, F, paratype, UQF78682 from UQL5447-3, x2. E, TS; F, LS.

closely spaced, often in contact, and unequal in size with diameters of adult corallites ranging from 6 to 22 mm. Corallite walls smooth, thin, 0.1 to 0.2 mm wide, without prominent septal grooves and interseptal ridges. Septa in two orders, (18-33)x2 in adult corallites; moderately dilated in dissepimentarium and very thin in tabularium; non-carinate, but often with rather coarse septal surface and slightly wavy in dissepimentarium. Major septa rarely reaching axis, unequal in length; mostly withdrawn from axis; minor septa half as long as radius or slightly longer, just extending into tabularium.

Dissepimentarium composed of two to five rows of globose dissepiments, axially inclined near tabularium and more or less horizontal near walls; in young corallites (De < 6 mm), dissepimentarium composed of only two or three

series ofglobose dissepiments, which are regular in size and one above another. Tabularium wide; tabulae complete or incomplete, horizontal or slightly concave axially, frequently with down-turned edges, and often supplemented with adaxially inclined peripheral tabcllae in adult stage; in young corallitcs, tabulae commonly complete and regularly concave; tabulac rather widely spaced, vertically 9-14 tabulae per I0 ram.

Remarks. The type material of Disphyllum (D.) couvinense Tsien 1969, from the base of the Horizon Co2b (Eifelian), Belgium has major septa only about two-thirds of the corallite radius and the corallites have diameters of 12 to 15 mm. The new species can be distinguished from D. (D.) couvinense by having a wider range of corallite diameters and more frequent com-

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plete tabulae, especially in juvenile forms.

Family PHILLIPSASTREIDAE Hill 1954

Phillipsastrea d'Orbigny 1849

1849 Phillipsasteea d'Orbigny, p.2. 1850 Phillipsastraea d'Orbisny, p.107. 1850 Lithostrotion d'Orbigrty, p.106,param. 1851 Smithia Milne-Edwards & Haime, p. 171, p.421. 1855 Medusaephytlum Roemer, p.33. 71881 Pseudoacercularia Schl~et, p.84. 1940 StreptastraeaLang, Smith& Thomas, p.125.

Type species (SD, by Milne-Edwards & Haime, 1851). Astrea (Siderastrea) hennahii Lonsdale, 1840, p.697,partim, only pl.58, fig.3,3b, not 3a; late Givetian limestones, Barton Quarry, Torquay.

Diagnosis. See McLean (1989, p.239).

Remarks. Horseshoe dissepiments are only irregularly developed and typically mark the inner margin of dissepimentarium in Phillipsastrea as shown by the type specimen ofPh. hennahii. A careful review of the species included in Phillipsastrea revealed two groups within the genus. One is represented by the type species, Phillipsastrea hennahii, with horseshoe dissepiments at the inner margin of dissepimentarium and globose dissepiments gently abaxially inclined, trabecular fans very narrow over the horseshoe dissepiments and the trabeculae in the outer part of dissepimentarium nearly vertical. The other group is represented by Phillipsastrea oculoides Hill 1942b and Ph. producta Jin & He 1982 in which horseshoe dissepiments are developed near the middle part of the dissepimentarium with globose dissepiments on both side forming arched dissepimental floors and the trabecular fans are relatively wider (see Strusz, 1965, p.565, text-fig.22).

Many authors (such as Scrutton, 1968; Hill, 1981) regarded or doubtfully regarded Pachyphyllum Milne-Edwards & Haime 1851, as a junior subjective synonym of Phillipsastrea. Scrutton (1968) remarked that '...with regard to the considerable disparity in size between the re- spective type species, the existence of species of intermediate size must not be overlooked'.

Sorauf(1978) regarded Pachyphyllum as a separate genus and six criteria were used to differen- tiate species of Pachyphyllum and Phillipsastrea; he suggested that'...these six criteria must be uti- lized in concert, and only in certain geographic areas can one rely on a single character'. He em- phasized the presence of a uniform row of horseshoe dissepiments at the inner margin of dissepimentariurn in Pachyphyllum. He included in Pachyphyllum a group of species with large tabularium, represented by the type species and a group of smaller forms, such as P woodmani (Hall & Whitfield 1873). On the other hand, in most American species of Pachyphyllum the inner walls of the corallites are strongly projecting above the intervening spaces; the coralla of these forms are usually not as large. McLean (1986, 1989) restricted Pachyphyllum to those species with coralla characterized by a small number of large corallites projecting well above the surface of the main, massive part of the coraUum, a uniform collar of horseshoe dissepiments, a larger diameter tabularium and coarser trabeculae. McLean (1986) employed Medusaephyllum Roerner 1855 for species with the internal mor- phology of Pachyphyllum, but lacking the so- called bouchardi growth form. Later, he (1989) correctly regarded Medusaephyllum to be a junior synonym of Phillipsastrea.

Phillipsastrea macuiosa Hill 1942a

1942a Phillipsastrea maculosa Hill, p. 153, pl.3, fig.5a-b. 1954b Phillipsastrea maculosa; Hill, p.l O7, pl.6, fi& l a-b.

Holotype. SU7268, Hill, 1942a, pl.3, fig.5a-b; Sulcor Limestone, late Emsian to early Eifelian, Tamworth, New South Wales.

Phillipsastrea sp. cf. P. maculosa Hill 1942a (Fig. 16H-J)

Material. UQF78642 from UQL5445.

Description. Thamnasterioid corallum, massive, exterior shape and overall size unknown; neigh- bowing corallites about 7 to 13 mm apart from axis to axis. Epitheca of individual corallites totally absent. Septa in two orders, 15 to 18 in

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each, radially arranged, relatively thin in outer part of dissepimentarium, occasionally discontinuous and sinuous midway between Iabularia, strongly dilated, somewhat fusiform in inner part of dissepimentarium, and more or less continuous from one corallite to the next; thickest part of sepia corresponding to trabecular fans and horseshoe dissepiments in longitudinal section, often laterally contiguous forming incomplete inner walls near inner margin of dissepimentarium. Sepia with very rough sides and some separation of trabeculae, and irregularly carinate. Major septa long, reaching or nearly reaching axis, often with distal ends slightly curved, very thin in tabularium; minor septa also 10ng, just extending into Iabularium.

Dissepimentarium wide, composed ofnumer- 0us, horizontally based subglobose or globose dissepiments. An irregular series of small horseshoe dissepiments is well developed near inner margin of dissepimentarium, and ot~en with one or two rows ofadaxially inclined, small globose dissepiments at inner side of horseshoe dissepiments. Symmetrical, narrow trabecular fans are centred over the row of horseshoe dissepiments; trabeculae coarse, about 0.2 mm wide, rhipidacanths, poorly preserved. Tabularium about 4 to 4.5 mm in diameter; tabulae incomplete, and horizontally floored, with their upper surface slightly domed, vertically about 35 to 40 tabulae per 10 ram.

Remarks. The specimen is comparable with the type material of Phillipsastrea maculosa Hill 1942a, from the Sulcor Limestone of New South Wales, but differing from it in having longer major sepia, which are more or less continuous in the tabularium and often reaching the axis.

The present specimen shows some resemblance to the specimens referred to Phillipsastrea ibergensis (Roemer 1855) from the Mount Frome Limestone (late Emsian to early Eifelian), New South Wales described by Wright (1966), in its similar number of septa, and diameter of tabularium, but the latter has a uniform row of horseshoe dissepiments, hounding the tabularium and more heavily carinate septa.

The type material of Phillipsastrea oculoides Hill 1942b, from the Garra Formation has shorter major sepia, which do not often project into the tabularium, and typically, gently concave Iabulae, but as noticed by Strusz (1965), Iabulae vary from sagging to moderately domed in the holotype.

Protomaegeea Rozkowska 1956

1956 Protomacgeea Rozkowska, p.280.

Type species (OD): Protomacgeea dobruchnensis Rozkowska, 1956, p.281, figs 2-8; Grzeg- orzowice beds, No. 3, Eifelian, the Holy Cross Mts, Poland.


Remarks. The specimen figured by Rozkowska (1956, pp.281,282, figs 2-4) as the holotype of the type species Protomacgeea dobruchnensis Rozkowska 1956 was not numbered and was recorded as being from different beds. The specimen shown in fig. 2 is from the strongly detrital marls of Grzegorzowice beds No. 2. The longitudinal section (fig. 3) and the transverse section were recorded from the limestone of Grzegorzowice beds No. 3 and could be from different specimens.

Wright (1966) reported the occurrence of Protomacgeea in the Mount Frome Limestone (late Emsian-early Eifelian) of the Mudgee district, New South Wales. Hill et al. (1967), Jell (1967) and Jell & Hill (1969) reported a new species of Protomacgeea from the Chinaman Creek Limestone, Broken River Province, north Queensland. A poorly preserved specimen was also recorded from a conglomerate clast at the base of the Drummond Group, Ukalunda, north Queensland (Jell & Hill, 1969). However, neither of these two Australian species has been for- mally published. They have a relatively larger diameter compared with the type species from Poland. Jell (1967) studied the sepial microstruc- ture of the material from the Broken River, and noted its rhipidacanthine trabeculae arranged in symmetrical fans.

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Protomacgeea minor sp. nov. (Fig. 16A-E)

Etymology. From the Latin minor - small; referring to the small size ofcorallites in this species.

Material. Holotyp¢ UQF78612 from UQL5445- 4, lower part of Mount Podge Limestone, Mount Podge area. Paratypes UQF78599, UQF78601- 10, UQF78613, UQF79538 and UQF78618-25, all from UQL5445-4.

Diagnosis. Very small trochoid corallum, with maximum diameter of 7 mm; septa in two orders, ( 14-2 l)x2, strongly dilated peripherally, and embedded into a rather thick outer wall, and laterally contiguous forming a peripheral stereozone; major septa long, connecting at axis to form an axial columella; dissepimentarium consisting of one series of hoseshoe dissepiments; tabulae complete and slightly concave.

Description. Small trochoid coralla, solitary, with apical angle of 400 to 450; corallites straight or slightly curved, with maximum diameter 7 ram; corallite surface smooth, with very weak trans- versely wrinkled constrictions, no septal grooves and interseptal ridges observed. Walls rather thick, 0.15 mm wide, composed of horizontally arranged fine sclerenchyme fibres, with a distinctive boundary and with bases of trabecular septa imbeded in sclerenchymal tissue. Calice not very deep, two-fiRhs of whole length of corallite, with flat or slightly concave floor; in the holotype, which is a corallite of 4 mm height and 2.6 nun maximum diameter, the calice is 2 mm wide and 2 mm deep; in a specimen CLIQF78599) of 6 mm maximum diameter and 12 ram height, the calice is 4 nun wide near the rim and 3 mm deep, occasionally a calicular boss develops. Septa in two orders (13-21)x2, more or less bisymmetrically arranged; strongly dilated peripherally forming a wide peripheral stereozone of O.5 to 1.0 nun wide. Major septa

long, reaching or nearly reaching axis, thinner in tabularium, slightly thickened at distal ends and connecting to form an axial columella or ring-like axial structure leaving a very small space at centre of O.2 to 0.3 nun diameter. Minor septa 0.5 to 1.5 nun long, two-fifths as long as radius, about the width of peripheral stereozone. Cardinal fossula indistinctive, but occasionally occurring in late stage, with cardinal septum shortened; in a paratype, a small axial structure formed by joining of the axial ends ofsepta opens to the cardinal fossula; one specimen showing flat side in early stage, in which distal ends of all septa are laterally contiguous to form axial s~ucture; cardinal septum on flat side is longer than others, and extends into ring-like structure, di- viding it into two parts.

Trabeculae very well preserved, rhipidacanths, 0.15 to 0.2 mm wide in longitudinal section. Symmetrical trabecular fans centred over single row of horseshoe dissepiments, no peripheral horizontal dissepiments observed. Tabulae complete, slightly concave, between peripheral stereozone and axial structure, vertically 0.3 to 0.5 mm apart.

Remarks. The new species shows some resemblance to the undescribed species of Proto- macgeea from the Broken River Group (Hill et al., 1967; Jell & Hill, 1969) in respect to their trabecular pattern. The present species has smaller diameter corallites, well-developed axial structure, indistinctive cardinal fossula and lacks peripheral horizontal dissepiments.

Protomacgeea dobruchnensis is similar in size and number of septa, but has shorter septa, distinctive cardinal fossula, well-developed flat dissepiments and no axial structure.

The species of Protomaegeea from the Mount Frome Limestone, Mudgee district, New South Wales (Wright, 1966) has a larger diameter (up to 22 mm), more septa (20-38x2), and a distinc-

Fig. 16. A-E, Protomacgeea minor sp. nov. A, paratype, UQF78599 from UQL5445-4, LS, x8. B, paratype, UQF78601 from UQL5445-4, 'IS, x9. C, holotype, UQF78612 from UQL5445-4, LS, xl0. D, paratype, UQF79538 from UQL5445-4, TS, xl0. E, paratype, UQF78605 from UQL5445-4, TS, xl0. F, G, ThamnophyUum sp., UQF79110 from UQL5445-4, xl0. F, TS; G, LS. H-J, Phillipsastrea sp. cf. P. maculosa Hill 1942a, UQF78642 from UQL5445. H, TS, x2; I, TS, x8; J, LS, x8.

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tive cardinal fossula. The dilated axial ends of the major septa, except for the short cardinal septum, form an elevated, arcuate, eccentric columella. There are rare peripheral flat dissepiments, and tabulae are lacking.

Thamnophyllum Penecke 1894

71894 Thamnophyllum Penecke, p.596. 1909 Phacellophyllum G~ch, p.102. 71977 Newpetrozium Jia/n Jia etai., p.151. 1982 Thamnophylloides Jin & He, p.117.

Type species (SD, by Lang & Smith, 1935). Thamnophyllum stachei Horn & Penecke in Penecke, 1894, p.594, pl.8, figs 1-3, pl.11, figs 1, 2; late Emsian to early Eifelian, near Graz, Austria.

Diagnosis. See McLean (1989, p.243).

Remarks. Schoupp6 (I958), Fliigel (1956a, 1956b), Rozkowska (1957), Soshkina (1951, 1952, 1954), Scrutton (1968), Birenheide (1978) and McLean (1989) regarded Phacellophyllum as a junior synonym of Thamnophyllum, and Wang (1950) considered Thamnophyllum as a subgenus of Phacellophyllum. Stumm (1949), Hill (1954a, 1956, 1981), McLaren (1959), Jia (in Jia et al., 1977), He (1978), Jell (1967) and Jin & He (1982) retained or doubtfully retained Thamnophyllum and Phacellophyllum as separate genera. According to their general concep- tion, such as Jia (in Jia et al., 1977), He (1978), and Jin & He (1982), Thamnophyllum could be distinguished from Phacellophyllum only by its shorter, more dilated septa and complete tabulae. Hill (1981) suggested that Phacellophyllum could be distinguished from Tharanophyilum by its well-presented outer row of fiat dissepiments, and frequent occurrence of peripheral tabulae. The type species of Thamnophyllum has the dissepimentarium largely concealed by the sclerenchyme tissue of the peripheral stereozone and without fiat dissepiments. McLean (1989) remarked that the morphological differences between the type species of Thamnophyllum and Phacellophyllum could not be accorded generic rank.

Thamnophyllum sp. (Fig. 16F, G)

Material. UQF79110 from UQL5445-4.

Description. Very slender cylindrical fragments, possibly from fasciculate corallum. CoraUites with diameter about 3.5 mm, no budding observed; epitheca very thin, weak septal grooves and interseptal ridges. Septa poorly differenti- ated into two orders, subequal, moderately thickened, with prominent, median dark lamellae, fusiform, noncarinate, short, 32 in number in a corallite of 3.2 mm diameter; major septa about 0.6 mm long, one-third as long as radius; minor septa only slightly shorter and thinner than major.

Dissepimentarium biserial and composed of an outer single row of horizontal fiat dissepiments and an inner single row of horseshoe dissepiments, which are about 0.25 to 0.3 mm wide and 0.3 mm high. Tabulae complete, horizontal, about 15 to 16 tabulae per 10 mm vertically. Trabeculae rhipidacanthine, fanned over the horseshoe dissepiments.

Remarks. This specimen is very close to the species of Thamnophyllum from the Broken River Province (Jell, 1967), but the latter has relatively longer major septa and larger diameter (5.5 mm in the holotype) corallites. Stratigraphically, the Broken River material probably occurs higher than the material from the Mount Podge Lime- stone described above.

The specimen is also close to those from the Guitang Member (late Emsian part), Beiliu For- marion of Guangxi described as Thamnophyllum uniense Soshkina by Yu & Kuang (1982), but the latter has a larger corallite diameter and slightly more convex, fiat dissepiments.

Acknowledgments This paper represents part of the author's Ph.D. thesis submitted to the University of Queensland. The studies were supported initially by an Academia Sinica Overseas Postgraduate Schol- arship and then a University of Queeusland Post- graduate Research Scholarship.

I am very grateful to Dr J. S. Jell, who sug-

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gested the project and gave constant advice and encouragement, and to Dr R. W. Day, the Chief

Government Geologist of Queensland, who supported me with transportation and other facili-

ties during my field trips. S.C. Lang, J. J. Draper, L. M. Gunther and T. P. T. McLennan provided much assistance and kindness during my field work. Many thanks also to Dr R. Mawson and Professor J. A. Talent of Maequarie University for identification of the brachiopods and provid-

ing unpublished data on conodonts from the Mount Podge Limestone. I wish to thank Drs J. E. Sorauf, A. J. Wright, A .E.H. Pedder and B. D. Webby for their helpful comments on the various versions of this manuscript.

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ALCHERINGA D E V O N I A N R U G O S E C O R A L S 233

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A P P E N D I X

Locality register o f the described material

All L-numbers refer to University of Queensland localities (UQL) and are registered under the Geologi- cal Museum locality catalogue system. Grid references are in parentheses following the L-numbers and refer to the Ewan, 1:100,000 map of Australia Na- tional Topographic map Series, sheet 8059, indicated by EW. Each continuous stratigraphic section is represented by one L-number with the stratigraphic unit number hyphenated to indicate the sequential order of the collection and the position in the section.

L5444 (EW872657). Mount Podge Limestone, seven small limestone outcrops, east of the dam, about 100 m south of the railway line, about 7.8 km northwest of l.arnona Homestead, Mount Podge.

L5445-1 (EW887652). Top of the Laroona Forma- tion, southwest side of Mount Podge. L5445-2. At the base of the Mount Podge Limestone, 60 m northeast of L5445-1. L5445-3. Mount Podge Limestone, 30 m northeast ofL5445-2. L5445-4. Mount Podge Limestone, 42 m northeast of L5445-3. L5445-6. Mount Podge Limestone, 78 m northeast and then 160 m northwest of L5445-4. L5445-8. Mount Podge Limestone, 81 m northeast of L5445-6. L5445-10. Mount Podge Limestone, 42 m northeast of L5445-8. L5445-12 (EW890655). Top of the Mount Podge Limestone, 22 m northeast of L5445-10.

L5447-1 (EW915638). Laroona Formation, conglomerate and sandstone, in Running Creek, 4 km N.N.W. of Laroona Homestead. The section is northward along the east bank of the creek. L5447-2. At the base of the Mount Podge Limestone, 347 m north of the base of L5447-1. L5447-3. Mount Podge Limestone, 117 m north of L5447-2. L5447-4. Mount Podge Limestone, 94 m north of L5447-3. L5447-5. Mount Podge Limestone, 64 m north of L5447-4. L5447-6. Mount Podge Limestone, 146 m north of L5447-5, in fault contact with L54~7-5. L5447-7. Mount Podge Limestone, 64 m north of L5447-6. L5447-8. Mount Podge Limestone, 24 m north of L5447-7. L5447-9. Mount Podge Limestone, 44 m north of L5447-8. L5447-10. Mount Podge Limestone, 17 m north of L5447-9, in fault contact with L5447-9. L5447-11 (EW913648). Top of the Mount Podge Limestone, 5 m north of L5447-10.

L5450-1 (EW892662). Mount Podge Limestone, northwest side of Mount Podge.

L5459 (EW890650). Mount Podge Limestone, Carlinastraea callosa beds, south side of Mount Podge.

L5460 (EW889651). Mount Podge Limestone, south side of Mount Podge, in fault contact with L5459.

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late emsian rugose corals of the mount podge area, burdekin basin, north queensland

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