Journal of Asian Earth Sciences 19 (2001) 177±194

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Middle Permian brachiopods from central Peninsular Malaysia Ð faunal af®nities between Malaysia and west Cambodia Masatoshi Sone a,1,*, Mohd Shafeea Leman b, Guang R. Shi a a

School of Ecology and Environment, Deakin University, Rusden Campus, Clayton, Vic. 3168, Australia School of Environmental Sciences and Natural Resources, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia

b

Accepted 27 April 2000

Abstract A moderately diverse Permian brachiopod fauna is described from a new rock unit, the Bera Formation, in the Bera District, central Pahang, Peninsular Malaysia. The fauna consists of 19 taxa, including 14 genera and 17 (both identi®ed and unidenti®ed) typically Tethyan species. The fauna appears to be correlative on the basis of brachiopods with the Neoschwagerina-Yabeina fusulinid Zones in Indochina and South China. In particular, it has strong linkage to Member C (Yabeina beds) of the Sisophon Limestone, west Cambodia. This is indicated by three of the Bera species Ð Urushtenoidea chaoi (Ching), Spyridiophora gubleri Termier and Termier, and Transennatia termierorum sp. nov., being shared with the Cambodian fauna. A possible early Capitanian (Middle Permian) age is proposed for the Bera brachiopod fauna. q 2001 Elsevier Science Ltd. All rights reserved.

1. Introduction and regional geology This paper describes the largest known Permian brachiopod fauna from the so-called Central Basin (or the Central Belt) of Peninsular Malaysia, and reports on the southernmost occurrence of such a fauna in that region. Most of the fossils examined in this study were collected by Sone and Leman in the Bera District in February 1998. Additional material came from a collection in the palaeontology laboratory of the National University of Malaysia (Universiti Kebangsaan Malaysia), collected originally by Leman. All described specimens are registered in the same university with pre®x UKM-F. The regional geology of the Bera District is known from only two brief reports (Cook and Suntharalingam, 1970; MacDonald, 1970). The rock unit from which the current fossil horizon is studied has been named the Bera Formation (Leman et al., in press). The area where it occurs was previously considered to be part of the Triassic Semantan Formation Ð for example, on the most recent of®cial geological map of Peninsular Malaysia (Geological Survey of Malaysia, 1985). The Semantan Formation, as revised, outcrops west of the Bera Formation, which in turn is * Corresponding author. E-mail address: [email protected] (M. Sone). 1 Present address: Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia.

¯anked on the east by ?Jurassic to Cretaceous continental sediments of the Bertangga Sandstone, but exposures to east and west are poor (Leman et al., in press). The northern and southern boundaries of the Bera Formation are also uncertain, but in the north it is associated with an andesitic volcanic unit thought to be of a Late Permian age (Cook and Suntharalingam, 1970). The occurrence of the fusulinid, Parafusulina sp., from a small limestone outcrop near Tasik Bera implies that the lower part of the Bera Formation may extend down into the Kungurian (upper Early Permian) (Cook and Suntharalingam, 1970).

2. Stratigraphy The fossils of this report come from the ªSungai Bera sectionº (grid reference WF151446; the National Map Malaysia 1:50 000 Sheet 4157) exposed on the eastern side of Bera Road, 17.3 km south of Felda Sebertak junction (Fig. 1). This section is equivalent to Locality BF2 of Leman et al. (in press). The outcrop was originally prepared for construction of a resort. The exposed strata extend approximately 90 m vertically and 300 m laterally, and appear to have undergone gentle but pervasive soft-sediment deformation westwards. It is thus dif®cult to trace the sequence from one side to the other. The general strike is N50±708W and dip 70±908SW. The lithology consists mainly of mudstone±siltstone interbedded with sandstone,

1367-9120/01/$ - see front matter q 2001 Elsevier Science Ltd. All rights reserved. PII: S 1367-912 0(00)00026-2

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Fig. 1. Possible extent of the Bera Formation and the location of the Sungai Bera section (modi®ed after Leman et al., in press).

becoming dominantly tuffaceous in the upper part of the section. The surface is bleached soft sediment. More outcrops occur along the road; these are much smaller than the Sungai Bera section. Three major fossiliferous stratigraphic intervals can be discriminated; these are referred to as Assemblages A±C in ascending order (Fig. 2). Assemblage A is predominantly a coquina of shell debris in grey mudstone; it often contains fossiliferous calcareous concretions. This assemblage holds the richest brachiopod deposit in this report. Assemblage B has less common brachiopods in nearly white, ®ne- to medium-grained sandstone. Assemblage C consists principally of abundant lyttoniids in purple to black tuffaceous mudstone and siltstone. The brachiopods are listed in Table 1. The genus Transennatia is the most abundant, although most specimens are too poorly preserved for satisfactory illustration. The three assemblages share few taxa, and may appear to represent distinct faunules, but sampling was not exhaustive. The three assemblages are considered as a single fauna in the following discussion. 3. Correlation and age At present, there is no globally accepted Permian chronological standard. The time-scale compiled by the International Subcommision on Permian Stratigraphy (Jin et al., 1997) is applied in this study, with slight modi®cation to fusulinid biozonation for more appropriate correlation within South-East Asia (Fig. 3). The brachiopod fauna of the Sungai Bera section appears to correlate with the other

brachiopod faunas in the Neoschwagerina-Yabeina Zones of the Middle Permian, most likely to a lower Capitanian, This is based on the three brachiopod species shared with Member C (Yabeina beds) of the Sisophon Limestone, west Cambodia. The Permian strata of west Cambodia are exposed mainly in the Sisophon and Battambang regions (Fig. 4), as limestone hills with intercalations of tuff, shale and mudstone strata, collectively called the Sisophon Limestone (Ishii et Table 1 Brachiopod species from each assemblage of the Sungai Bera section Species

Assemblages A

Derbyia sp. Urushtenoidea chaoi (Ching, 1963) Urushtenoidea sp. Transennatia termierorum sp. nov. Transennatia sp. indet. Spinomarginifera sp. Echinauris sp. Spyridiophora gubleri Termier and Termier, 1970a Kozlowskia sp. Phricodothyris sp. Orthotetes aff. picta Fang, 1983 Strophalosiina sp. Linoproductus sp. Rhynchonellid family indet. Orthothetina cf. iljinae Sokol'skaya, 1965 Gubleria aff. ninglangensis Fang and Jiang, 1992 Gubleria sp. Eolyttonia? sp. Spiriferinid family indet.

B

X X X X X X X X X

C X X

X X X X

X X X X X

M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

Fig. 2. Stratigraphic log of the Sungai Bera section with indications of each Assemblage A to C (modi®ed after Leman et al., in press).

al., 1969). Brachiopods from the studied by several palaeontologists 1914; Chi-Thuan, 1961; Termier 1970a, 1970b). Ishii et al. (1969)

limestone have been (e.g., Mansuy, 1913, and Termier, 1960, listed 50 brachiopod

179

species from their collections gained from that unit; Nakamura (1979a) later revised the list. Based on lithology, Ishii et al. (1969) divided the limestone into four members, A±D in ascending order (Fig. 3). This division is supported by fusulinid faunas, supplemented by coral and brachiopod data. Because the four members are de®ned on lithologies and because a few key fusulinid species range through more than one member, the boundaries of each member cannot be precisely assigned chronologically. The most likely biostratigraphic ranges of the four members are indicated in Fig. 3, based on studies of Ishii et al. (1969) and Waterhouse (1976). Member C consists of shale and calcareous mudstone with calcareous nodules, sandwiched by the limestone facies of Members B and D (Ishii et al., 1969). It contains abundant brachiopods, and is characterised by fusulinid species such as Yabeina asiatica, Lepidolina multiseptata, Sumatrina longissima and Verbeekina verbeeki. Y. asiatica is the primitive form of the genus, suggesting the lowest part of the Yabeina Zone (Ishii, 1966). L. multiseptata and S. longissima are typical species of the middle Midian (the Y. globosa Zone) of the eastern Tethys (Leven, 1992, 1993). Species of Sumatrina and Verbeekina are not known to extend up to the upper Midian (see Kobayashi, 1997). The Midian Stage is approximately equivalent to the Capitanian Stage (Jin et al., 1997), although the lower boundary of the former is probably slightly older than that of the latter, as pointed out by Leven and Grant-Mackie (1997). Therefore, an early Capitanian age is the most de®nable for Member C of the Sisophon Limestone. The most important species in the Bera fauna is Urushtenoidea chaoi (Ching) whose many reported occurrences are con®ned to the Kuhfengian Stage (Wordian) of South China and Member C of the Sisophon Limestone. It is one of the index fossils for the lower Maokouan Sub-series in the Nanjing Hills, southeastern China (He and Shi, 1996). An equally important form is Spyridiophora gubleri Termier and Termier. It has previously been found only in the Yabeina beds of the Sisophon Limestone (Mansuy, 1913; Chi-Thuan, 1961; Termier and Termier, 1970a). Nakamura (1979a) also listed its occurrence in Member C. Transennatia termierorum sp. nov. is elsewhere known only in the Yabeina horizon of the Sisophon Limestone (Termier and Termier, 1970a). These three species indicate strong linkage between the Bera fauna and that in Member C (the Yabeina Zone) of the Sisophon Limestone. Species of Orthotetes, Gubleria, Phricodothyris and Orthothetina in the Bera fauna closely resemble those recorded in the Gnishik-Khachik beds of the Trans-Caucasus and in the Maokou horizons of South China. In particular, the species of Orthotetes and Gubleria are alike to those in west Yunnan, southwestern China. Additionally, the Bera fauna displays some similarity to the Capitanian brachiopods of the Lengwu Formation, Zhejiang Province, South China (see Liang, 1990). Almost all Bera genera occur in the Lengwu fauna, including relatively rare taxa such as

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Fig. 3. Age of the Sungai Bera section brachiopods and possible correlation with other horizons in the Central Basin of Peninsular Malaysia, the Sisophon Limestone of west Cambodia, and South China. Only part of the Early Permian is shown. Time-scale after Jin et al. (1997); fusulinid biozonation and the stratigraphy of localities in the Central Basin adopted from Fontaine et al. (1994); Jengka Pass, Lst. ˆ Limestone Member, S & S. ˆ Sandstone-shale Member; Aring Formation is partly adapted from Yanagida and Aw (1979); Sumalayang Limestone is from Igo et al. (1979); divisions of the Sisophon Limestone are after Ishii et al. (1969).

Strophalosiina and Spyridiophora; the latter are uncommon or absent in the underlying and overlying Maokou and Wuchiaping Formations. There are no species in common between the Bera and Lengwu faunas. This may be due to slightly different ages, or slightly different climatic conditions, or both. No genus of the Bera fauna is particularly suggestive of a Late Permian age. Transennatia, Orthothetina and Gubleria are regularly seen in the Wuchiapingian Stage of South China and in the Djul®an Stage of the Trans-Caucasus. However, some species of these genera occur in Middle Permian rocks of South China, west Cambodia, Japan, south Primorie and possibly Timor. The presence of Urushtenoidea in both Assemblages A and C is good evidence for precluding the possibility of a Late Permian age. The genus has never been found in postKuhfengian horizons in South China; its closest relative Uncisteges persisted only into the lower part of the overlying Lengwuan horizon; its youngest occurrence, on present data, is in Member C of the Sisophon Limestone. Spyridiophora and Kozlowskia are consistent with a pre-Wuchiapingian age. These genera are dominant in the Early Permian, and have never been unequivocally reported from Late Permian rocks. Their youngest occurrence may in fact be in the Lengwu Formation.

Fig. 4. Possible distributions of the Indochina and East Malaya terranes, and locations of the Bera District and the Sisophon and Battambang regions: 1, Uttaradit-Nan Suture; 2, Raub-Bentong Line (modi®ed after Metcalfe, 1998).

M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

There is a generally accepted view that large lyttoniids such as Leptodus and Gubleria are typically Late Permian (e.g., Shen and Shi, 1996) and, in the case of Oldhamina, this seems to be true. It is only partly true for brachiopod faunas from the Trans-Caucasus and South China, where the Lengwu fauna is now known to contain numerous lyttoniids, but notably without Oldhamina. Many lyttoniid forms occur in the Middle Permian of Indochina (e.g., Mansuy, 1912, 1913, 1914; Termier and Termier, 1960; Chi-Thuan, 1961), and in the Neoschwagerina horizon of the Sosio Limestone, Italy (e.g., Rudwick and Cowen, 1967). Lyttoniids are in fact more common in the Middle rather than Late Permian of Japan, with several reports from the lower Kanokuran Series (e.g., Tazawa, 1976, 1987; Tazawa and Matsumoto, 1998). The presence of numerous lyttoniids in Assemblage C is thus not necessarily indicative of a Late Permian age; it may in fact be the Middle Permian. Permian strata are prominent in the Central Basin of Peninsular Malaysia. Ages with respect to the age of the Sungai Bera section are indicated in Fig. 3. These strata are mostly limestones dated by fusulinids, corals, algae and/or radiolarians. In contrast, only a few Permian brachiopods have been described, despite many reported occurrences (e.g., Jones et al., 1966; Gobbett, 1973; Fontaine et al., 1994). Igo (1964) and Nakamura (in Nakazawa, 1973) described some poorly preserved brachiopods of little biostratigraphic value. Yanagida and Aw (1979) described a small suite of Permian brachiopods from Kelantan; they considered the fauna to be late Djul®an. The so-called ªLeptodus Shaleº fauna in north-central Pahang has been known for over a half century (MuirWood, 1948, p. 5, footnote; Jones et al., 1966). Leman (1993, 1994) reported 49 species of brachiopods from the same fauna, but they have not been described. Based on the presence of abundant lyttoniids, the fauna was thought to be Late Permian, but this is questionable. Thus, based on brachiopods, it is still dif®cult to make precise correlations within the Central Basin. Nevertheless, considering available data from palaeontology and regional geology, the Sungai Bera section is likely to more or less align stratigraphically with the nearest limestone units at Kampung Awah and Jengka Pass (Fig. 3). Both the limestones yield Yabeina asiatica which is known elsewhere only in the Sisophon Limestone (Ishii, 1966; Igo, 1967; Ishii et al., 1969; Fontaine et al., 1994). North of the Bera District, small outcrops of late Middle Permian age have been studied in recent years (e.g., Fontaine et al., 1994; Jasin et al., 1995). There are thus prospects for better understanding of Permian correlations in central Pahang, speci®cally with respect to the Bera brachiopod fauna. 4. Palaeogeographic implications All taxa identi®ed to generic and speci®c levels accord with this being a warm-water fauna. This is indicated by

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Orthothetina, Transennatia, Gubleria, Spinomarginifera, Urushtenoidea and Strophalosiina, all of which are typical Tethyan elements. The close af®nities between the Bera and Sisophon faunas were discussed above. Relationships with the Middle Permian brachiopods of South China are less obvious. Differences in Middle Permian brachiopods between Indochina and South China have been pointed out by Nakamura et al. (1985). It is, therefore, assumed that the Central Basin with the Bera fauna was located in the same or nearby climatic zone as west Cambodia during the age-interval in question, but that there were slight differences in ecology or geographic position relative to tropical South Chinese Cathaysia, hindering faunal interchange. This conclusion is consistent with the tectonic model of Metcalfe (1996a, 1996b, 1998), namely, that the East Malaya terrane accommodating the Central Basin was part of the Indochina terrane (Fig. 4).

5. Systematic palaeontology (M. Sone) The systematic study follows the supra-ordinal classi®cation of Williams et al. (1996), and the classi®cations of Williams and Brunton (1993) for the Strophomenida, Brunton et al. (1995) and Brunton and Lazarev (1997) for the Productida, and Carter et al. (1994) for the Spiriferida and Spiriferinida. The suborder Strophalosiidina Schuchert, 1913 refers to Brunton (written communication, 27 January 2000). Class Strophomenata Williams et al., 1996 È pik, 1934 Order Strophomenida O Suborder Orthotetidina Waagen, 1884 Superfamily Orthotetoidea Waagen, 1884 Family Orthotetidae Waagen, 1884 Subfamily Orthotetinae Waagen, 1884 Genus Orthotetes Fischer de Waldheim, 1829 Orthotetes aff. picta Fang, 1983 Fig. 5; 1±2. cf. Orthotetes picta Fang, 1983, p. 94. pl. 1, ®gs. 4±6. Description. Ventral shell medium for genus, 27 mm wide and 26.5 mm long with greatest width at hinge; outline Ushaped; pro®le fairly ¯at; ears not demarcated; interarea very low; umbo inconspicuous; rectimarginate; plications absent. Ornament of moderate costellae with angular crests, increasing by intercalation, numbering approximately 18 per 10 mm at anterior margin; growth lines few, well de®ned, with two high ones at mid-length and near the anterior margin, with the anteriormost one extending from the end of the hinge. Ventral interior with long median septum, extending to nearly mid-valve, slightly thickened anteriorly, height unknown; spondylium minute, laterally semi-ovate; dental ridges uniting to form a spondylium;

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M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

adductor ®eld weakly concave. Details of dorsal valve unknown. Remarks. The ¯attened ventral shell with a small apical spondylium and a median septum suggests Orthotetes. Its U-shaped outline, low in¯ation and rectimarginate commissure makes it close to O. picta Fang (1983, p. 94. pl. 1, ®gs. 4±6) from the Neoschwagerina horizon of the Xiaoxinzhai section in Gengma, west Yunnan. There are de®nable, but subtle differences in the condition of the costellation Ð the Yunnan species displays slightly coarser (7±8 in 5 mm) and more intercalated costellae. This, however, may be due to differing stages of their maturity; the larger Bera valve seems older ontogenetically with more growth lines. It is thought to be close to O. picta, but additional material is required for con®dent identi®cation. No other previously described species appears close to the present form. Family Derbyiidae Stehli, 1954 Subfamily Derbyiinae Stehli, 1954 Genus Derbyia Waagen, 1884 Derbyia sp. Fig. 5; 3±4 Remarks. The ventral shell with a high and thick median septum bisecting the umbo is indicative of Derbyia, but it is small for the genus, almost equidimensional (about 23 mm wide to 22 mm long), and the hingeline is the widest part. The outline is U-shaped, the ears are not differentiated, and the pro®le is convex in the median region, but contrastingly ¯at on the ¯anks. The interarea is low and narrow, and cardinal extremities are obtuse. The umbo is prominent. The shell is rectimarginate without plications, and is undistorted. The septum is relatively short. Costae are coarse, low and blunt, increasing by intercalation, with width fairly uneven. The present species is characterised by the umbonal region being prominent, raised and demarcated by a strong concentric line. D. altestriata Waagen illustrated by Fantini Sestini (1965, p. 38, pl. 3, ®g. 7) from the upper Ruteh Limestone, north Iran, displays a very similar elevated umbo with a primary lamina. It differs from the Bera form in possessing a more transverse outline and ¯attened median region. Family Meekellidae Stehli, 1954 Subfamily Meekellinae Stehli, 1954 Genus Orthothetina Schellwien, 1900

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Orthothetina cf. iljinae Sokol'skaya, 1965 Fig. 5; 5±6 cf. Meekella cf. baschkirica Tschernyschew; Mansuy, 1914, p. 22, pl. 2, ®g. 15. cf. Orthothetina iljinae Sokol'skaya, 1965, p. 204, pl. 30, ®gs. 1a and b, 2. cf. Orthothetina iljinae Sokol'skaya; Tong, 1978, p. 214, pl. 77, ®gs. 9a±c. Description. Shell medium sized, approximately 46 mm wide and 47 mm long, with greatest width at mid-valve; outline sub-pyramidal; pro®le gently convex; ¯anks smoothly rounded; sulcus and plications absent; umbonal region ¯attened, not recurved. Ornament of radial capillae, moderately strong, even in width, numbering 18±20 per 10 mm at anterior margin. Concentric crenulation present near the frontal margin, 4 mm in maximum width Ð growth lamella otherwise not de®ned on surface. Ventral interior with pair of dental plates, high, arising directly from valve ¯oor, extending at least one-third of valve-length, gently divergent internally, slightly converging apically. Dorsal characters unknown. Remarks. The non-plicate ventral valve with strong, parallel dental plates clearly indicates Orthothetina. The present form is characterised by a sub-pyramidal outline and low in¯ation. It closely resembles O. iljinae Sokol'skaya (1965, p. 204, pl. 30, ®gs. 1a and b, 2) from the Gnishik and Khachik Formations of the Trans-Caucasus by lacking growth lines but instead having a de®ned concentric elevation anteriorly. O. iljinae, as illustrated by Tong (1978, p. 214, pl. 77, ®gs. 9a±c) from the Limestone Member of the upper Chihsia Formation, Guizhou, South China, has somewhat more ¯attened shells than the TransCaucasian form. O. phetchabunensis Yanagida (1964, p. 14, pl. 2, ®gs. 5 and 6) from the upper Middle Permian of central Thailand is similar in outline and size, but its costellation is slightly stronger than the Bera form. The Thai species displays concentric rugae over the ventral surface, a feature lacking in the present form. A Cambodian shell illustrated by Mansuy (1914, p. 22, pl. 2, ®g. 15) as Meekella cf. baschkirica Tschernyschew from Phnom Takream comes close to the Bera shell in having a sub-pyramidal outline and lacking concentric lines. The specimen is, however, much smaller than the Bera form, so precise identi®cation is dif®cult. It

Fig. 5. 1±2, Orthotetes aff. picta Fang, internal and external moulds of ventral shell showing a trace of spondylium with dental ridges, £ 1.5, UKM-F258, Assemblage B. 3±4, Derbyia sp., rubber cast of ventral exterior, ventral internal mould, £ 2.0, UKM-F259, Assemblage A. 5±6, Orthothetina cf. iljinae Sokol'skaya, rubber cast of ventral exterior, ventral internal mould showing a pair of dental plates, £ 1.5, UKM-F260, Assemblage C. 7±14, 16±18, Urushtenoidea chaoi (Ching); 7±11, rubber cast of dorsal interior, dorsal external mould Ð ventral, left lateral views, and anterior view showing mould of short trail with re¯ected dorsal hollow spines, £ 2.0, enlarged anterior view showing delicate concentric laminae, £ 4.5, UKM-F261, Assemblage A. 12± 14, broken ventral internal mould Ð ventral, anterior and lateral views, £ 2.0, UKM-F262, Assemblage A. 16±18, internal mould of ventral trail Ð ventral view showing trace of ªmarginal fenceº and ªtransverse ridgeº, lateral views (right and left), £ 1.5, UKM-F263, Assemblage C. 15, Urushtenoidea sp., dorsal external mould, £ 2.0, UKM-F264, Assemblage A. 19, Strophalosiina sp., rubber cast of dorsal exterior with stereozone, £ 1.5, UKM-F265, Assemblage B.

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shows no plication, hence is more likely Orthothetina rather than Meekella. Order Productida Sarytcheva and Sokol'skaya, 1959 Suborder Strophalosiidina Schuchert, 1913 Superfamily Aulostegoidea Muir-Wood and Cooper, 1960 Family Aulostegidae Muir-Wood and Cooper, 1960 Subfamily Chonosteginae Muir-Wood and Cooper, 1960 Genus Urushtenoidea Jing (Jin) and Hu, 1978 Discussion. Urushtenoidea has a complicated spine arrangement on the anterior margin of both valves. The genus has considerable similarity to Urushtenia Likharev. The former, however, differs from the latter most obviously in having scaly concentric lamellae (composed of spinous structure) on the ventral trail. The ventral geniculation is much stronger in Urushtenoidea; its angle often exceeds 608. Urushtenoidea has a ¯attened dorsal disc with a poorly de®ned trail, whereas Urushtenia has a geniculate dorsal valve with a moderately long trail. Species of Urushtenoidea commonly show ®ner costae than Urushtenia. Occurrence. Urushtenoidea has hitherto been recorded only from the eastern Tethys of South China, Cambodia, Laos and Japan. This report con®rms the presence of the genus in Peninsular Malaysia. Urushtenoidea is limited to the Xiangboan to Kuhfengian Stages (Roadian to Wordian) of South China, Members A±C of the Sisophon Limestone, the Parafusulina-Neoschwagerina horizons of Laos, and the Lower Kanokuran Series of Japan, all of which are correlated with the Guadalupian Series. Urushtenoidea chaoi (Ching (Jin), 1963), Fig. 5; 7±14, 16±18 Urushtenia chaoi Ching, 1963, p. 15, pl. 1, ®gs., 1±4, 9± 12, 16; pl. 2, ®gs. 7±8, 11±17. cf. Urushtenia chenanensis (Chan); Tong, 1978, p. 218, pl. 78. ®gs. 16a±c. Urushtenia chaoi Ching; Tong, 1978, p. 218, pl. 78, ®g. 18a±d. Urushtenoidea chaoi (Ching); Jing (Jin) and Hu, 1978, p. 116, pl. 2, ®g. 10. Urushtenoidea chaoi (Ching); Nakamura, 1979b, p. 230, pl. 2, ®g. 4. Urushtenoidea chaoi (Ching); Hu, 1983, pl. 3, ®gs. 6a±c. Urushtenia chaoi Ching; Zhang et al., 1983, p. 266, pl. 93, ®gs. 12a±e. Urushtenoidea chaoi (Ching); Zeng, 1987, pl. 1, ®gs. 1± 5, 8±14, 16±19, 26±27, 29±30; pl. 2, ®gs. 4, 7, 9. Description. Shell average- to large-sized for genus, 19.5± 23 mm wide and 16±18 mm long, with maximum width at mid-length; outline cylindrical; ears not differentiated; median fold not well de®ned. Ventral valve moderately geniculate; trail long; umbo unknown. Dorsal valve almost

¯at on disc, trail about 2 mm. Radial costae well marked on trail but poor on disc, numbering about 40; terminations of dorsal costae developing into hollow spines anteriorly along edge of re¯exed marginal ledge; spines incurved dorsally at approximately 35±408; concentric wrinkles poorly de®ned on disc; concentric laminar delicate over dorsal trail, dense and regular (Fig. 5; 9±11). Dorsal interior with robust bi®d, sessile cardinal process; lobes sub-ovate, large; alveolus absent; posterior platform weakly in¯ated; buttress plates absent; muscle scars large, palmate, hanging anteriorwards; breviseptum narrow and long; brachial ridges weak and small; ear baf¯es strong; marginal rim moderately elevated. In specimen UKM-F263 (Fig. 5; 16±18), moulds of dorsal marginal endospines (`marginal fence spines' of Zeng, 1987, p. 502) individually appear as cylindrical hollows lying on the interior of costal crests. A row of endospines collectively forming a high `marginal fence' (term from Jing (Jin) and Hu, 1978, p. 258) behind the ventral trail. The spines are laterally supported by the band of the `marginal fence transverse ridge' (term from Zeng, 1987, p. 502) at mid-length of spines Ð appearing as a fracture in this specimen. Remarks. Three shells (one dorsal and two ventral) were available. The present form is referred to Urushtenoidea mainly because of its short dorsal trail and spinous nature of the ventral trail. It is broadly similar to the type species, U. chaoi (Ching). There are minor differences from the Chinese form. The Bera form has slightly ®ner costae and a less strongly elevated posterior platform. The present form is very much like the Cambodian form, U. chaoi of Zeng (1987), in having comparable hanging palmate-shaped adductor scars. Zeng's materials are from the Neoschwagerina margaritae bed of the Sisophon region. Jing (Jin) and Hu (1978), in de®ning the genus, referred to it having a large alveolus. This feature is not developed on the present material, and is not always apparent in other congeneric Chinese or Cambodian species. Occurrence. U. chaoi has been reported from Member C of the Sisophon Limestone, Cambodia, and the Maokou Formation, Kuhfeng Formation, and Hsiaochiang Limestone of South China, all horizons correlative with the Neoschwagerina-Yabeina Zones. Urushtenoidea sp. Fig. 5; 15 Urushtenoidea maceus (Ching); Nakamura, 1979b, p. 227, (not pl. 1, ®gs. 1±4), pl. 2, ®gs. 1±3. Remarks. One dorsal valve referred to Urushtenoidea has a short dorsal trail (1.5 mm). It is medium-sized for the genus (21 mm wide and 12 mm long). It clearly differs from U. chaoi in possessing a more transverse outline and slightly coarser costae. Its cardinal process lobes are globular in shape, unlike the semi-oval shape of those in U. chaoi,

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suggesting that this is a second species of Urushtenoidea in Assemblage A. Its raised ventral ears are well impressed on the mould. Three specimens illustrated by Nakamura (1979b, p. 227, pl. 2, ®gs. 1±3) as U. maceus (Ching) each from Members A, B and C of the Sisophon Limestone are close to the present form in outline and costae, and are probably conspeci®c. In addition, Jing (Jin) and Hu (1978) placed U. maceus in Uncisteges, but Nakamura (1979b) has not accepted this. Urushtenia khmeriana Termier and Termier (1970b, p. 446, text-®gs. 2A±D, pl. 29, ®gs. 1±7) from Phnom Te Bon, Sisophon, has a ¯attened dorsal shell and scaly laminae on its ventral trail. It is thus doubtlessly a species of Urushtenoidea. It has costae and raised ventral ears similar to those of the present form, but no dorsal view is available. Genus Strophalosiina Likharev, 1935 Strophalosiina sp. Fig. 5; 19 Remarks. A single mould of a dorsal exterior was available. The shell is large for the genus (26 mm wide and 18.5 mm long), transverse, with a semi-circular anterior margin, moderate geniculation. It has a well-de®ned dorsal trail (3.5 mm long) with no concentric wrinkles, unlike that of Urushtenoidea. The stereozone is smooth, seemingly very thick, wide and reaching the hinge; it is widest (2 mm) in front. Ornament of ®ne and sharp costae with weak and irregular rugae are present on the disc but are absent from the trail. No dorsal external spines are discernible. The large transverse shell with the wide and thick stereozone suggests Strophalosiina. The stereozone does not seem to have any re¯ected hollow spines or funnels, unlike those of other genera of Chonosteginae such as Urushtenia and Chonosteges. The present form shares several similarities with the type species, S. tibetica (Diener) from Tibet, Cambodia, Timor, Iran and the Trans-Caucasus. The Malaysian species, however, differs most obviously in being larger and having ®ner costae Ð at least 45 in number, whereas typical S. tibetica has about 30. S. zhejiangensis Liang (1990, p. 154, pl. 18, ®gs. 9±20) from the lower Lengwu Formation, China, is less transverse than the Bera form, and has many large nodules on the dorsal disc. Superfamily Lyttonioidea Waagen, 1883 Family Lyttoniidae Waagen, 1883 Subfamily Lyttoniinae Waagen, 1883 Genus Gubleria Termier and Termier, 1960 Comments. Gubleria was originally proposed by Termier and Termier (1960) with the type species, G. disjuncta from the Sisophon Limestone. The genus is characterised most importantly by the discontinuous nature of its ventral median septum and corresponding dorsal median holes. Cooper and Grant (1974, p. 411) and Grant (1976, p.

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162), however, asserted that Gubleria is invalid, being immature, or a variation of Leptodus Kayser. In this study, on the contrary, numerous specimens of large lyttoniids, thought to belong to two species, were collected. Almost all specimens possess more or less developed median incisions or segments, indicative of Gubleria, whereas no form clearly assignable to Leptodus was found. This seems to contradict to the view of Cooper and Grant (1974) that Gubleria has individual rather than a generic characters. Gubleria aff. ninglangensis Fang and Jiang, 1992 Fig. 6; 1±8 cf. Gubleria ninglangensis Fang and Jiang, 1992, p. 327, pl. 1, ®gs. 3±5. Description. Shell size varied, average width 19 mm and length more than 57 mm; conical, spatulate, weakly twisted; outline long and narrow, umbo strongly obtuse, gradually expanding and ¯attening towards mid-length, then twisted anteriorly; lateral pro®le ¯at to slightly convex. Ventral valve interior with obsolete muscle ®eld; median septum highest medianly, faintly continuous, rough incisions de®ned anteriorly; lateral septa symmetrical, short and thin, numerous Ð 4 to 5 in 10 mm, at least 18 for longest but still incomplete specimen UKM-F287 (Figs. 6; 7); interseptal spaces much wider than septal ridges, deeply grooved along septal margins. Dorsal internal plates ¯at and long, laterally grooved along margins; median trough segmented irregularly. No trace of papilla observed. Other details unknown. Remarks. Seven specimens (®ve ventral and two dorsal internal moulds) are identi®ed as a species of Gubleria by possessing a discontinuous or segmented median line on both ventral and dorsal valves. This species is characterised by its elongate shape. G. ninglangensis Fang and Jiang (1992, p. 327, pl. 1, ®gs. 3±5) from the lower Emishan Basalt Formation (lower Capitanian), west Yunnan, is very close in outline and septal apparatus; it is believed to be closely allied to the present form. Collemataria tilita Liang (1990, p. 229, pl. 40, ®g. 2; pl. 41, ®g. 11) from the lower Lengwu Formation also has a similar outline with an anteriorly segmented median septum, and is possibly allied as well. The type species, G. disjuncta Termier and Termier (1960, p. 241, text-pl. 3, ®gs. 11±14; ®g. 1), is represented by a couple of fragmentary internal plates. They are ®nely papillose and show median holes that are irregular in size and shape. Other details are unknown, preventing useful comparison. Gubleria sp. Fig. 6; 9±12 Remarks. Three ventral shells are assigned to Gubleria. The shell is medium sized (about 36 mm wide and more than 47 mm long). The outline is ovate, with an umbonal angle

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Fig. 6. 1±8, Gubleria aff. ninglangensis Fang and Jiang; 1±2, ventral internal mould of umbonal coneÐventral and lateral views, £ 1.5, UKM-F282, 3, anterior part of ventral internal mould, £ 1.5, UKM-F283, 4, ventral internal mould, £ 1.5, UKM-F284, 5, internal mould of dorsal internal plate showing segmentation of median slit, £ 1.5, UKM-F285, 6, internal mould of dorsal internal plate, £ 1.5, UKM-F286, 7, ventral internal mould, £ 1.5, UKM-F287, 8, ventral internal mould of two imbricating shells, £ 1.5, UKM-F288. 9±12, Gubleria sp.; 9, ventral internal mould displaying anteriorly segmented median septum, £ 1.5, UKM-F289, 10, ventral internal mould, £ 1.5, UKM-F290, 11±12, ventral internal mould, rubber cast of ventral exterior, £ 1.5, UKM-F291. 13±14, Eolyttonia? sp., fragment of ventral internal mould, £ 1.0, £ 3.0, UKM-F292, (All specimens are from Assemblage C).

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ranging 80±1058, and the pro®le is fairly ¯at. They may belong to Gubleria aff. ninglangensis because of their similar septal apparatus. The present material, however, is much wider and has greater umbonal angles. Specimen UKMF291 is problematic in that it seems to have an incurved beak (?) pointing dorsally (Fig. 6; 11). The present form super®cially recalls Leptodus nobilis (Waagen). The chief differences, however, concern the median septum. The Bera form displays a weakly raised median septum with segmentation. Mature individuals of L. nobilis typically have a high, thick, continuous, bladelike median septum. This feature is well illustrated by Noetling (1905, pl. 17, ®gs. 1 and 2; pl. 18, ®gs. 1 and 2) for specimens of L. nobilis from the Wargal and Chhidru Formations of the Salt Range, Pakistan. Genus Eolyttonia Fredericks, 1924 Eolyttonia? sp. Fig. 6; 13±14 Remarks. A fragment of a ventral internal mould exhibits the distinctive ¯uting of its angustilobate septal apparatus, suggesting possible location in Eolyttonia. It is small (9 mm in width) for the genus, but its true length is uncertain. It is longitudinally conical in shape, narrow and slightly wider anteriorly. The ventral interior has well-formed septa with crests broadly grooved and low. The lateral septa are transverse, and the interseptal troughs are rounded. It may be a juvenile, although the internal structure is relatively well expressed. Suborder Productidina Waagen, 1883 Superfamily Productoidea Gray, 1840 Family Productellidae Schuchert, 1929 Subfamily Marginiferinae Stehli, 1954 Tribe Paucispiniferini Muir-Wood and Cooper, 1960 Genus Transennatia Waterhouse, 1975 Transennatia termierorum sp. nov. Fig. 7; 1±10, 13±15 Spyridiophora ? timorensis Termier and Termier, 1970a, p. 58, pl. 5, ®g. 1, text-®gs. 1±4. Etymology. After Professors Henri Termier and GenevieÁve Termier, who greatly contributed to knowledge on the South-East Asian Permian brachiopods, and ®rst described this species. Type specimen. The specimen illustrated as Spyridiophora ? timorensis Termier and Termier (1970a, pl. 5, ®g. 1) (not Hamlet) is designated as the holotype. Diagnosis. Shell medium-sized, geniculation abrupt, sulcus and fold strong anteriorly, ears proportionally large and projecting, costae notably converging, branching, and sturdy on the trail, coarse reticulation present on the disc. Description. Shell size ranging from 13±23.5 mm wide, 9±14.5 mm long, more than 8.5 mm high, commonly small;

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outline sub-quadrate, widest part at hinge; ears large, convex and costate; ventral umbo pointed; geniculation abrupt with angle about 508; dorsal disc fairly ¯attened, semi-circular in shape, almost without a fold, notably reticulate; trail long. Surface coarsely ornamented by sturdy costae, branching on the ¯anks, converging in the sulcus and on the fold, with crests rounded, and similarly rounded, wide and shallow interspaces; reticulation con®ned to disc, concentric rugae faint on posterior part of the trail. Ventral interior with wide and ¯at platform, well elevated, extending from apex; muscle scar obsolete or obscure. Remarks. This form is characterised by its abrupt and strong geniculation and extended ears. It is most like Spyridiophora ? timorensis Termier and Termier (1970a, pl. 5, ®g. 1, text-®gs. 1±4) from Sisophon, Cambodia, but lacks the row of spine bases along the ears seen in the Cambodian material; this could be an artifact of preservation. Termier and Termier (1970a) originally de®ned their timorensis as a synonym of Productus gratiosus Waagen, as illustrated by Broili (1916, pl. 2, ®gs. 4±5, 7±13), from the Basleo, Koeka bei Koeafeoe and Bitauni areas of Timor. However, Grant (1976, p. 134) asserted that only the Basleo specimens belong to Transennatia, the others being Retimarginifera. This view was followed by Archbold (1984, p. 115) and Archbold and Bird (1989, p. 108). Waterhouse (1978, p. 119) considered Termiers' timorensis to be a junior synonym and homonym of Productus timorensis Hamlet. T. timorensis is close to T. termierorum in size and ornament. The former can, however, be distinguished from the latter by having much smaller ears and a strongly folded dorsal disc. There is admittedly considerable variation among the materials of T. timorensis as understood by Archbold and Bird (1989, p. 108). For instance, the Kasliu form of Archbold and Bird (1989, p. 107, ®gs. 3E±I) is exceptionally large and has ®ne ornament, whereas the BasleoWesleo forms of Hamlet (1928, pl. 2, ®gs. 2±4) have strong ornament and a deep fold. The small specimen in ®g. 7 of Diener (1897, pl. 3) identi®ed as P. gratiosus from the Chitichun Limestone, Tibet, has a deep sinus and sturdy costae; it strongly resembles T. termierorum. This Tibetan material is possibly conspeci®c, although it is slightly wider in outline. T. insculpta Grant (1976, p.135, pl. 32, ®gs. 1± 37; pl. 33, ®gs. 1±16) from the Rat Buri Limestone of southern Thailand is similar. Its geniculation is, however, not as abrupt as that of the new species. T. termierorum differs from the type species, T. gratiosa (Waagen, 1884, pl. 72, ®gs. 3±7) from the Salt Range, Pakistan, most obviously in having larger ears. The geniculation of T. gratiosa varies from shell to shell; smaller ones tend to be much less geniculate. This trend is probably common in the genus; it occurs also in T. insculpta Grant. The small but highly geniculate Bera shells may, therefore, be mature forms. Occurrence. This form was originally recorded from the Yabeina bed of a limestone hill, Phnom Takream, in the Battambang region (Termier and Termier, 1970a, p. 58), suggesting that its occurrence is con®ned to an upper part

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of Member B to Member D, most likely Member C of the Sisophon Limestone. Transennatia sp. indet. Fig. 7; 11±12 Remarks. The small conjoined shells (9 mm wide and 7.5 mm long) with no geniculation is probably a juvenile. It, nevertheless, displays features diagnostic of the genus, such as strong reticulation, tiny ears and branching costae. It occurs in Assemblage C. Tribe Marginiferini Muir-Wood and Cooper, 1960 Genus Spinomarginifera Huang, 1932 Spinomarginifera sp. Fig. 7; 16±17 Remarks. One squashed mould of a dorsal interior was available. The distinctive appearance of its muscle ®eld suggests alliance with Spinomarginifera. The shell is small for the genus; its dorsal disc is about 13 mm wide and 13.5 mm long. It seems to have a row of endospines around the anterior half of the disc (characteristic of the genus) but this is poorly preserved. There is too little information for speci®c assignment. Subfamily Overtoniinae Muir-Wood and Cooper, 1960 Tribe Costispiniferini Muir-Wood and Cooper, 1960 Genus Echinauris Muir-Wood and Cooper, 1960 Echinauris sp. Fig. 7; 18 Remarks. One internal mould of a dorsal disc was available. It is medium-sized (18 mm wide and 15.5 mm long). Its panlike appearance with a low and short median septum indicates Echinauris. As common in the genus, brachial ridges are not de®ned. The marginal ridge is moderate. The external features are not unknown, but rugose radiations occur internally, probably corresponding to the external spines. Considering the poorly developed internal structure, it may be an immature shell, although it is not small in size. A Cambodian shell ®gured by Termier and Termier (1970b, p. 450, ®g. 6) as E. opuntia (Waagen) has a longer median septum than the Bera form. Productus khmerianus Mansuy (1914, p. 17, pl. 6, ®gs. 3a±d) from Phnom Roang, Sisophon, possibly belongs to Echinauris, as suggested by Waterhouse and Piyasin (1970, p. 130) and Grant (1976, p. 123). Prior to them, Ishii et al. (1969, p. 48) reported the

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same species as Echinauris khmerianus from Member D of the Sisophon Limestone. Its interior is not known, hindering comparison with the Bera material, although the size is close. Family Productidae Gray, 1840 Subfamily Productinae Gray, 1840 Tribe Spyridiophorini Muir-Wood and Cooper, 1960 Genus Spyridiophora Cooper and Stehli, 1955 Spyridiophora gubleri Termier and Termier, 1970a Fig. 7; 19 Productus gratiosus Waagen; Mansuy, 1913, p. 115, pl. 13, ®g. 1. Productus gratiosus Waagen; Chi-Thuan, 1961 p. 276, pl. 5, ®gs. 8 and 9. Spyridiophora gubleri Termier and Termier, 1970a, p. 60, pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G±I. Lectotype. A holotype was not selected by Termier and Termier (1970a). Their specimen of conjoined valves (pl. 5, ®gs. 4±6) is here selected as the lectotype for this species. Description. About medium size for genus, 15.5 mm wide and about 9 mm long, widest at hinge and of similar width at mid-length; outline sub-rectangular with ears well ¯attened and ornamented; anterior margin broadly emarginate; pro®le gently concave; trail short; geniculation weak, restricted to median region between ¯anks; median fold notably shallow, poorly delimited on visceral disc, very broad on anterior trail. The umbonal region is missing, so true length and height are uncertain. Ornament distinctively reticulate; radial costae strong, coarse, rounded, of even width, converging slightly in the sulcus, numbering about 16 at mid-length, increasing occasionally by bifurcation; intertroughs of similar width, rounded; concentric rugae strong, but ®ner than costae; external spines not de®ned. Remarks. The present dorsal specimen is damaged but is suf®ciently well preserved to retain the main characteristics of Spyridiophora gubleri. The Bera form is almost identical to all the Cambodian shells ®gured by Mansuy (1913, pl. 13, ®g. 1), Chi-Thuan (1961, pl. 5, ®gs. 8 and 9) and Termier and Termier (1970a, pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G± I). In addition, Colani (1919, p. 10, pl. 1, ®g. 2) re-illustrated the Mansuy's (1913) Spyridiophora specimen named Productus gratiosus for comparison with her Vietnamese gratiosus; the latter is probably a species of Transennatia. Spyridiophora is known to bear the distinctive elevation

Fig. 7. 1±10, 13±15, Transennatia termierorum sp. nov.; 1±4, ventral internal mould Ð ventral, lateral (right), anterior, and lateral (left) views, £ 3.0, UKMF266, 5±8, dorsal external mould with accompanying natural cast of shell remaining anteriorly Ð posterior, ventral, anterior, and lateral views, £ 3.0, UKMF267, 9±10, dorsal external mould Ð posterior and anterior views, £ 3.0, UKM-F268, 13±15, dorsal external mould Ð ventral, anterior, and posterior views, the last showing a hole corresponding to the ventral beak, £ 2.0, UKM-F269, Assemblage A. 11±12, Transennatia sp. indet., dorsal external mould, dorsal internal mould resting on ventral external mould, £ 3.0, UKM-F270, Assemblage C. 16±17, Spinomarginifera sp., dorsal internal mould and rubber cast, £ 3.0, UKM-F271, Assemblage A. 18, Echinauris sp., rubber cast displaying dorsal disc interior, £ 2.0, UKM-F272, Assemblage A. 19, Spyridiophora gubleri Termier and Termier, incomplete dorsal external mould, £ 3.0, UKM-F273, Assemblage A. 20±23, Spiriferinid family indet., incomplete mould of a ventral interior Ð ventral, lateral, posterior and dorsal views, the last showing interarea, £ 3.5, UKM-F274, Assemblage C.

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Fig. 8. 1±11, Kozlowskia sp.; 1±5, ventral internal mould of semi-mature shell (Arrows indicate spine bases.) Ð ventral, posterior, lateral (right and left) and anterior views, £ 2.5, UKM-F275, 6±11, internal mould of conjoined shells Ð ventral, posterior, lateral (right) and anterior views of ventral interior, and dorsal view and rubber cast of dorsal interior, £ 2.0, UKM-F276, Assemblage A. 12±14, Linoproductus sp.; 12, ventral internal mould, £ 1.5, UKM-F277, 13, ventral internal mould, £ 1.5, UKM-F278, 14, ventral internal mould, £ 1.5, UKM-F279, Assemblage B. 15, Rhynchonellid family indet, incomplete ventral internal mould, £ 2.0, UKM-F280, Assemblage B. 16±17, Phricodothyris sp., rubber cast of ventral exterior, £ 2.0, enlarged section showing microornament, £ 6.0, UKM-F281, Assemblage A.

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of the brachial adductor scars, called a `spyridium' (term proposed by Cooper and Stehli, 1955, p. 472). It is well developed in the type species, S. distincta Cooper and Stehli from West Texas. By comparison, the spyridium of S. gubleri, as illustrated by Termier and Termier (1970a, pl. 5, ®g. 6), is less salient. Grant (1976, p. 136) regarded the Chi-Thuan's (1961) materials as Transennatia instead of Spyridiophora, but made no mention of the Termier and Termier (1970a) designation. Waterhouse (1978, p. 119) compared the Mansuy's (1913) material to his Nepalese Transennatia sp., but does not seem to have accepted the Termiers' work. However, considering the fact that the Termiers' material clearly displays the typical spyridium, the present form is placed in Spyridiophora without hesitation. Productus craticulatus Reed (1931, p. 11, pl. 1, ®gs., 1 and 2) from the Neoschwagerina craticulifera bed of the Bamian Limestone, Afghanistan, obviously belongs to Spyridiophora, judged from the description and illustrations. It appears to be close to S. gubleri, especially as regards its ornament. The South-East Asian forms are more transverse than the Afghan form, but this may be no more than within the same species. Because of its distinctive reticulation and transverse outline, Productus margaritatus Mansuy (1913, p. 28, pl. 2, ®g. 6) from Kham-keut, Laos, is possibly a species of Spyridiophora. The same species was illustrated by Huang (1932, p. 30, pl. 1, ®gs. 22±24) from the Late Permian of Guizhou, South China. This species has ®ner reticulation than S. gubleri. Grant (1976, p. 136) noted that this species is unlike Transennatia. Liang (1990) proposed two species of Spyridiophora, namely S. huadongensis (p. 159, pl. 24, ®gs. 1±21) and S. plegma (p. 161, pl. 24, ®gs. 22 and 23), both from the Lengwu Formation. They both differ from S. gubleri in having much ®ner ornament. Occurrence. S. gubleri has previously been found only in the Sisophon Limestone at Phnom Tup in the Sisophon region (Mansuy, 1913; Chi-Thuan, 1961), and at Phnoms Anseh, Totung and Takream in the Battambang region (Termier and Termier, 1970a). Nakamura (1979a) also recorded this species in Member C of the same formation. Species of Spyridiophora are extremely rare in the Tethyan Province. They have been recognised only in the Early to Middle Permian of South China, Indochina, Afghanistan and the Carnic Alps. Tribe Kozlowskiini Brunton et al., 1995 Genus Kozlowskia Fredericks, 1933 Kozlowskia sp. Fig. 8; 1±11 Remarks. This species is represented by two specimens of mature and semi-mature shells. The shells are slightly large for the genus, 20 mm wide, 15 mm long and the disc is 8 mm high for the mature specimen UKM-F276. The body cavity is of moderate size. The cardinal process is sessile, and the

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breviseptum is short. The brachial ridges are located very distally. Two small dorsal endospines are discerned. Generic assignment to Kozlowskia is con®rmed by the distinctive pattern of the ventral strut spines. Three large stout spines are seen on the ventral valve of the semi-mature specimen UKM-F275 Ð one on each lateral slope (Fig. 8; 2±4) and one on the anterior of the disc (Fig. 8; 5). This is further supported by the appearance of the dorsal interior, speci®cally the strong ear baf¯es, large pair of adductors, high breviseptum and dorsal marginal ridge forming a wide platform. The indistinct or absent sulcus and very weak ventral ornament also characterise the genus. Two species of Kozlowskia, K. cornuta Grant (1976, p. 118, pl. 26, ®gs. 24±29; pl. 27, ®gs. 35±38) and K. opipara Grant (1976, p. 121, pl. 28, ®gs. 1±27), were found in the Rat Buri Limestone of southern Thailand. Their dorsal interiors differ from the Bera form. Grant (1976, p. 118) noted that species of Kozlowskia were restricted to the Pennsylvanian and Early Permian (Wolfcampian); his two Thai species were the only Asian occurrences known at the time of his study. Since then, only two additional Asian occurrences have been reported, both from Early Permian horizons in Xinjiang, northwestern China (Wang and Yang, 1993; Wang, 1995), both being reports of the type species, K. capacii d'Orbigny. Waterhouse (1981, p. 13) indicated that Marginifera sensu Diener (1897, pl. 4, ®gs. 11±13; pl. 5, ®gs. 1 and 2) from the Chitichun Limestone (Kalabaghian) of Tibet may be Kozlowskia. However, as con®rmed by Waterhouse (written communication, 24 July 1999), this was in error because he previously discussed the Diener's specimens in terms of Marginifera, as a species separate from the type species, M. typica (see Waterhouse and Piyasin, 1970, p. 127; Waterhouse, 1978, p. 30). Marginifera and Kozlowskia often resemble each other considerably, as pointed out by Cooper and Grant (1975, p. 969). The two genera, however, can be differentiated by the very different spine patterns on the ventral valve. They are now classi®ed into the different families by Brunton et al. (1995). Superfamily Linoproductoidea Stehli, 1954 Family Linoproductidae Stehli, 1954 Subfamily Linoproductinae Stehli, 1954 Genus Linoproductus Chao, 1927 Linoproductus sp. Fig. 8; 12±14 Remarks. Three impressions of the ventral interior display characteristic costellae of Linoproductus. The shells are medium to large for the genus, ranging 29±40 mm wide and 30±40 mm long. They are semi-ovate to semi-elongate in outline, and lack a median fold. Costellae are irregular and uneven, numbering 7±8 in 5 mm medianly. Rugae are present only on the lateral trail as broad wrinkles. Parallel transverse lines intersect costellae in the largest specimen UKM-F278 (Fig. 8; 13).

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Spiriferinid family indet. Fig. 7; 20±23 Remarks. A single mould of a ventral interior is the only spiriferinid in the Bera fauna. It is small, spiriferiform, with simple, strong plicae indicating Spiriferinidina. The shell is about 9.5 mm wide, 7.5 mm long and 4.5 mm high in size, and has three ribs on each ¯ank. Acknowledgements Fig. 9. Comparison of microornament among three Phricodothyris species (all illustrations £ 25), 1, Phricodothyris sp. in this paper, 2, P. pyriformis Pavlova (1969, text-®g. 58), 3, P. rotiformis Liang (1990, pl. 63, ®g. 23).

Class Rhynchonellata Williams et al., 1996 Order Rhynchonellida KuÈhn, 1949 Rhynchonellid family indet. Fig. 8; 15 Remarks. A poorly preserved impression of a ventral interior is the only rhynchonellid in the Bera fauna. It is small (approximately 20 mm wide and 16 mm long), and has simple, weak costae Ð four in the sulcus and ®ve on each ¯ank. Order Spiriferida Waagen, 1884 Suborder Delthyridina Ivanova, 1972 Superfamily Reticularioidea Waagen, 1884 Family Elythidae Fredericks, 1924 Subfamily Phricodothyridinae Caster, 1939 Genus Phricodothyris George, 1932 Phricodothyris sp. Fig. 8; 16±17; 9; 1 Remarks. One incomplete ventral specimen was available. The shell is sub-orbicular, medium-sized (30 mm wide and about 22 mm long), with double-barrelled (biramous) spinebases, suggesting location in Phricodothyris. It is broadly convex without sulcation, with its umbo moderately high and obtuse at about 908. The spine arrangement of the Bera shell is similar to that of P. pyriformis Pavlova (1969, p. 93, pl. 8, ®gs. 1±3) (Fig. 9; 2), but differs slightly in that the Bera species has a third row of spine bases (Fig. 9; 1) and slightly coarser concentric laminae. P. rotiformis Liang (1990, p. 290, pl. 63, ®gs. 17±23) from the lower Lengwu Formation, South China, is also close; it seems to have a similar spine arrangement of three rows (Fig. 9; 3). Only a mould of the micro-ornament was illustrated for the Chinese species; the exact shape of its spines is, therefore, uncertain. Order Spiriferinida Ivanova, 1972 Suborder Spiriferinidina Ivanova, 1972

M. Sone's research was carried out for his B.Sc. Honours degree at Deakin University in 1998±99, and was ®nanced by his parents, Masao and Toshiko Sone. He is indebted to Prof. N.W. Archbold, Dr. S. Shen and Mr. Z. Chen for their invaluable discussions. Drs. J.B. Waterhouse, I. Metcalfe and Prof. J.A. Talent are thanked for critically reading the manuscript. Dr. Ahmad Jantan and Prof. Ibrahim Komoo are acknowledged for providing ®nancial supports from their government IPRA Projects 02±02±02±0004 and 02±02± 02±0015, respectively. This paper was completed while M. Sone was supported by a fellowship of LESTARI. References Archbold, N.W., 1984. Western Australian occurrences of the Permian brachiopod genus Retimarginifera. Alcheringa 8 (1/2), 113±122. Archbold, N.W., Bird, P.R., 1989. Permian Brachiopoda from near Kasliu village, West Timor. Alcheringa 13 (1/2), 103±123. Broili, F., 1916. Die permischen Brachiopoden von Timor. In: Wanner, J. (Ed.), PalaÈontologie von Timor 7, Stuttgart, pp. 1±104. Brunton, C.H.C., Lazarev, S.S., Grant, R.E., 1995. A review and new classi®cation of the brachiopod order Productida. Palaeontology 38 (4), 915±936. Brunton, C.H.C., Lazarev, S.S., 1997. Evolution and classi®cation of the Productellidae (Productida), Upper Paleozoic brachiopods. Journal of Paleontology 71 (3), 381±394. Carter, J.L., Johnson, J.G., Gourvennec, R., Hou, Hong-fei, 1994. A revised classi®cation of the spiriferid brachiopods. Carnegie Museum, Annals 63 (4), 327±374. Caster, K.E., 1939. A Devonian fauna from Columbia. Bulletin of American Paleontology 24 (83), 1±218, pls. 1±14. Chao, Y.T., 1927. Productidae of China, Part 1: Producti. Palaeontolgia Sinica, Series B 5 (2), 1±244, 16 pls. Chi-Thuan, T.T., 1961. Les brachiopodes permiens du Phnom-Tup (Sisophon Ð Cambodge). Annales de la Faculte des Sciences, Universite de Saigon 1961, 267±308, 8 pls. (in French). Ching, Y., 1963. Urushtenia from the Lower Permian of China. Acta Palaeontologica Sinica 11 (1), 1±31, 2 pls. (in Chinese with English abstract). Colani, M., 1919. Sur quelques fossiles Ouralo-Permiens de Hongay. Service GeÂologique de l'Indochine, Bulletin 6 (5), 1±27, 2 pls. (in French). Cook, R.H., Suntharalingam, T., The geology and mineral resources of Pahang Tenggara (Sectors ªEº and ªFº). Geological Survey Malaysia Annual Report 1970, pp. 104±116. Cooper, G.A., Grant, R.E., 1974. Permian brachiopods of West Texas, II. Smithsonian Contributions to Paleobiology 15, 233±793, pls. 24±191. Cooper, G.A., Grant, R.E., 1975. Permian brachiopods of West Texas, III (Part 1 and 2). Smithsonian Contributions to Paleobiology 19, 794± 1920, pls. 192±502.

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