Middle Permian Brachiopods from Central Peninsular Malaysia Ð Faunal Af®Nities Between Malaysia and West Cambodia

Journal of Asian Earth Sciences 19 (2001) 177±194 www.elsevier.nl/locate/jseaes

Middle Permian brachiopods from central Peninsular Malaysia Ð faunal af®nities between Malaysia and west Cambodia

Masatoshi Sonea,1,*, Mohd Shafeea Lemanb, Guang R. Shia

aSchool of Ecology and Environment, Deakin University, Rusden Campus, Clayton, Vic. 3168, Australia bSchool of Environmental Sciences and Natural Resources, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia 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 ¯anked on the east by ?Jurassic to Cretaceous continental sediments of the Bertangga Sandstone, but exposures to east This paper describes the largest known Permian brachio- and west are poor (Leman et al., in press). The northern and pod fauna from the so-called Central Basin (or the Central southern boundaries of the Bera Formation are also uncer- Belt) of Peninsular Malaysia, and reports on the southern- tain, but in the north it is associated with an andesitic vol- most occurrence of such a fauna in that region. Most of the canic unit thought to be of a Late Permian age (Cook and fossils examined in this study were collected by Sone and Suntharalingam, 1970). The occurrence of the fusulinid, Leman in the Bera District in February 1998. Additional Parafusulina sp., from a small limestone outcrop near material came from a collection in the palaeontology labora- Tasik Bera implies that the lower part of the Bera Formation tory of the National University of Malaysia (Universiti may extend down into the Kungurian (upper Early Permian) Kebangsaan Malaysia), collected originally by Leman. All (Cook and Suntharalingam, 1970). described specimens are registered in the same university with pre®x UKM-F. The regional geology of the Bera District is known from 2. Stratigraphy only two brief reports (Cook and Suntharalingam, 1970; MacDonald, 1970). The rock unit from which the current The fossils of this report come from the ªSungai Bera fossil horizon is studied has been named the Bera Formation sectionº (grid reference WF151446; the National Map (Leman et al., in press). The area where it occurs was Malaysia 1:50 000 Sheet 4157) exposed on the eastern previously considered to be part of the Triassic Semantan side of Bera Road, 17.3 km south of Felda Sebertak junction Formation Ð for example, on the most recent of®cial geolo- (Fig. 1). This section is equivalent to Locality BF2 of gical map of Peninsular Malaysia (Geological Survey of Leman et al. (in press). The outcrop was originally prepared Malaysia, 1985). The Semantan Formation, as revised, for construction of a resort. The exposed strata extend outcrops west of the Bera Formation, which in turn is approximately 90 m vertically and 300 m laterally, and appear to have undergone gentle but pervasive soft-sedi- * Corresponding author. ment deformation westwards. It is thus dif®cult to trace E-mail address: [email protected] (M. Sone). the sequence from one side to the other. The general strike 1 Present address: Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, is N50±708W and dip 70±908SW. The lithology consists Malaysia. mainly of mudstone±siltstone interbedded with sandstone,

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 brachiopod faunas in the Neoschwagerina-Yabeina Zones section. The surface is bleached soft sediment. More of the Middle Permian, most likely to a lower Capitanian, outcrops occur along the road; these are much smaller This is based on the three brachiopod species shared with than the Sungai Bera section. Member C (Yabeina beds) of the Sisophon Limestone, west Three major fossiliferous stratigraphic intervals can be Cambodia. discriminated; these are referred to as Assemblages A±C The Permian strata of west Cambodia are exposed mainly in ascending order (Fig. 2). Assemblage A is predominantly in the Sisophon and Battambang regions (Fig. 4), as lime- a coquina of shell debris in grey mudstone; it often contains stone hills with intercalations of tuff, shale and mudstone fossiliferous calcareous concretions. This assemblage holds strata, collectively called the Sisophon Limestone (Ishii et the richest brachiopod deposit in this report. Assemblage B has less common brachiopods in nearly white, ®ne- to Table 1 medium-grained sandstone. Assemblage C consists princi- Brachiopod species from each assemblage of the Sungai Bera section pally of abundant lyttoniids in purple to black tuffaceous Species Assemblages mudstone and siltstone. The brachiopods are listed in Table 1. The genus Trans- ABC ennatia is the most abundant, although most specimens are Derbyia sp. X too poorly preserved for satisfactory illustration. The three Urushtenoidea chaoi (Ching, 1963) X X assemblages share few taxa, and may appear to represent Urushtenoidea sp. X distinct faunules, but sampling was not exhaustive. The Transennatia termierorum sp. nov. X three assemblages are considered as a single fauna in the Transennatia sp. indet. X following discussion. Spinomarginifera sp. X Echinauris sp. X Spyridiophora gubleri Termier and Termier, 1970a X Kozlowskia sp. X 3. Correlation and age Phricodothyris sp. X Orthotetes aff. picta Fang, 1983 X At present, there is no globally accepted Permian chron- Strophalosiina sp. X Linoproductus sp. X ological standard. The time-scale compiled by the Interna- Rhynchonellid family indet. X tional Subcommision on Permian Stratigraphy (Jin et al., Orthothetina cf. iljinae Sokol'skaya, 1965 X 1997) is applied in this study, with slight modi®cation to Gubleria aff. ninglangensis Fang and Jiang, 1992 X fusulinid biozonation for more appropriate correlation Gubleria sp. X within South-East Asia (Fig. 3). The brachiopod fauna of Eolyttonia? sp. X Spiriferinid family indet. X the Sungai Bera section appears to correlate with the other M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 179 species from their collections gained from that unit; Naka- mura (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 biostra- tigraphic 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®n- able for Member C of the Sisophon Limestone. The most important species in the Bera fauna is Urushte- noidea 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. Transenna- tia 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-Cauca- sus and in the Maokou horizons of South China. In particu- 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). lar, the species of Orthotetes and Gubleria are alike to those in west Yunnan, southwestern China. Additionally, the Bera al., 1969). Brachiopods from the limestone have been fauna displays some similarity to the Capitanian brachio- studied by several palaeontologists (e.g., Mansuy, 1913, pods of the Lengwu Formation, Zhejiang Province, South 1914; Chi-Thuan, 1961; Termier and Termier, 1960, China (see Liang, 1990). Almost all Bera genera occur in 1970a, 1970b). Ishii et al. (1969) listed 50 brachiopod the Lengwu fauna, including relatively rare taxa such as 180 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

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. Limestone Member, S & S. 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 condi- tions, 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 post- Kuhfengian 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 Fig. 4. Possible distributions of the Indochina and East Malaya terranes, and never been unequivocally reported from Late Permian locations of the Bera District and the Sisophon and Battambang regions: 1, rocks. Their youngest occurrence may in fact be in the Uttaradit-Nan Suture; 2, Raub-Bentong Line (modi®ed after Metcalfe, Lengwu Formation. 1998). M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 181 There is a generally accepted view that large lyttoniids Orthothetina, Transennatia, Gubleria, Spinomarginifera, such as Leptodus and Gubleria are typically Late Permian Urushtenoidea and Strophalosiina, all of which are typical (e.g., Shen and Shi, 1996) and, in the case of Oldhamina, Tethyan elements. The close af®nities between the Bera and this seems to be true. It is only partly true for brachiopod Sisophon faunas were discussed above. Relationships with faunas from the Trans-Caucasus and South China, where the Middle Permian brachiopods of South China are less the Lengwu fauna is now known to contain numerous lytto- obvious. Differences in Middle Permian brachiopods niids, but notably without Oldhamina. Many lyttoniid forms between Indochina and South China have been pointed occur in the Middle Permian of Indochina (e.g., Mansuy, out by Nakamura et al. (1985). It is, therefore, assumed 1912, 1913, 1914; Termier and Termier, 1960; Chi-Thuan, that the Central Basin with the Bera fauna was located in 1961), and in the Neoschwagerina horizon of the Sosio the same or nearby climatic zone as west Cambodia during Limestone, Italy (e.g., Rudwick and Cowen, 1967). Lyt- the age-interval in question, but that there were slight differ- toniids are in fact more common in the Middle rather than ences in ecology or geographic position relative to tropical Late Permian of Japan, with several reports from the lower South Chinese Cathaysia, hindering faunal interchange. Kanokuran Series (e.g., Tazawa, 1976, 1987; Tazawa and This conclusion is consistent with the tectonic model of Matsumoto, 1998). The presence of numerous lyttoniids in Metcalfe (1996a, 1996b, 1998), namely, that the East Assemblage C is thus not necessarily indicative of a Late Malaya terrane accommodating the Central Basin was part Permian age; it may in fact be the Middle Permian. of the Indochina terrane (Fig. 4). 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 5. Systematic palaeontology (M. Sone) and/or radiolarians. In contrast, only a few Permian brachio- The systematic study follows the supra-ordinal classi®ca- pods have been described, despite many reported occur- tion of Williams et al. (1996), and the classi®cations of rences (e.g., Jones et al., 1966; Gobbett, 1973; Fontaine et Williams and Brunton (1993) for the Strophomenida, Brun- al., 1994). Igo (1964) and Nakamura (in Nakazawa, 1973) ton et al. (1995) and Brunton and Lazarev (1997) for the described some poorly preserved brachiopods of little bio- Productida, and Carter et al. (1994) for the Spiriferida and stratigraphic value. Yanagida and Aw (1979) described a Spiriferinida. The suborder Strophalosiidina Schuchert, small suite of Permian brachiopods from Kelantan; they 1913 refers to Brunton (written communication, 27 January considered the fauna to be late Djul®an. 2000). The so-called ªLeptodus Shaleº fauna in north-central Pahang has been known for over a half century (Muir- Class Strophomenata Williams et al., 1996 Wood, 1948, p. 5, footnote; Jones et al., 1966). Leman Order Strophomenida OÈ pik, 1934 (1993, 1994) reported 49 species of brachiopods from the Suborder Orthotetidina Waagen, 1884 same fauna, but they have not been described. Based on Superfamily Orthotetoidea Waagen, 1884 the presence of abundant lyttoniids, the fauna was thought Family Orthotetidae Waagen, 1884 to be Late Permian, but this is questionable. Thus, based Subfamily Orthotetinae Waagen, 1884 on brachiopods, it is still dif®cult to make precise correla- Genus Orthotetes Fischer de Waldheim, 1829 tions within the Central Basin. Nevertheless, considering Orthotetes aff. picta Fang, 1983 available data from palaeontology and regional geology, Fig. 5; 1±2. 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 lime- cf. Orthotetes picta Fang, 1983, p. 94. pl. 1, ®gs. 4±6. stones yield Yabeina asiatica which is known elsewhere only in the Sisophon Limestone (Ishii, 1966; Igo, 1967; Description. Ventral shell medium for genus, 27 mm wide Ishii et al., 1969; Fontaine et al., 1994). North of the Bera and 26.5 mm long with greatest width at hinge; outline U- District, small outcrops of late Middle Permian age have shaped; pro®le fairly ¯at; ears not demarcated; interarea been studied in recent years (e.g., Fontaine et al., 1994; very low; umbo inconspicuous; rectimarginate; plications Jasin et al., 1995). There are thus prospects for better absent. Ornament of moderate costellae with angular crests, understanding of Permian correlations in central Pahang, increasing by intercalation, numbering approximately 18 speci®cally with respect to the Bera brachiopod fauna. 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 4. Palaeogeographic implications the end of the hinge. Ventral interior with long median septum, extending to nearly mid-valve, slightly thickened All taxa identi®ed to generic and speci®c levels accord anteriorly, height unknown; spondylium minute, laterally with this being a warm-water fauna. This is indicated by semi-ovate; dental ridges uniting to form a spondylium; 182 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 183 adductor ®eld weakly concave. Details of dorsal valve Orthothetina cf. iljinae Sokol'skaya, 1965 unknown. Fig. 5; 5±6 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 commis- cf. Meekella cf. baschkirica Tschernyschew; Mansuy, sure makes it close to O. picta Fang (1983, p. 94. pl. 1, ®gs. 1914, p. 22, pl. 2, ®g. 15. 4±6) from the Neoschwagerina horizon of the Xiaoxinzhai cf. Orthothetina iljinae Sokol'skaya, 1965, p. 204, pl. 30, section in Gengma, west Yunnan. There are de®nable, but ®gs. 1a and b, 2. subtle differences in the condition of the costellation Ð the cf. Orthothetina iljinae Sokol'skaya; Tong, 1978, p. 214, Yunnan species displays slightly coarser (7±8 in 5 mm) and pl. 77, ®gs. 9a±c. more intercalated costellae. This, however, may be due to differing stages of their maturity; the larger Bera valve Description. Shell medium sized, approximately 46 mm seems older ontogenetically with more growth lines. It is wide and 47 mm long, with greatest width at mid-valve; thought to be close to O. picta, but additional material is outline sub-pyramidal; pro®le gently convex; ¯anks required for con®dent identi®cation. No other previously smoothly rounded; sulcus and plications absent; umbonal described species appears close to the present form. region ¯attened, not recurved. Ornament of radial capillae, moderately strong, even in width, numbering 18±20 per 10 Family Derbyiidae Stehli, 1954 mm at anterior margin. Concentric crenulation present near Subfamily Derbyiinae Stehli, 1954 the frontal margin, 4 mm in maximum width Ð growth Genus Derbyia Waagen, 1884 lamella otherwise not de®ned on surface. Ventral interior Derbyia sp. with pair of dental plates, high, arising directly from valve Fig. 5; 3±4 ¯oor, extending at least one-third of valve-length, gently divergent internally, slightly converging apically. Dorsal Remarks. The ventral shell with a high and thick median characters unknown. septum bisecting the umbo is indicative of Derbyia, but it is Remarks. The non-plicate ventral valve with strong, small for the genus, almost equidimensional (about 23 mm parallel dental plates clearly indicates Orthothetina.The wide to 22 mm long), and the hingeline is the widest part. present form is characterised by a sub-pyramidal outline The outline is U-shaped, the ears are not differentiated, and and low in¯ation. It closely resembles O. iljinae the pro®le is convex in the median region, but contrastingly Sokol'skaya (1965, p. 204, pl. 30, ®gs. 1a and b, 2) from ¯at on the ¯anks. The interarea is low and narrow, and the Gnishik and Khachik Formations of the Trans-Caucasus cardinal extremities are obtuse. The umbo is prominent. by lacking growth lines but instead having a de®ned The shell is rectimarginate without plications, and is undis- concentric elevation anteriorly. O. iljinae, as illustrated by torted. The septum is relatively short. Costae are coarse, low Tong (1978, p. 214, pl. 77, ®gs. 9a±c) from the Limestone and blunt, increasing by intercalation, with width fairly Member of the upper Chihsia Formation, Guizhou, South uneven. China, has somewhat more ¯attened shells than the Trans- The present species is characterised by the umbonal Caucasian form. region being prominent, raised and demarcated by a strong O. phetchabunensis Yanagida (1964, p. 14, pl. 2, ®gs. 5 concentric line. D. altestriata Waagen illustrated by Fantini and 6) from the upper Middle Permian of central Thailand is Sestini (1965, p. 38, pl. 3, ®g. 7) from the upper Ruteh similar in outline and size, but its costellation is slightly Limestone, north Iran, displays a very similar elevated stronger than the Bera form. The Thai species displays umbo with a primary lamina. It differs from the Bera form concentric rugae over the ventral surface, a feature lacking in possessing a more transverse outline and ¯attened median in the present form. A Cambodian shell illustrated by region. Mansuy (1914, p. 22, pl. 2, ®g. 15) as Meekella cf. baschkirica Tschernyschew from Phnom Takream comes close to Family Meekellidae Stehli, 1954 the Bera shell in having a sub-pyramidal outline and lacking Subfamily Meekellinae Stehli, 1954 concentric lines. The specimen is, however, much smaller Genus Orthothetina Schellwien, 1900 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. 184 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 shows no plication, hence is more likely Orthothetina rather ¯at on disc, trail about 2 mm. Radial costae well marked on than Meekella. trail but poor on disc, numbering about 40; terminations of dorsal costae developing into hollow spines anteriorly along Order Productida Sarytcheva and Sokol'skaya, 1959 edge of re¯exed marginal ledge; spines incurved dorsally at Suborder Strophalosiidina Schuchert, 1913 approximately 35±408; concentric wrinkles poorly de®ned Superfamily Aulostegoidea Muir-Wood and Cooper, on disc; concentric laminar delicate over dorsal trail, dense 1960 and regular (Fig. 5; 9±11). Dorsal interior with robust bi®d, Family Aulostegidae Muir-Wood and Cooper, 1960 sessile cardinal process; lobes sub-ovate, large; alveolus Subfamily Chonosteginae Muir-Wood and Cooper, 1960 absent; posterior platform weakly in¯ated; buttress plates Genus Urushtenoidea Jing (Jin) and Hu, 1978 absent; muscle scars large, palmate, hanging anteriorwards; breviseptum narrow and long; brachial ridges weak and Discussion. Urushtenoidea has a complicated spine arrange- small; ear baf¯es strong; marginal rim moderately elevated. ment on the anterior margin of both valves. The genus has In specimen UKM-F263 (Fig. 5; 16±18), moulds of considerable similarity to Urushtenia Likharev. The former, dorsal marginal endospines (`marginal fence spines' of however, differs from the latter most obviously in having Zeng, 1987, p. 502) individually appear as cylindrical scaly concentric lamellae (composed of spinous structure) hollows lying on the interior of costal crests. A row of on the ventral trail. The ventral geniculation is much stron- endospines collectively forming a high `marginal fence' ger in Urushtenoidea; its angle often exceeds 608. Urushte- (term from Jing (Jin) and Hu, 1978, p. 258) behind the noidea has a ¯attened dorsal disc with a poorly de®ned trail, ventral trail. The spines are laterally supported by the whereas Urushtenia has a geniculate dorsal valve with a band of the `marginal fence transverse ridge' (term from moderately long trail. Species of Urushtenoidea commonly Zeng, 1987, p. 502) at mid-length of spines Ð appearing show ®ner costae than Urushtenia. as a fracture in this specimen. Occurrence. Urushtenoidea has hitherto been recorded Remarks. Three shells (one dorsal and two ventral) were only from the eastern Tethys of South China, Cambodia, available. The present form is referred to Urushtenoidea Laos and Japan. This report con®rms the presence of the mainly because of its short dorsal trail and spinous nature genus in Peninsular Malaysia. Urushtenoidea is limited to of the ventral trail. It is broadly similar to the type species, the Xiangboan to Kuhfengian Stages (Roadian to Wordian) U. chaoi (Ching). There are minor differences from the of South China, Members A±C of the Sisophon Limestone, Chinese form. The Bera form has slightly ®ner costae and the Parafusulina-Neoschwagerina horizons of Laos, and the a less strongly elevated posterior platform. The present form Lower Kanokuran Series of Japan, all of which are corre- is very much like the Cambodian form, U. chaoi of Zeng lated with the Guadalupian Series. (1987), in having comparable hanging palmate-shaped adductor scars. Zeng's materials are from the Neoschwager- Urushtenoidea chaoi (Ching (Jin), 1963), ina margaritae bed of the Sisophon region. Jing (Jin) and Fig. 5; 7±14, 16±18 Hu (1978), in de®ning the genus, referred to it having a large alveolus. This feature is not developed on the present mate- Urushtenia chaoi Ching, 1963, p. 15, pl. 1, ®gs., 1±4, 9± rial, and is not always apparent in other congeneric Chinese 12, 16; pl. 2, ®gs. 7±8, 11±17. or Cambodian species. cf. Urushtenia chenanensis (Chan); Tong, 1978, p. 218, Occurrence. U. chaoi has been reported from Member C pl. 78. ®gs. 16a±c. of the Sisophon Limestone, Cambodia, and the Maokou Urushtenia chaoi Ching; Tong, 1978, p. 218, pl. 78, ®g. Formation, Kuhfeng Formation, and Hsiaochiang Lime- 18a±d. stone of South China, all horizons correlative with the Urushtenoidea chaoi (Ching); Jing (Jin) and Hu, 1978, p. Neoschwagerina-Yabeina Zones. 116, pl. 2, ®g. 10. Urushtenoidea chaoi (Ching); Nakamura, 1979b, p. 230, Urushtenoidea sp. pl. 2, ®g. 4. Fig. 5; 15 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 maceus (Ching); Nakamura, 1979b, p. Urushtenoidea chaoi (Ching); Zeng, 1987, pl. 1, ®gs. 1± 227, (not pl. 1, ®gs. 1±4), pl. 2, ®gs. 1±3. 5, 8±14, 16±19, 26±27, 29±30; pl. 2, ®gs. 4, 7, 9. Remarks. One dorsal valve referred to Urushtenoidea has a Description. Shell average- to large-sized for genus, 19.5± short dorsal trail (1.5 mm). It is medium-sized for the genus 23 mm wide and 16±18 mm long, with maximum width at (21 mm wide and 12 mm long). It clearly differs from U. mid-length; outline cylindrical; ears not differentiated; chaoi in possessing a more transverse outline and slightly median fold not well de®ned. Ventral valve moderately coarser costae. Its cardinal process lobes are globular in geniculate; trail long; umbo unknown. Dorsal valve almost shape, unlike the semi-oval shape of those in U. chaoi, M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 185 suggesting that this is a second species of Urushtenoidea in 162), however, asserted that Gubleria is invalid, being Assemblage A. Its raised ventral ears are well impressed on immature, or a variation of Leptodus Kayser. the mould. In this study, on the contrary, numerous specimens of Three specimens illustrated by Nakamura (1979b, p. 227, large lyttoniids, thought to belong to two species, were pl. 2, ®gs. 1±3) as U. maceus (Ching) each from Members collected. Almost all specimens possess more or less devel- A, B and C of the Sisophon Limestone are close to the oped median incisions or segments, indicative of Gubleria, present form in outline and costae, and are probably conspe- whereas no form clearly assignable to Leptodus was found. ci®c. In addition, Jing (Jin) and Hu (1978) placed U. maceus This seems to contradict to the view of Cooper and Grant in Uncisteges, but Nakamura (1979b) has not accepted this. (1974) that Gubleria has individual rather than a generic Urushtenia khmeriana Termier and Termier (1970b, p. 446, characters. text-®gs. 2A±D, pl. 29, ®gs. 1±7) from Phnom Te Bon, Sisophon, has a ¯attened dorsal shell and scaly laminae on Gubleria aff. ninglangensis Fang and Jiang, 1992 its ventral trail. It is thus doubtlessly a species of Urushte- Fig. 6; 1±8 noidea. It has costae and raised ventral ears similar to those of the present form, but no dorsal view is available. cf. Gubleria ninglangensis Fang and Jiang, 1992, p. 327, pl. 1, ®gs. 3±5. Genus Strophalosiina Likharev, 1935 Strophalosiina sp. Description. Shell size varied, average width 19 mm and Fig. 5; 19 length more than 57 mm; conical, spatulate, weakly twisted; outline long and narrow, umbo strongly obtuse, gradually Remarks. A single mould of a dorsal exterior was available. expanding and ¯attening towards mid-length, then twisted The shell is large for the genus (26 mm wide and 18.5 mm anteriorly; lateral pro®le ¯at to slightly convex. Ventral long), transverse, with a semi-circular anterior margin, valve interior with obsolete muscle ®eld; median septum moderate geniculation. It has a well-de®ned dorsal trail highest medianly, faintly continuous, rough incisions (3.5 mm long) with no concentric wrinkles, unlike that of de®ned anteriorly; lateral septa symmetrical, short and Urushtenoidea. The stereozone is smooth, seemingly very thin, numerous Ð 4 to 5 in 10 mm, at least 18 for longest thick, wide and reaching the hinge; it is widest (2 mm) in but still incomplete specimen UKM-F287 (Figs. 6; 7); inter- front. Ornament of ®ne and sharp costae with weak and septal spaces much wider than septal ridges, deeply grooved irregular rugae are present on the disc but are absent from along septal margins. Dorsal internal plates ¯at and long, later- the trail. No dorsal external spines are discernible. ally grooved along margins; median trough segmented irregu- The large transverse shell with the wide and thick stereo- larly. No trace of papilla observed. Other details unknown. zone suggests Strophalosiina. The stereozone does not seem Remarks. Seven specimens (®ve ventral and two dorsal to have any re¯ected hollow spines or funnels, unlike those internal moulds) are identi®ed as a species of Gubleria by of other genera of Chonosteginae such as Urushtenia and possessing a discontinuous or segmented median line on Chonosteges. The present form shares several similarities both ventral and dorsal valves. This species is characterised with the type species, S. tibetica (Diener) from Tibet, by its elongate shape. G. ninglangensis Fang and Jiang Cambodia, Timor, Iran and the Trans-Caucasus. The Malay- (1992, p. 327, pl. 1, ®gs. 3±5) from the lower Emishan sian species, however, differs most obviously in being larger Basalt Formation (lower Capitanian), west Yunnan, is and having ®ner costae Ð at least 45 in number, whereas very close in outline and septal apparatus; it is believed to typical S. tibetica has about 30. S. zhejiangensis Liang be closely allied to the present form. Collemataria tilita (1990, p. 154, pl. 18, ®gs. 9±20) from the lower Lengwu Liang (1990, p. 229, pl. 40, ®g. 2; pl. 41, ®g. 11) from the Formation, China, is less transverse than the Bera form, and lower Lengwu Formation also has a similar outline with an has many large nodules on the dorsal disc. 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 Superfamily Lyttonioidea Waagen, 1883 by a couple of fragmentary internal plates. They are ®nely Family Lyttoniidae Waagen, 1883 papillose and show median holes that are irregular in size Subfamily Lyttoniinae Waagen, 1883 and shape. Other details are unknown, preventing useful Genus Gubleria Termier and Termier, 1960 comparison.

Comments. Gubleria was originally proposed by Termier Gubleria sp. and Termier (1960) with the type species, G. disjuncta Fig. 6; 9±12 from the Sisophon Limestone. The genus is characterised most importantly by the discontinuous nature of its ventral Remarks. Three ventral shells are assigned to Gubleria. The median septum and corresponding dorsal median holes. shell is medium sized (about 36 mm wide and more than 47 Cooper and Grant (1974, p. 411) and Grant (1976, p. mm long). The outline is ovate, with an umbonal angle 186 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

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). M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 187 ranging 80±1058, and the pro®le is fairly ¯at. They may outline sub-quadrate, widest part at hinge; ears large, belong to Gubleria aff. ninglangensis because of their simi- convex and costate; ventral umbo pointed; geniculation lar septal apparatus. The present material, however, is much abrupt with angle about 508; dorsal disc fairly ¯attened, wider and has greater umbonal angles. Specimen UKM- semi-circular in shape, almost without a fold, notably reti- F291 is problematic in that it seems to have an incurved culate; trail long. Surface coarsely ornamented by sturdy beak (?) pointing dorsally (Fig. 6; 11). costae, branching on the ¯anks, converging in the sulcus The present form super®cially recalls Leptodus nobilis and on the fold, with crests rounded, and similarly rounded, (Waagen). The chief differences, however, concern the wide and shallow interspaces; reticulation con®ned to disc, median septum. The Bera form displays a weakly raised concentric rugae faint on posterior part of the trail. Ventral median septum with segmentation. Mature individuals of interior with wide and ¯at platform, well elevated, extend- L. nobilis typically have a high, thick, continuous, blade- ing from apex; muscle scar obsolete or obscure. like median septum. This feature is well illustrated by Remarks. This form is characterised by its abrupt and Noetling (1905, pl. 17, ®gs. 1 and 2; pl. 18, ®gs. 1 and 2) strong geniculation and extended ears. It is most like Spyr- for specimens of L. nobilis from the Wargal and Chhidru idiophora ? timorensis Termier and Termier (1970a, pl. 5, Formations of the Salt Range, Pakistan. ®g. 1, text-®gs. 1±4) from Sisophon, Cambodia, but lacks the row of spine bases along the ears seen in the Cambodian Genus Eolyttonia Fredericks, 1924 material; this could be an artifact of preservation. Termier Eolyttonia? sp. and Termier (1970a) originally de®ned their timorensis as a Fig. 6; 13±14 synonym of Productus gratiosus Waagen, as illustrated by Broili (1916, pl. 2, ®gs. 4±5, 7±13), from the Basleo, Koeka Remarks. A fragment of a ventral internal mould exhibits bei Koeafeoe and Bitauni areas of Timor. However, Grant the distinctive ¯uting of its angustilobate septal apparatus, (1976, p. 134) asserted that only the Basleo specimens suggesting possible location in Eolyttonia. It is small (9 mm belong to Transennatia, the others being Retimarginifera. in width) for the genus, but its true length is uncertain. It is This view was followed by Archbold (1984, p. 115) and longitudinally conical in shape, narrow and slightly wider Archbold and Bird (1989, p. 108). Waterhouse (1978, p. anteriorly. The ventral interior has well-formed septa with 119) considered Termiers' timorensis to be a junior syno- crests broadly grooved and low. The lateral septa are trans- nym and homonym of Productus timorensis Hamlet. verse, and the interseptal troughs are rounded. It may be a T. timorensis is close to T. termierorum in size and orna- juvenile, although the internal structure is relatively well ment. The former can, however, be distinguished from the expressed. latter by having much smaller ears and a strongly folded dorsal disc. There is admittedly considerable variation Suborder Productidina Waagen, 1883 among the materials of T. timorensis as understood by Arch- Superfamily Productoidea Gray, 1840 bold and Bird (1989, p. 108). For instance, the Kasliu form Family Productellidae Schuchert, 1929 of Archbold and Bird (1989, p. 107, ®gs. 3E±I) is excep- Subfamily Marginiferinae Stehli, 1954 tionally large and has ®ne ornament, whereas the Basleo- Tribe Paucispiniferini Muir-Wood and Cooper, 1960 Wesleo forms of Hamlet (1928, pl. 2, ®gs. 2±4) have strong Genus Transennatia Waterhouse, 1975 ornament and a deep fold. The small specimen in ®g. 7 of Transennatia termierorum sp. nov. Diener (1897, pl. 3) identi®ed as P. gratiosus from the Chit- Fig. 7; 1±10, 13±15 ichun Limestone, Tibet, has a deep sinus and sturdy costae; it strongly resembles T. termierorum. This Tibetan material Spyridiophora ? timorensis Termier and Termier, 1970a, is possibly conspeci®c, although it is slightly wider in p. 58, pl. 5, ®g. 1, text-®gs. 1±4. outline. T. insculpta Grant (1976, p.135, pl. 32, ®gs. 1± 37; pl. 33, ®gs. 1±16) from the Rat Buri Limestone of south- Etymology. After Professors Henri Termier and GenevieÁve ern Thailand is similar. Its geniculation is, however, not as Termier, who greatly contributed to knowledge on the abrupt as that of the new species. T. termierorum differs South-East Asian Permian brachiopods, and ®rst described from the type species, T. gratiosa (Waagen, 1884, pl. 72, this species. ®gs. 3±7) from the Salt Range, Pakistan, most obviously in Type specimen. The specimen illustrated as Spyridio- having larger ears. The geniculation of T. gratiosa varies phora ? timorensis Termier and Termier (1970a, pl. 5, ®g. from shell to shell; smaller ones tend to be much less geni- 1) (not Hamlet) is designated as the holotype. culate. This trend is probably common in the genus; it Diagnosis. Shell medium-sized, geniculation abrupt, occurs also in T. insculpta Grant. The small but highly sulcus and fold strong anteriorly, ears proportionally large geniculate Bera shells may, therefore, be mature forms. and projecting, costae notably converging, branching, and Occurrence. This form was originally recorded from the sturdy on the trail, coarse reticulation present on the disc. Yabeina bed of a limestone hill, Phnom Takream, in the Description. Shell size ranging from 13±23.5 mm wide, Battambang region (Termier and Termier, 1970a, p. 58), 9±14.5 mm long, more than 8.5 mm high, commonly small; suggesting that its occurrence is con®ned to an upper part 188 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 189 of Member B to Member D, most likely Member C of the same species as Echinauris khmerianus from Member D of Sisophon Limestone. the Sisophon Limestone. Its interior is not known, hindering comparison with the Bera material, although the size is Transennatia sp. indet. close. Fig. 7; 11±12 Family Productidae Gray, 1840 Remarks. The small conjoined shells (9 mm wide and 7.5 Subfamily Productinae Gray, 1840 mm long) with no geniculation is probably a juvenile. It, Tribe Spyridiophorini Muir-Wood and Cooper, 1960 nevertheless, displays features diagnostic of the genus, such Genus Spyridiophora Cooper and Stehli, 1955 as strong reticulation, tiny ears and branching costae. It Spyridiophora gubleri Termier and Termier, 1970a occurs in Assemblage C. Fig. 7; 19

Tribe Marginiferini Muir-Wood and Cooper, 1960 Genus Spinomarginifera Huang, 1932 Productus gratiosus Waagen; Mansuy, 1913, p. 115, pl. Spinomarginifera sp. 13, ®g. 1. Fig. 7; 16±17 Productus gratiosus Waagen; Chi-Thuan, 1961 p. 276, pl. 5, ®gs. 8 and 9. Remarks. One squashed mould of a dorsal interior was Spyridiophora gubleri Termier and Termier, 1970a, p. 60, available. The distinctive appearance of its muscle ®eld pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G±I. suggests alliance with Spinomarginifera. The shell is small for the genus; its dorsal disc is about 13 mm wide Lectotype. A holotype was not selected by Termier and and 13.5 mm long. It seems to have a row of endospines Termier (1970a). Their specimen of conjoined valves (pl. around the anterior half of the disc (characteristic of the 5, ®gs. 4±6) is here selected as the lectotype for this species. genus) but this is poorly preserved. There is too little infor- Description. About medium size for genus, 15.5 mm wide mation for speci®c assignment. and about 9 mm long, widest at hinge and of similar width at mid-length; outline sub-rectangular with ears well ¯attened Subfamily Overtoniinae Muir-Wood and Cooper, 1960 and ornamented; anterior margin broadly emarginate; Tribe Costispiniferini Muir-Wood and Cooper, 1960 pro®le gently concave; trail short; geniculation weak, Genus Echinauris Muir-Wood and Cooper, 1960 restricted to median region between ¯anks; median fold Echinauris sp. notably shallow, poorly delimited on visceral disc, very Fig. 7; 18 broad on anterior trail. The umbonal region is missing, so true length and height are uncertain. Ornament distinctively Remarks. One internal mould of a dorsal disc was available. reticulate; radial costae strong, coarse, rounded, of even It is medium-sized (18 mm wide and 15.5 mm long). Its pan- width, converging slightly in the sulcus, numbering about like appearance with a low and short median septum indi- 16 at mid-length, increasing occasionally by bifurcation; cates Echinauris. As common in the genus, brachial ridges intertroughs of similar width, rounded; concentric rugae are not de®ned. The marginal ridge is moderate. The exter- strong, but ®ner than costae; external spines not de®ned. nal features are not unknown, but rugose radiations occur Remarks. The present dorsal specimen is damaged but is internally, probably corresponding to the external spines. suf®ciently well preserved to retain the main characteristics Considering the poorly developed internal structure, it of Spyridiophora gubleri. The Bera form is almost identical may be an immature shell, although it is not small in size. to all the Cambodian shells ®gured by Mansuy (1913, pl. 13, A Cambodian shell ®gured by Termier and Termier ®g. 1), Chi-Thuan (1961, pl. 5, ®gs. 8 and 9) and Termier (1970b, p. 450, ®g. 6) as E. opuntia (Waagen) has a longer and Termier (1970a, pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G± median septum than the Bera form. Productus khmerianus I). In addition, Colani (1919, p. 10, pl. 1, ®g. 2) re-illustrated Mansuy (1914, p. 17, pl. 6, ®gs. 3a±d) from Phnom Roang, the Mansuy's (1913) Spyridiophora specimen named Sisophon, possibly belongs to Echinauris, as suggested by Productus gratiosus for comparison with her Vietnamese Waterhouse and Piyasin (1970, p. 130) and Grant (1976, p. gratiosus; the latter is probably a species of Transennatia. 123). Prior to them, Ishii et al. (1969, p. 48) reported the 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, UKM- F266, 5±8, dorsal external mould with accompanying natural cast of shell remaining anteriorly Ð posterior, ventral, anterior, and lateral views, £ 3.0, UKM- F267, 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. 190 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

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. M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 191 of the brachial adductor scars, called a `spyridium' (term breviseptum is short. The brachial ridges are located very proposed by Cooper and Stehli, 1955, p. 472). It is well distally. Two small dorsal endospines are discerned. developed in the type species, S. distincta Cooper and Stehli Generic assignment to Kozlowskia is con®rmed by the from West Texas. By comparison, the spyridium of S. distinctive pattern of the ventral strut spines. Three large gubleri, as illustrated by Termier and Termier (1970a, pl. stout spines are seen on the ventral valve of the semi-mature 5, ®g. 6), is less salient. Grant (1976, p. 136) regarded the specimen UKM-F275 Ð one on each lateral slope (Fig. 8; Chi-Thuan's (1961) materials as Transennatia instead of 2±4) and one on the anterior of the disc (Fig. 8; 5). This is Spyridiophora, but made no mention of the Termier and further supported by the appearance of the dorsal interior, Termier (1970a) designation. Waterhouse (1978, p. 119) speci®cally the strong ear baf¯es, large pair of adductors, compared the Mansuy's (1913) material to his Nepalese high breviseptum and dorsal marginal ridge forming a wide Transennatia sp., but does not seem to have accepted the platform. The indistinct or absent sulcus and very weak Termiers' work. However, considering the fact that the ventral ornament also characterise the genus. Termiers' material clearly displays the typical spyridium, Two species of Kozlowskia, K. cornuta Grant (1976, p. the present form is placed in Spyridiophora without 118, pl. 26, ®gs. 24±29; pl. 27, ®gs. 35±38) and K. opipara hesitation. Grant (1976, p. 121, pl. 28, ®gs. 1±27), were found in the Productus craticulatus Reed (1931, p. 11, pl. 1, ®gs., 1 Rat Buri Limestone of southern Thailand. Their dorsal inter- and 2) from the Neoschwagerina craticulifera bed of the iors differ from the Bera form. Grant (1976, p. 118) noted Bamian Limestone, Afghanistan, obviously belongs to that species of Kozlowskia were restricted to the Pennsylva- Spyridiophora, judged from the description and illustra- nian and Early Permian (Wolfcampian); his two Thai tions. It appears to be close to S. gubleri, especially as species were the only Asian occurrences known at the regards its ornament. The South-East Asian forms are time of his study. Since then, only two additional Asian more transverse than the Afghan form, but this may be no occurrences have been reported, both from Early Permian more than within the same species. Because of its distinctive horizons in Xinjiang, northwestern China (Wang and Yang, reticulation and transverse outline, Productus margaritatus 1993; Wang, 1995), both being reports of the type species, Mansuy (1913, p. 28, pl. 2, ®g. 6) from Kham-keut, Laos, is K. capacii d'Orbigny. possibly a species of Spyridiophora. The same species was Waterhouse (1981, p. 13) indicated that Marginifera illustrated by Huang (1932, p. 30, pl. 1, ®gs. 22±24) from sensu Diener (1897, pl. 4, ®gs. 11±13; pl. 5, ®gs. 1 and 2) the Late Permian of Guizhou, South China. This species has from the Chitichun Limestone (Kalabaghian) of Tibet may ®ner reticulation than S. gubleri. Grant (1976, p. 136) noted be Kozlowskia. However, as con®rmed by Waterhouse that this species is unlike Transennatia. Liang (1990) (written communication, 24 July 1999), this was in error proposed two species of Spyridiophora, namely S. because he previously discussed the Diener's specimens in huadongensis (p. 159, pl. 24, ®gs. 1±21) and S. plegma terms of Marginifera, as a species separate from the type (p. 161, pl. 24, ®gs. 22 and 23), both from the Lengwu species, M. typica (see Waterhouse and Piyasin, 1970, p. Formation. They both differ from S. gubleri in having 127; Waterhouse, 1978, p. 30). Marginifera and Kozlowskia much ®ner ornament. often resemble each other considerably, as pointed out by Occurrence. S. gubleri has previously been found only in Cooper and Grant (1975, p. 969). The two genera, however, the Sisophon Limestone at Phnom Tup in the Sisophon can be differentiated by the very different spine patterns on region (Mansuy, 1913; Chi-Thuan, 1961), and at Phnoms the ventral valve. They are now classi®ed into the different Anseh, Totung and Takream in the Battambang region families by Brunton et al. (1995). (Termier and Termier, 1970a). Nakamura (1979a) also recorded this species in Member C of the same formation. Superfamily Linoproductoidea Stehli, 1954 Species of Spyridiophora are extremely rare in the Tethyan Family Linoproductidae Stehli, 1954 Province. They have been recognised only in the Early to Subfamily Linoproductinae Stehli, 1954 Middle Permian of South China, Indochina, Afghanistan Genus Linoproductus Chao, 1927 and the Carnic Alps. Linoproductus sp. Fig. 8; 12±14 Tribe Kozlowskiini Brunton et al., 1995 Genus Kozlowskia Fredericks, 1933 Remarks. Three impressions of the ventral interior display Kozlowskia sp. characteristic costellae of Linoproductus. The shells are Fig. 8; 1±11 medium to large for the genus, ranging 29±40 mm wide and 30±40 mm long. They are semi-ovate to semi-elongate Remarks. This species is represented by two specimens of in outline, and lack a median fold. Costellae are irregular mature and semi-mature shells. The shells are slightly large and uneven, numbering 7±8 in 5 mm medianly. Rugae are for the genus, 20 mm wide, 15 mm long and the disc is 8 mm present only on the lateral trail as broad wrinkles. Parallel high for the mature specimen UKM-F276. The body cavity transverse lines intersect costellae in the largest specimen is of moderate size. The cardinal process is sessile, and the UKM-F278 (Fig. 8; 13). 192 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 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

M. Sone's research was carried out for his B.Sc. Honours Fig. 9. Comparison of microornament among three Phricodothyris species degree at Deakin University in 1998±99, and was ®nanced (all illustrations £ 25), 1, Phricodothyris sp. in this paper, 2, P. pyriformis by his parents, Masao and Toshiko Sone. He is indebted to Pavlova (1969, text-®g. 58), 3, P. rotiformis Liang (1990, pl. 63, ®g. 23). 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 Class Rhynchonellata Williams et al., 1996 manuscript. Dr. Ahmad Jantan and Prof. Ibrahim Komoo are Order Rhynchonellida KuÈhn, 1949 acknowledged for providing ®nancial supports from their Rhynchonellid family indet. government IPRA Projects 02±02±02±0004 and 02±02± Fig. 8; 15 02±0015, respectively. This paper was completed while M. Sone was supported by a fellowship of LESTARI. Remarks. A poorly preserved impression of a ventral interior is the only rhynchonellid in the Bera fauna. It is small References (approximately 20 mm wide and 16 mm long), and has simple, weak costae Ð four in the sulcus and ®ve on each Archbold, N.W., 1984. Western Australian occurrences of the Permian ¯ank. brachiopod genus Retimarginifera. Alcheringa 8 (1/2), 113±122. Archbold, N.W., Bird, P.R., 1989. Permian Brachiopoda from near Kasliu Order Spiriferida Waagen, 1884 village, West Timor. Alcheringa 13 (1/2), 103±123. Suborder Delthyridina Ivanova, 1972 Broili, F., 1916. Die permischen Brachiopoden von Timor. In: Wanner, J. Superfamily Reticularioidea Waagen, 1884 (Ed.), PalaÈontologie von Timor 7, Stuttgart, pp. 1±104. Family Elythidae Fredericks, 1924 Brunton, C.H.C., Lazarev, S.S., Grant, R.E., 1995. A review and new classi®cation of the brachiopod order Productida. Palaeontology 38 Subfamily Phricodothyridinae Caster, 1939 (4), 915±936. Genus Phricodothyris George, 1932 Brunton, C.H.C., Lazarev, S.S., 1997. Evolution and classi®cation of the Phricodothyris sp. Productellidae (Productida), Upper Paleozoic brachiopods. Journal of Fig. 8; 16±17; 9; 1 Paleontology 71 (3), 381±394. Carter, J.L., Johnson, J.G., Gourvennec, R., Hou, Hong-fei, 1994. A revised Remarks. One incomplete ventral specimen was available. classi®cation of the spiriferid brachiopods. Carnegie Museum, Annals 63 (4), 327±374. The shell is sub-orbicular, medium-sized (30 mm wide and Caster, K.E., 1939. A Devonian fauna from Columbia. Bulletin of Amer- about 22 mm long), with double-barrelled (biramous) spine- ican Paleontology 24 (83), 1±218, pls. 1±14. bases, suggesting location in Phricodothyris. It is broadly Chao, Y.T., 1927. Productidae of China, Part 1: Producti. Palaeontolgia convex without sulcation, with its umbo moderately high Sinica, Series B 5 (2), 1±244, 16 pls. and obtuse at about 908. The spine arrangement of the Chi-Thuan, T.T., 1961. Les brachiopodes permiens du Phnom-Tup (Siso- phon Ð Cambodge). Annales de la FaculteÂ des Sciences, UniversiteÂ de Bera shell is similar to that of P. pyriformis Pavlova Saigon 1961, 267±308, 8 pls. (in French). (1969, p. 93, pl. 8, ®gs. 1±3) (Fig. 9; 2), but differs slightly Ching, Y., 1963. Urushtenia from the Lower Permian of China. Acta in that the Bera species has a third row of spine bases (Fig. 9; Palaeontologica Sinica 11 (1), 1±31, 2 pls. (in Chinese with English 1) and slightly coarser concentric laminae. P. rotiformis abstract). Liang (1990, p. 290, pl. 63, ®gs. 17±23) from the lower Colani, M., 1919. Sur quelques fossiles Ouralo-Permiens de Hongay. Service GeÂologique de l'Indochine, Bulletin 6 (5), 1±27, 2 pls. (in Lengwu Formation, South China, is also close; it seems to French). have a similar spine arrangement of three rows (Fig. 9; 3). Cook, R.H., Suntharalingam, T., The geology and mineral resources of Only a mould of the micro-ornament was illustrated for the Pahang Tenggara (Sectors ªEº and ªFº). Geological Survey Malaysia Chinese species; the exact shape of its spines is, therefore, Annual Report 1970, pp. 104±116. uncertain. 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 Order Spiriferinida Ivanova, 1972 (Part 1 and 2). Smithsonian Contributions to Paleobiology 19, 794± Suborder Spiriferinidina Ivanova, 1972 1920, pls. 192±502. M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 193

Cooper, G.A., Stehli, F.G., 1955. New genera of Permian brachiopods from Ishii, K., Kato, M., Nakamura, K., 1969. Permian limestones of West West Texas. Journal of Paleontology 29 (3), 469±474, pls. 52±54. Cambodia Ð lithofacies and biofacies (contribution to the geology Diener, C., 1897. Himalayan fossils. The Permocarboniferous fauna of and palaeontology of Cambodia, Part 3). Palaeontological Society of Chitichun No.1. Palaeontologia Indica, Series 15 1 (13), 1±105. Japan, Special Papers 14, pp. 41±55, pls. 9±11. Fang, R., 1973. The Early Permian Brachiopoda from Xiaoxinzhai of Ivanova, E.A., 1972. Osnovnyye zakonomernosti evolyutsii spiriferid Gengma, Yunnan, and its geological signi®cance. Contribution to the (Brachiopoda) [main features of spiriferid evolution (Brachiopoda)]. Geology of the Qinghai-Xizang (Tibet) Plateau 11, 110±119, 7 pls. (in Paleontologicheskii Zhurnal 1972 (3), 28±42 (in Russian). Chinese with English abstract). Jasin, B., Said, U., Rahman, R.A., 1995. Late Middle Permian radiolarian Fang, R., Jiang, N., 1992. Discovery and its geological signi®cance of from the Jengka area, central Pahang, Malaysia. Journal of Southeast Gubleria from Permian Emeishan Basalt Formation in Ninglang Asian Earth Sciences 12 (1/2), 79±83. Xipiaoluo of northwestern Yunnan. Yunnan Geology 11 (4), 325± Jing (Jin). Y., Hu, S.Z., 1978. Brachiopods of the Kuhfeng Formation in 329, 1 pl. (in Chinese with English abstract). south Anhui and Nanking Hills. Acta Palaeontologica Sinica 17 (2), Fantini Sestini, N., 1965. Bryozoans, brachiopods and molluscs from Ruteh 101±127, 4 pls. (in Chinese with English abstract). Limestone (Permian). The geology of the Upper Djadjerud and Lar Jin, Y., Wardlaw, B.R., Glenister, B.F., Kotlyar, G.V., 1997. Permian Valleys (North Iran) II, Palaeontology. Rivista Italiana di Paleontologia chronostratigraphic subdivisions. Episodes 20 (1), 10±15. e Stratigra®a 71 (1), 13±108, pls. 2±8. Jones, C.R., Gobbett, D.J., Kobayashi, T., 1966. Summary of fossil record Fischer de Waldheim, G., 1829. Quelques fossiles du Gouvernement de in Malaya and Singapore 1900±1965. Geology and Palaeontology of Moscou. SocieÂteÂ ImpeÂriale des Naturalistes de Moscou Bulletin 1 (12), Southeast Asia 2, 309±359. 375±376. Kobayashi, F., 1997. Middle Permian biogeography based on fusulinacean Fontaine, H., Ibrahim, A., Khoo, H.P., Nguyen, D.T., Vachard, D., 1994. faunas. In: Ross, C.A., Ross, J.R.P., Brenckle, P.L. (Eds.), Late Paleo- Geological Papers Volume 4. Geological Survey of Malaysia, Ministry zoic Foraminifera; their Biostratigraphy, Evolution and Paleoecology, of Primary Industries, pp. 1±175. and the Mid-Carboniferous Boundary. Cushman Foundation for Fora- Fredericks, G.N., 1924. Ussuriiskii verkhnii paleozoi. I. Brachiopoda miniferal Research Special Publication vol. 36, pp. 73±76. (Upper Paleozoic of the Ussuriland, I: Brachiopoda.). Materialy po KuÈhn, O., 1949. Lehrbuch der PalaÈozoologie. Schweizerbart'sche Verlags- Geologii i Poleznym Iskopaemym Dal'nego Vostoka 28, 1±53, 1 pl. buchhandlung, Stuttgart, 326 pp. (in Russian). Leman, M.S., 1993. Upper Permian brachiopods from northwest Pahang, Fredericks, G.N., 1933. Paleontologicheskie etiudy, 4: O nekotorykh Malaysia. In: Thanasuthipitak, T. (Ed.), Proceedings of the Interna- verkhne-paleozoiskikh brakhiopodakh Evrazii (palaeontological tional Symposium on Biostratigraphy of Mainland Southeast Asia: notes, 4: on some Upper Paleozoic brachiopods of Eurasia). Materialy Facies & Paleontology. Chang Mai, Thailand, pp. 203±218. Tsentral'nogo Nauchno-Issledovatel'skogo Geologo-Razvedochnogo Leman, M.S., 1994. The signi®cance of Upper Permian brachiopods from Instituta (TSNIGRI), Paleontolgiia i Stratigra®ia Sbornik 2, 24±33 (in Merapoh area, northwest Pahang. Geological Society of Malaysia Russian). Bulletin 35, 113±121. Geological Survey of Malaysia, 1985. Geological Map of Peninsular Leman, M.S., Mohamed, K.R., Sone, M. On the new Permian Bera Forma- Malaysia, 8th ed. (1:500,000). Directorate of Geological Survey Malay- tion from the Bera District, Pahang, Malaysia, Proceedings of the sia, Kuala Lumpur. Annual Geological Conference 2000. 8±9 September 2000, Penang. George, T.N., 1932. The British Carboniferous reticulate Spiriferidae. Geological Society of Malaysia. In press Geological Society of London Quarterly Journal 88, 516±575, pls. Leven, E.Ya.(Ja.), 1992. Problems of Tethyan Permian stratigraphy. Inter- 31±35. national Geology Review 34 (10), 976±985. Gobbett, D.J., 1973. Chapter 4, Upper Paleozoic. In: Gobbett, D.J., Hutch- Leven, E.Ya.(Ja.), 1993. Phylogeny of the sumatrinids and questions on the ison, C.S. (Eds.), Geology of the Malay Peninsula (West Malaysia and zonal subdivision of the Midian Stages of the Permian System. Paleon- Singapore). Wiley/Interscience, New York, pp. 61±96. tological Journal 27 (3), 29±35. Grant, R.E., 1976. Permian brachiopods from southern Thailand. Journal of Leven, E.Ja., Grant-Mackie, J.A., 1997. Permian fusulinid foraminifera Paleontology (Paleontological Society Memoir 9) 50 (3), 1±269. from Wherowhero Point, Orua bay, Northland, New Zealand. New Gray, J.E., 1840. Synopsis of the contents of the British Museum, 42nd ed. Zealand Journal of Geology and Geophysics 40, 473±486. London, 308 pp. Liang, W., 1990. Lengwu Formation of Permian and its brachiopod fauna in Hamlet, B., 1928. Permische Brachiopoden, Lamellibranchiaten und Zhejiang Province. People's Republic of China, Ministry of Geology Gastropoden von Timor. Jaarboek van het Mijnwezen in Neder- and Mineral Resources, Geological Memoirs, Series 2. 10. Geological landsch-IndieÈ 56 (2), 1±115, 12 pls. Publishing House, Beijing, pp. 1±522, 84 pls. (in Chinese with English He, X., Shi, G.R., 1996. The sequence of Permian brachiopod assemblages summary). in South China. In: Cooper, P., Jin, J. (Eds.), Proceedings of the Third Likharev, B., 1935. Bemerkungen uÈber einige oberpalaÈozoische Brachio- International Brachiopod Congress, 2±5 September 1995, Subdury, poden. Zentralblatt fuÈer Mineralogie, Geologie und PaÈlaeontologie, Ont., Canada, pp. 111±115. Abt. B 9, 369±373. Hu, S., 1983. Brachiopods from the Hsiaochiangpien Limestone, southern MacDonald, S., 1970. Geology and mineral resources of the Lake Chini Ð Jiangxi. Acta Palaeontologica Sinica 22 (3), 338±345, pls. 1±3 (in Sungei Bera Ð Sungei Jeram area of south-central Pahang. Geological Chinese with English abstract). Survey of Malaysia, Map Bulletin 1, pp. 1±44. Huang, T.K., 1932. Late Permian Brachiopoda of southwestern China. Mansuy, H., 1912. Mission du Laos 1, GeÂologie des environs de Luang- Palaeontologia Sinica, Series B 9 (1), 1±113, 9 pls. Prabang. Service GeÂologique de l'Indochine MeÂmoires 1 (4), 1±32, pls. Igo, H., 1964. Permian fossils from northern Pahang, Malaya. Geology and 1±7 (in French). Paleontology of Southeast Asia 1, 191±207, pl. 2. Mansuy, H., 1913. Faunes des Calcaires aÁ Productus de l'Indochine, Igo, H., 1967. Some Permian fusulinids from Pahang, Malaya. Geology and PremieÁre SeÂrie. Service GeÂologique de l'Indochine MeÂmoires 2 (4), Palaeontology of Southeast Asia 3, 30±38, pls. 4±9. 1±133, 13 pls. (in French). Igo, H., Rajah, S.S., Kobayashi, T., 1979. Permian fusulinacean from the Mansuy, H., 1914. Faunes des Calcaires aÁ Productus de l'Indochine, Deux- Sungai Sedili area, Johore, Malaysia. Geology and Palaeontology of ieÁme SeÂrie. Service GeÂologique de l'Indochine MeÂmoires 3 (3), 1±59, Southeast Asia 20, 95±118, pls. 15±26 pls. 1±7 (in French). Ishii, K., 1966. On some fusulinids and other Foraminifera from the Metcalfe, I., 1996a. Pre-cretaceous evolution of SE Asian terranes. In: Hall, Permian of Pahang, Malaya. Journal of Geosciences, Osaka City R., Blundell, D.J. (Eds.), Tectonic Evolution of Southeast Asia, Geolo- University 9 (4), 131±136, pls. 5±6. gical Society Special Publication 106, pp. 97±122. 194 M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194

Metcalfe, I., 1996b. Gondwanaland dispersion, Asian accretion and evolution Tazawa, J., 1987. Kamiyase-chiku Kaseki Chyosa Houkoku-syo (the report of Eastern Tethys. Australian Journal of Earth Sciences 43, 605±623. of the fossil survey in the Kamiyase area). Publication of the Kesennuma Metcalfe, I., 1998. Palaeozoic and Mesozoic geological evolution of the SE Educational Committee, Miyagi Prefecture, pp. 1±37 (in Japanese). Asian region: multidisciplinary constraints and implications for biogeo- Tazawa, J., Matsumoto, T., 1998. Middle Permian brachiopods from the graphy. In: Hall, R., Holloway, J.D. (Eds.), Biogeography and Geolo- Oguradani Formation, Ise district, Hida Gaien Belt, Central Japan. gical Evolution of SE Asia. Backbuys, Leiden, pp. 25±41. Science Reports of Niigata University, Series E (Geology), 13, 1±19 Muir-Wood, H.M., 1948. Malayan Lower Carboniferous Fossils and their Termier, H., Termier, G., 1960. Contribution aÁ la classi®cation des Brachio- Bearing on the Visean Palaeogeography of Asia. British Museum podes: le lophophore des Collolophides nov. ord. Appendice: les Oldha- (Natural History), London, pp. 1±118. minideÂs du Cambodge. SocieÂteÂ GeÂologique de France Bulletin, Series, 7 Muir-Wood, H.M., Cooper, G.A., 1960. Morphology, classi®cation and life 1 (3), 233±244, 2 pls. (in French). habits of the Productoidea (Brachiopoda). Geological Society of Amer- Termier, H., Termier, G., 1970a. Le genre Spyridiophora (Brachiopodes ican Memoir 81, 1±447, 135 pls. ProductoõÈdeÂs) dans le Permien asiatique. SocieÂteÂ GeÂologique du Nord, Nakamura, K., 1979a. Kenkyu Seika Houkokusho, ªHimalaya Zenentai Annales 90 (2), 57±62 1 pl. (in French). oyobi Honpousan Perumuki Wanzokurui no Hikaku Kenkyuº (the Termier, H., Termier, G., 1970b. Les ProductoõÈdeÂs du Djoul®en (Permien report of research results, ªcomparison between Permian brachiopods supeÂrieur) dans la TeÂthys orientale: essai sur l'agonie d'un phylum. from the Himalaya Frontal Margin and Japanº). Monbu-syo Science SocieÂteÂ GeÂologique du Nord, Annales 90 (4), 443±461, pls. 29±31 Research Subsidy (General Science Category C) for the Syowa Year 53 (in French with English abstract). (in Japanese), unpublished. Tong, Z., 1978. Brachiopoda, Carboniferous±Permian. Palaeontological Nakamura, K., 1979b. Additional occurrences of Urushtenoidea (brachio- Atlas of Southwestern China, Sichuan Province, 2. Geological Publish- pods) from the Permian of Asia. Journal of Faculty of Science, ing House, Beijing, pp. 10±267, pls. 77±92 (in Chinese). Hokkaido University, Series 4. Geology and Mineralogy 19 (1/2), Waagen, W., 1883. Salt Range Fossils: Productus Limestone fossils, Part 4 221±233, 3 pls. (2), Brachiopoda Palaeontologia Indica, Series 13 (1), 391±546. Nakamura, K., Shimizu, D., Liao, Z., 1985. Permian palaeobiogeography of Waagen, W., 1884. Salt Range Fossils: Productus Limestone fossils, Part brachiopods based on the faunal provinces. In: Nakazawa, K., Dickins, 4(2) Brachiopoda. Palaeontologia Indica, Series 13 (1), 547±728. J.M. (Eds.), The Tethys. Tokai University Press, Tokyo, pp. 185±198. Wang, C., 1995. Brachiopod fauna from the Kangkelin Formation in Nakazawa, K., 1973. On the Permian fossils from Jengka Pass, Pahang, Akesu, the Xinjiang Autonomous Region. Journal of Changchun Malay Peninsula. Tohoku University, Science Report Series 2 (Geol- University of Earth Sciences 25 (1), 15±23, 4 pls. (in Chinese with ogy), Special Volume 6 (Hatai Memorial Volume), 277±296, pls. 30± English abstract). 31 Wang C., Yang, S., 1993. Brachiopod fauna around the Carboniferous± Noetling, F., 1905. Untersuchungen uÈber die Familie Lyttoniidae Waag. Permian boundary from the Balikelike Formation in Keping, Xinjiang. emend. Noetling. Palaeontographica 51, 129±153, pls. 15±18. Journal of Changchun University of Earth Sciences 23 (1), 1±9, 3 pls. OÈ pik, A.A. 1934. UÈ ber Klitamboniten. Acta Commentationes, Series A, 26 (in Chinese with English abstract). (3), 1±239, 48 pls. Tartu University, Dorpat. Waterhouse, J.B., 1975. New Permian and Triassic brachiopod taxa. Papers Pavlova, E.E., 1969. Razvitie brakhiopod semeistva Reticulariidae (devel- of Department of Geology, University of Queensland 7 (1), 1±23, pls. opment of brachiopods of the family Reticulariidae). Akademia Nauk 1±2. SSSR, Palaeontologicheskii Institut, Trudy 120, pp. 1±128 (in Russian). Waterhouse, J.B., 1976. Permian brachiopod correlations for South-east Reed, F.R.C., 1931. Upper Carboniferous fossils from Afghanistan. Asia. Geological Society of Malaysia Bulletin 6, 187±210. Palaeontologia Indica 19, 1±39, 4 pls. Waterhouse, J.B., 1978. Permian Brachiopoda and Mollusca from north- Rudwick, M.J.S., Cowen, R., 1967. The functional morphology of some west Nepal. Palaeontographica, Abt. A 160 (1±6), 1±175, 26 pls. aberrant strophomenide brachiopods from the Permian of Sicily. Soci- Waterhouse, J.B., 1981. Early Permian brachiopods from Ko Yao Noi and eta paleontologica Italiana, Bollettino 6 (2), 113±176, pls. 32±43. near Krabi, southern Thailand. In: Waterhouse, J.B., Pitakpaivan, K., Sarytcheva, T.G., Sokol'skaya, A.N., 1959. O klassi®katsin lozhnoporis- Mantajit, N. (Eds.), The Permian Stratigraphy and Palaeontology of tykh brachiopod (on the classi®cation of pseudopunctate brachiopods). Southern Thailand. Geological Survey of Thailand (Memoir vol. Vol. 125, pp. 181±184. Akademiia Nauk SSSR, Doklady. 4(2)), pp. 43±213. Schellwien, E., 1900. BeitraÈge zur Systematik der Strophomeniden des Waterhouse J.B., Piyasin, S., 1970. Mid-Permian brachiopods from Khao oberen Palaeozoicum. Neues Jahrbuch fuÈer Mineralogie, Geologie Phrik, Thailand. Palaeontographica, Abt. A 135 (3±6), 83±197, pls. und PalaÈeontologie 190, 1±15, 1 pl. 14±32 Schuchert, C., 1913. Class Brachiopoda. The Lower Devonian deposits of Williams, A., Brunton, C.H.C., 1993. Role of shell structure in the classi- Maryland. Maryland Geological Survey, Baltimore. 290±449. ®cation of the Orthotetidine brachiopods. Palaeontology 36 (4), 931± Schuchert, C., 1929. Brachiopoda (generum et genotyporum index et 966. bibliographia). In: Schuchert, C. and LeVene, C.M. (Eds.), Fossilium Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L.E., Popov, L., Catalogus. I. Animalia. W. Junk, Berlin. 140 pp. 1996. A supra-ordinal classi®cation of the Brachiopoda Philosophical Shen, S., Shi, G.R., 1996. Diversity and extinction patterns of Permian Translations of the Royal Society of London. Biological Science 351, Brachiopoda of South China. Historical Biology 12, 93±110. 1171±1193. Sokol'skaya, A.N., 1965. Brachiopoda: Strophomenida. In: Ruzhensev, Yanagida, J., 1964. Permian brachiopods from central Thailand. Faculty of V.E., Sarytcheva, T.G. (Eds.), Razvitie i smena morskikh organismov Sciences, Kyushu University Memoirs, Series D. Geology 15 (1), 1±22, na rubezhe paleozoia i mezozoia (the development and change of pls. 1±3 marine organisms at the Palaeozoic±Mesozoic boundary). Akademia Yanagida, J. Aw, P.C., 1979. Upper Carboniferous, Upper Permian and Nauk SSSR, Palaeontologicheskii Institut, Trudy 108, pp. 203±209 (in Triassic brachiopods from Kelantan, Malaysia. Geology and Palaeon- Russian). tology of Southeast Asia 20, 119±141, pls. 27±29 Stehli, F.G., 1954. Lower Leonardian Brachiopoda of the Sierra Zhang, Y., Fu, L., Ding, P., 1983. Phylum Brachiopoda. In: Palaeontolo- Diablo. Bulletin of the American Museum of Natural History gical Atlas of Northwest China (Shaanxi±Gansu±Ningxia Provinces), 105, 257±358. Upper Palaeozoic 2. Geological Publishing House, pp. 244±425, pls. Tazawa, J., 1976. The Permian of Kesennuma, Kitakami Mountains: a 88±143 (in Chinese). preliminary report. Earth Science (Chikyu Kagaku). Journal of the Zeng, Q., 1987. Some structures and life habits of Urushtenoidea. Acta Association for the Geological Collaboration in Japan 30 (3), 175± Palaeontologica Sinica 26 (4), 497±503, 2 pls. (in Chinese with English 185, 3 pls. abstract).