J. Paleont., 75(2), 2001, pp. 274-283 Copyright ? 2001, The Paleontological Society 0022-3360/01/0075-274$03.00

A LOPINGIAN (LATE PERMIAN) BRACHIOPOD FAUNA FROM THE QUBUERGA FORMATION AT SHENGMI IN THE MOUNT QOMOLANGMA

REGION OF SOUTHERN XIZANG (TIBET), CHINA SHUZHONG SHEN,' 2 N. W. ARCHBOLD,2 AND G. R. SH12

'Nanjing Institute of Geology & Palaeontology, Chinese Academy of Science, 39 eastern Beijing Road, Nanjing, Jiangsu 210008, P. R. China, and 2School of Ecology and Environment, Deakin University, Rusden Campus, 662 Blackburn Road, Clayton, Victoria 3168, Australia

<zshen@deakin.edu.au>, <narchie@deakin.edu.au>, <grshi @deakin.edu.au>

ABSTRACT-A small fauna of 11 species belonging to 10 genera of Permian Brachiopoda from the lower part of the Qubuerga Formation outcropping near Shengmi village in the Qomolangma region of southern Xizang (Tibet) is figured and new taxa are described. New taxa are Quinquenella semiglobosa and Costatumulus shengmiensis. The fauna is most likely of Wuchiapingian (Djhulfian) age as indicated by the majority of the brachiopod species.

INTRODUCTION

THE INACCESSIBILITY of the area near Mt. Qomolangma (Mt. Everest) of Xizang (Tibet) has resulted in relatively

little geological work being undertaken in the area prior to this study. Muir-Wood and Oakley (1941) first reported a small col- lection of poorly preserved fossils from the Mt. Qomolangma region, and confirmed the existence of the Permian in this region. In 1952, Li Pu completed a field survey over most areas of eastern Xizang and collected some Permian brachiopods near Mt. Qom- olangma that were later described by Ding (1962) as Neospirifer kubeiensis Ding, N. tibetensis Ding, N. moosakhailensis (David- son), Costiferina indica (Waagen), and Spiriferella salteri Tscher- nyschew.

In 1964, a scientific investigation team of the Chinese Academy of Sciences completed a field expedition in the Qomolangma re- gion. Brachiopods collected by this team were described by Zhang and Jin (1976) and subsequently summarized by Jin (1985).

The brachiopod fauna described in the present paper was col- lected by the senior author in 1997 during a field expedition to the Mt. Qomolangma region about 430 km west from Lhasa, the capital of Xizang (Tibet). Tectonically, this region belongs to the Himalayan Terrane. The studied section is located on the north- western slope of Mt. Jinong (5469.9 m above sea level), about 40 km north of Mt. Qomolangma. A small village (87?09'E, 28?34'N) called Shengmi is at the foot of Mt. Jinong (Fig. 1). Brachiopods have never been described before from this locality, but four brachiopod species, identified as Spiriferella rajah (Salt- er), Marginifera himalayensis (Diener), Martinia sp., and Can- crinella sp., associated with the late Changhsingian conodonts Clarkina changxingensis Wang and Wang, C. deflecta Wang and Wang and C. subcarinata (Sweet) in the Permian-Triassic bound- ary beds, were mentioned by Yu and Dai (1985) and Rao and Zhang (1985).

STRATIGRAPHY

The Permian section in the Himalayan region of Xizang (Tibet) is commonly divided into two parts. The lower part of the section in the study area, named the Jilong Formation, consists of diam- ictites, sandstone, and siltstone (Yin and Guo, 1976) with "Ste- panoviella" brachiopod fauna (Jin, 1979) of probable Sakmarian age. The upper part of the section overlying the Jilong Formation by fault contact has been subdivided into the Qubu and Qubuerga Formations in ascending order (Yin and Guo, 1976). The Qubu Formation is mainly composed of quartzose sandstone interbed- ded with black shale and silty shale, and contains a Glossopteris flora (Hsii, 1976) at the Qubu section about 30 km north of Mt.

Qomolangma. This flora was regarded as early Late Permian (Wuchiapingian in the sense of Jin et al., 1997) in age, correlative with the Raniganj Beds in India (Hsii, 1976).

The Qubuerga Formation, lying above the Qubu Formation without obvious discordance in the Mt. Qomolangma region, is a sequence of siltstone interbedded with shale and bioclastic lime- stone containing abundant brachiopods, bryozoans, and corals. According to Yin and Guo (1976) and Yin (1997), three members have been recognized. The lower member is 118.3 m thick, and composed of brown sandstone, muddy siltstone, and shale. The middle member (120.5 m thick) is dominated by gray silty shale grading upwards to bioclastic siltstone interbedded with bioclastic limestone. The upper member, conformably overlain by the do- lomitic limestone of the Lower Triassic Tulung Formation, is 86.3 m thick, and consists of broadly monotonous dark gray shale with bivalves, plant stems, and unidentifiable ammonoids.

Rao and Zhang (1985) named the Permian strata at Shengmi as the Baga Formation (the topmost 7 m of the Selong Formation) and the Menduo Formation (the remaining part of the Selong Formation) in descending order. These two formations are not significantly different in lithology from the previously-defined Qubuerga Formation, hence this nomenclature is not used herein.

The brachiopods described in this paper come from the lower member of the Qubuerga Formation outcropping about 1 km northeast of Shengmi village (Fig. 2). The strata are deformed. Most of the brachiopods are represented by isolated valves, and are poorly preserved with worn surfaces indicating that the shells were abraded during transportation prior to deposition.

DISCUSSION OF AGE

The Qubuerga Formation in the Qomolangma region is most likely of Wuchiapingian (Djhulfian) to Changhsingian in age. An apparently Wuchiapingian flora in the immediately underlying Qubu Formation (Hsu, 1976) and the late Changhsingian cono- dont Clarkina changxingensis in the topmost beds of the Qub- uerga Formation (Yu and Dai, 1985) constrain the age. Eleven brachiopod species, belonging to 10 genera, have been identified from the present collection (Table 1). Although of low diversity the fauna most likely indicates a correlation with the Wuchiapin- gian (Djhulfian) Stage. However, the possibility of an early Changhsingian age cannot be ruled out.

Neochonetes (Sommeriella) strophomenoides (=N. convexa Liao, 1980) is one of the commonest species in the Lopingian of South China and is also found from the Wargal Formation in the Salt Range, Pakistan (Waagen, 1884). Conodonts (Wardlaw and Pogue, 1995; Wardlaw, 1997) and fusulinids (Pakistan-Japanese Working Group, 1985) indicate that the Wargal Formation is largely of Wuchiapingian (Djhulfian) age. Biplatyconcha grandis

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TABLE 1-Brachiopod species from near Shengmi, Xizang (Tibet).

Ventral Dorsal Con- Species valves valves joined

Quinquenella semiglobosa n.sp. 7 8 Neochonetes (S.) strophomenoides (Waagen) 8 1 Taeniothaerus sp. 2 Biplatyconcha grandis Waterhouse 2 2 Waagenoconcha? sp. - 1 Costatumulus shengmiensis n.sp. 3 2 Costatumulus sp. 1 - Pondospirifer cf. magnificus Waterhouse 2 Costatispirifer? sp. 2 - Fusispirifer sp. 2 2 Himathyris sp. - 1

FIGURE --Sketch map showing the fossil locality.

FIGURE 2-Stratigraphic section of the Permian System in the Qomo- langma region of Xizang (Tibet).

and Pondospirifer are known from the Nisal Member of the Senja Formation in northwest Nepal (Waterhouse, 1978). Costatumulus shengmiensis n. sp. is close to C. papilionata (Waterhouse) from the Nambdo Member; and Costatumulus sp. is similar to C. pol- liciformis Waterhouse from the Nisal Member of the Senja For- mation in northwest Nepal. The Nisal and Nambdo Members are within the Marginalosia kalikotei Zone, which was considered to be Dorashamian (=Changhsingian) by Waterhouse (1978, 1988). However, M. kalikotei possibly ranges from the Capitanian to the Changhsingian at the Selong Xishan section in Nyalam County, southern Xizang (Wang et al., 1989; Shen et al., 1998). A low diversity fauna from the middle part of the Selong Formation at the Selong Xishan section was described by Shi and Shen (1997) and considered to be Midian to early Djhulfian (Wuchiapingian) in age. The M. kalikotei Zone contains abundant Spiriferella ra- jah. This species is very common in the Wuchiapingian (Djhul- fian) of the Himalayan region (Angiolini in Garzanti et al., 1996).

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Waagenoconcha? sp. closely resembles W. purdoni (Davidson) from the Chhidru Formation, Salt Range, Pakistan of apparently late Wuchiapingian to Changhsingian age (Wardlaw, 1997; Jin et al., 1997). It is also comparable with W. imperfecta Prendergast (1935) of Djhulfian (Wuchiapingian) age (Archbold, 1998) in Western Australia in terms of its outline and profile (Archbold, 1993). Fusispirifer sp. is close morphologically to F. nitiensis (Diener) from the Productus Shales at Kiunglung near the Niti Pass (Diener, 1897), which are likely to be of Wuchiapingian age, and also from the Nisal Member of the Senja Formation in north- west Nepal (Waterhouse, 1978). This species is also comparable with F. marcouiformis Jin (=Ching) in Zhang and Jin (1976) from the Selong Formation in southern Xizang. Himathyris has been reported from the Kuling Shale of Spiti, India (Diener, 1897) and the Productus Shales of the Lissar Valley (Johar), which were referred to the Lamnimargus himalayensis Zone by Waterhouse (1985), also of Wuchiapingian (Djhulfian) age.

All specimens figured in this paper are registered and housed in the Natural Museum of Victoria, Melbourne, Australia (NMV P).

SYSTEMATIC PALEONTOLOGY

Family RUGOSOCHONETIDAE Muir-Wood, 1962 Subfamily QUINQUENELLINAE Archbold, 1981a

Genus QUINQUENELLA Waterhouse, 1975

Type species.-Quinquenella glabra Waterhouse, 1975, p. 2, pl. 1, figs. 1-3.

Diagnosis.-Quinquenellinid with oblique hinge spines; dorsal valve with variably developed median septum, long thin acces- sory septa and short strong lateral septa; surface smooth except for some concentric lines; deltidial plates and chilidium present.

Discussion.-Quinquenella was originally proposed and based on specimens from the Wuchiapingian (Djhulfian) of Popa, north- west Nepal (Waterhouse, 1975). It is characterized by oblique hinge spines, a variably developed but prominent median septum in the ventral valve, long thin accessory septa and short lateral septa in the dorsal valve, and smooth exteriors (Waterhouse, 1975; Archbold, 1981a). Small shells of Quinquenella have no median septum and possess pustules which are arranged regularly over the valve with no ridges (Archbold, 1981b), but mature dorsal valves have distinct, short lateral septa, long thin accessory septa and a median septum of variable length (Archbold, 1981b). Gla- brichonetina Waterhouse (1978) is considered to be a junior syn- onym of Quinquenella (Archbold, 1981b, p. 29) and is apparently based on juvenile specimens of Quinquenella.

QUINQUENELLA SEMIGLOBOSA new species Figure 3.1-3.9

Diagnosis.--Quinquenella with subquadrate outline, quadrate or slightly obtuse cardinal extremities and semiglobose profile.

Description.-Shell very small, subquadrate or slightly semi- circular in outline, semiglobose in profile; hinge equal to or slight- ly shorter than greatest width; cardinal extremities subquadrate; ears very small, not differentiated from venter; ventral valve

strongly convex in profile; maximum convexity at midvalve; beak inconspicuous; interarea low, broadly triangular; delthyrium small; sulcus absent; surface smooth except for concentric growth lines. Dorsal valve nearly flat or slightly concave; ears flat, no sulcus or fold. Ventral interior with a prominent median septum; median septum variable in length, thick and low at posterior half, extending to anterior half as a low ridge; anterior and lateral flank interior pustulose. Dorsal interior with distinct lateral septa, long accessory septa and median septum beginning from alveolus; al- veolus circular and small; radiating rows of elongate papillae well developed in between septa; median septum inconspicuous, may become a row of elongate papillae anteriorly; socket ridges thick, reducing laterally; cardinal process interiorly bilobed, exteriorly quadrilobed.

Etymology.-Semi- (Latin), half; globose-, spherical, referring to its profile.

Types.-Holotype, an internal mold of a ventral valve (NMV P303084); paratypes, an internal mold of a dorsal valve (NMV P303085), an external mold of a dorsal valve (NMV P303086), and an external mold of a ventral valve (NMV P303087).

Measurements.-See Table 2. Other material examined.-A ventral valve (NMV P303088),

three internal molds of ventral valves (NMV P303089-303091); six internal molds of dorsal valves (NMV P303092-303097); an external mold of ventral valve (NMV P303098).

Discussion.-The new species is similar to Quinquenella ku- waensis (Waterhouse, 1978) from the Kuwa Member of the Senja Formation in northwest Nepal in outline, but the Nepalese species is of very small size and possesses a less strongly convex ventral valve and somewhat flattened ears. The type species, Q. glabra Waterhouse (1975), also from the Senja Formation in northwest Nepal, differs from the present Tibetan species in its less strong convex profile and in having alate extremities with minute, acute ears (Waterhouse, 1978, p. 107). Q. magnifica Archbold (1981b) from the upper Lower Permian (Kungurian?) of Irian Jaya, In- donesia, and Q. australis Archbold (1981a) from the Kungurian of Western Australia are of relatively larger size in maturity, and are less convex, and possess more prominent and flattened ears than the present species.

Subfamily RUGOSOCHONETINAE Muir-Wood, 1962 Genus NEOCHONETES Muir-Wood, 1962

Subgenus NEOCHONETES (SOMMERIELLA) Archbold, 1982 NEOCHONETES (SOMMERIELLA) STROPHOMENOIDES

(Waagen, 1884) Figure 3.10-3.15

Chonetes strophomenoides WAAGEN, 1884, p. 628, pl. 58, fig. lOa-f; FRECH, 1911, p. 164, pl. 23, fig. 5a, b.

Neochonetes convexa LIAO, 1980, p. 257, pl. 5, figs. 18-22; WANG ET AL., 1982, p. 200, pl. 95, figs. 12, 13; Xu, 1987, p. 221, pl. 9, figs. 13, 25, 28; pl. 10, figs. 4-10.

Material examined.-Two external molds of ventral valves (NMV P303099, NMV P303100); Six internal molds of ventral

FIGURE 3-1-9, Quinquenella semiglobosa n. sp.; 1, latex of external mold of ventral valve, NMV P303087, paratype, X6; 2, internal mold of ventral valve, NMV P303089, X6; 3, ventral valve, NMV P303088, X5.8; 4, internal mold of ventral valve, NMV P303090, X6; 5, internal mold of ventral valve, NMV P303084, holotype, x6; 6, latex of internal mold of dorsal valve, NMV P303095, X5.3; 7, latex of external mold of dorsal valve showing the exteriorly quadrilobed cardinal process, NMV P303086, paratype, X5.7; 8, latex of internal mold of dorsal valve showing lateral septa, long accessory septa and short median septum, well developed elongate papillae in between septa, and interiorly bilobed cardinal process, NMV P303085, paratype, X6.4; 9, latex of internal mold of dorsal valve, NMV P303092, x5.8. 10-15, Neochonetes (Sommeriella) strophomenoides (Waagen); 10, latex of external mold of ventral valve, NMV P303099, X3.3; 11, internal mold of ventral valve, NMV P303102, X3.2; 12, internal mold of ventral valve, NMV P303106, X2.7; 13, internal mold of ventral valve, NMV P303101, X3.2; 14, internal mold of dorsal valve, NMV P303107, X3; 15, internal mold of ventral valve, NMV P303104, X2.6.

277

278 JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

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SHEN ET AL.-PERMIAN BRACHIOPOD FAUNA FROM CHINA (TIBET)

valves (NMV P303101-NMV P303106); an internal mold of dor- sal valve (NMV P303107).

Discussion.-The present specimens are close morphologically to N. convexa Liao (1980, p. 257, pl. 5, figs. 18-22) from the Lopingian of South China and Chonetes strophomenoides Waagen (1884, p. 628, pl. 58, fig. lOa-f) from the Wargal Formation in the Salt Range, Pakistan. Frech (1911) initially documented ma- terial from the Lopingian of South China as Chonetes stropho- menoides Waagen. C. strophomenoides was tentatively listed as a species of Waterhousiella by Archbold (1983). Waagenites spe- ciosus Waterhouse and Piyasin (1970) from the Kungurian-Ufi- mian of Thailand was designated as the type species of Water- housiella (Archbold, 1983). The moderately fine costellae (about 40 in total number near front margin of ventral valve) and the costellate ears of C. strophomenoides suggest that it is probably referable to Neochonetes. The shell outline and nature of the sul- cus of the present material indicate that it belongs to the subgenus Sommeriella. Waterhousiella is characterized by small size, coarse and simple costellae, non-costellate ears and relatively con- vex profile. Late Permian species recorded as "Waagenites" and Fusichonetes (Liao in Zhao et al., 1981) from the Lopingian of South China may be referable to Waterhousiella. Unfortunately, the internal details of N. strophomenoides are still unknown. N. convexa Liao (1980) is considered to be a synonym of N. stro- phomenoides as indicated by its similar size, shell outline, con- vexity, costellation and nature of the sulcus.

Neochonetes (Sommeriella) strophomenoides bears some re- semblance to another South Chinese Lopingian species, N. zhon- gyingensis Liao (1980, p. 257, pl. 5, figs. 10-13) in size and costellation; but N. zhongyingensis usually has a less strong de- veloped ventral convexity than N. strophomenoides and acute car- dinal extremities. Rugaria nisalensis (Waterhouse, 1978, p. 60, pl. 7, figs. 1-13) from the Nanguang Formation in northwest Nepal is also somewhat similar to the present species in outline, con- vexity and size, but the former has coarser, stronger and fewer costellae than N. strophomenoides. Sulcirugaria subquadrata Wa- terhouse (1983a) from the Pija Member of the Senja Formation in northwest Nepal is similar to the present species in outline, costellation, nature of the sulcus, short median septum in ventral valve, prominent brachial scars, and a long median septum in the dorsal valve. However, Sulcirugaria, according to the definition of Waterhouse (1983a), does not have lateral septa in the dorsal valve. Neochonetes (S.) hardmani Archbold (1993) (=N. sp. A, Archbold, 1981c, p. 125, fig. 12A-C) from the Wuchiapingian (Djhulfian) Cherrabun Member of the Hardman Formation of the Canning Basin, Western Australia, is also similar to the present species in outline, but is smaller and less convex in profile.

Family STROPHALOSIIDAE Schuchert, 1913 Subfamily STROPHALOSIINAE Schuchert, 1913

Genus BIPLATYCONCHA Waterhouse, 1983a

Platyconcha WATERHOUSE, 1975, p. 8, pl. 2, figs. 1-4; 1978, p. 73, p1. 10, figs. 11-14; pl. 11, figs. 1-5.

Biplatyconcha WATERHOUSE, 1983a, p. 125. Megalosia WATERHOUSE, 1988, p. 45, figs. 4, 5.

Discussion.-Platyconcha was originally proposed as a genus

of the family Waagenoconchidae Muir-Wood and Cooper by Wa- terhouse (1975) based on specimens from the Nisal Member of the Senja Formation in northwest Nepal. The absence of dorsal spines, the close-set dimples over dorsal valve, the relatively deep adductor scars in ventral valve, and the presence of a ventral interarea all differ from those of Waagenoconchidae, but indicate that the genus is a strophalosiid (Briggs, 1998, p. 113). The dorsal interior of Platyconcha was not well figured, but was described as possessing a bilobed cardinal process, low median septum and smooth adductors by Waterhouse (1975, 1978). The replacement of the name Biplatyconcha was proposed by Waterhouse (1983a, p. 125).

Megalosia subsequently proposed by Waterhouse (1988), with- out reference to Biplatyconcha, also based on the specimens from the Nisal Member of the Senja Formation in northwest Nepal is a junior synonym of Biplatyconcha (Briggs, 1998, p. 113).

Wyndhamia Booker, 1929, is closely allied to Biplatyconcha in terms of their similar internal features. However, the ventral or- nament of Wyndhamia includes thick spines interspersed with fine spines, whereas Biplatyconcha has small, close-set variable spines on the ventral valve according to Waterhouse (1978).

BILATYCONCHA GRANDIS (Waterhouse, 1975) Figure 4.1-4.4

Platyconcha grandis WATERHOUSE, 1975, p. 8, pl. 2, figs. 1-4; 1978, p. 73, pl. 10, figs. 11-14; pl. 11, figs. 1-5; BRIGGS, 1998, fig. 59.

Wyndhamia circularis Zhang (=Chang); ZHANG AND JIN, 1976, p. 166, pl. 2, figs. 1-15; text-fig. 3.

Megalosia chuluensis WATERHOUSE, 1988, p. 42, figs. 4, 5.

Material examined.-Two internal molds of ventral valves (NMV P303108, NMV P303109), and two worn dorsal valves (NMV P303110, NMV P303111).

Discussion.-Megalosia chuluensis Waterhouse (1988) is con- sidered to be a junior subjective synonym of Biplatyconcha gran- dis following Briggs (1998). Wyndhamia circularis Zhang (=Chang) in Zhang and Jin from the Selong Formation in south- ern Xizang (Tibet) is also referable to Biplatyconcha grandis in terms of similar outline, size, profile, short hinge, close-set dim- ples over the dorsal valve, and anteriorly thickened dorsal valve. The internal details of Wyndhamia circularis and Biplatyconcha grandis are comparable. The ventral spine pattern of Wyndhamia circularis is not preserved on the material of Zhang and Jin (1976). Aulosteges sp. from Suri Hill, Dingri, southern Xizang (Tibet) described by Yang and Zhang (1982, p. 311, pl. 1, figs. 2-4) is probably conspecific with the present species although the specimens are smaller in size.

Family ECHINOCONCHIDAE Stehli, 1954 Subfamily WAAGENOCONCHINAE Muir-Wood and Cooper, 1960

Genus WAAGENOCONCHA Chao, 1927 WAAGENOCONCHA? sp.

Figure 4.7-4.9 Material examined.-A severely worn internal mold of a con-

joined shell (NMV P303114). Discussion.-The lack of external ornament of the ventral

valve of this specimen makes this assignment questionable. The

FIGURE 4-1-4, Biplatyconcha grandis (Waterhouse); 1, internal mold of ventral valve, NMV P303108; 2, internal mold of ventral valve, NMV P303109; 3, dorsal valve with worn surface showing the median septum, NMV P303110; 4, dorsal exterior, NMV P303111. 5, 6, Taeniothaerus sp.; 5, dorsal interior, NMV P303112; 6, dorsal exterior showing dimples on the surface, NMV P303113. 7-9, Waagenoconcha? sp.; posterior, anterior, and lateral views of the internal mold of the ventral valve, NMV P303114. 10-14, Costatumulus shengmiensis n. sp.; 10, ventral valve, NMV P303116, paratype; 11, external mold of dorsal valve, NMV P303119; 12, external mold of dorsal valve, NMV P303117, paratype; 13, ventral valve, NMV P303118; 14, ventral valve, NMV P303115, holotype. 15, Costatumulus sp.; latex of ventral valve, NMV P303120, X4. (All figures life size unless otherwise stated)

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JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

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FIGURE 5-1-3, Pondospirifer cf. magnificus Waterhouse; 1, 2, ventral and posterior views of an incomplete ventral valve, NMV P303121; 3, a ventral valve with worn surface, NMV P303122. 4, 5, Costatispirifer? sp.; 4, external mold of the sulcal part of a ventral valve, NMV P303123; 5, latex of an external mold of a ventral valve, NMV P303124. 6, 7, Himathyris sp.; ventral and dorsal views of a conjoined shell, NMV P303129. 8-11, Fusispirifer sp.; 8, internal mold of an immature dorsal valve, NMV P303125, X1.7; 9, external mold of an immature ventral valve, NMV P303127, X1.8; 10, internal mold of an immature dorsal valve, NMV P303126, X2; 11, internal mold of ventral valve, NMV P303128. (All figures life size unless otherwise stated)

observed features of this single internal mold are comparable with those of Waagenoconcha imperfecta Prendergast (1935, p. 15, pl. 4, figs. 1-3; see also Archbold, 1993, p. 20, figs. 11A-H, 12A- K, 13A-G) from the Djhulfian (=Wuchiapingian) W. imperfecta Zone of Western Australia (Archbold, 1988), particularly the na- ture of the distinct sulcus, longitudinal curvature of the ventral valve, and the huge striated diductors in the ventral valve. W.

purdoni (Davidson, 1862, p. 31, pl. 11, fig. 5; Waagen, 1884, p. 705, pl. 73, figs. 1-3) from the Chhidru Formation in the Salt Range, Pakistan and also recorded from the Selong Formation in southern Xizang (Tibet) (Zhang and Jin, 1976, p. 176, pl. 5, figs. 1-3, 5, 6) is very close to the present specimen in size, outline, sulcus and profile, but further comparison is hampered because of the unknown surface ornament of the present material.

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SHEN ET AL.-PERMIAN BRACHIOPOD FAUNA FROM CHINA (TIBET)

TABLE 2-Measurements (in mm) of Quinquenella semiglobosa n.sp.

Registration no. Length Width Thickness Valves

NMV P303084 6.5 8.6 2.7 Ventral NMV P303085 6.2 8.1 Dorsal NMV P303086 6.1 9.2 Dorsal NMV P303087 7.3 9.2 2.2 Ventral NMV P303088 7.4 9.5 2.8 Ventral NMV P303089 6.3 8.6 2.5 Ventral NMV P303090 7.5 11.1 2.6 Ventral NMV P303091 7.3 10.1 2.7 Ventral NMV P303092 6.4 8.8 -Dorsal NMV P303093 5.6 8.6 Dorsal NMV P303094 7.1 8.7 Dorsal NMV P303095 6.3 9.3 - Dorsal NMV P303096 6.6 8.3 - Dorsal NMV P303097 6.3 7.7 - Dorsal NMV P303098 6.9 11.0 2.5 Ventral

Family LINOPRODUCTIDAE Stehli, 1954 Subfamily LINOPRODUCTINAE Stehli, 1954 Genus COSTATUMULUS Waterhouse, 1983b

COSTATUMULUS SHENGMIENSIS new species Figure 4.10-4.14

Diagnosis.-Large Costatumulus with very strong convexity, and prominent acute ears.

Description.-Shell large, 23.6 to 29.8 mm long, 29.4 to 54 mm wide and 12.8-22.0 mm thick; subquadrate or somewhat semielliptical in outline, strongly concavo-convex in profile; greatest width at hinge; ventral valve moderately geniculate; max- imum convexity in anterior part of venter; venter strongly inflated, somewhat globose in profile; beak low and thick, strongly in- curved; ears acute, slightly alate, flattened, and well demarcated from venter by a groove, and with strong rugae; umbonal slopes sharply inclined; sulcus totally absent; trail slightly flattened; dor- sal valve strongly geniculate; venter flat or slightly concave; trail and lateral margin sharply inclined; surface of both valves finely costellate; costellae numbering three in 2 mm on ventral trail; rugae stronger laterally, weakly across venter; spines not known along hinge or on ears, scattered on visceral disc and trail, with elongate base anteriorly up to 2 mm long, and sturdy swollen bases about 1 mm in diameter over trail; dorsal valve with dim- ples, but not spinose.

Etymology.-Referring to Shengmi, the type locality of the spe- cies.

Types.-Holotype, a nearly complete ventral valve (NMV P303115); paratypes, a complete ventral valve (NMV P303116) and a complete external mold of a dorsal valve (NMV P303117).

Other material examined.-A complete ventral valve (NMV P303118) and an external mold of a dorsal valve (NMV P303119).

Occurrence.-This species is from the lower part of the Qub- uerga Formation at Shengmi of Tingri County of southern Tibet.

Discussion.-This species is readily distinguished from other known species of Costatumulus by its large size, globose profile, and prominent acute ears. Costatumulus papilionata Waterhouse (1978, p. 109, pl. 21, figs. 11-21; pl. 22, figs. 1-3) from the Nambdo Member of the Senja Formation in northwest Nepal is close to the present species in its large size, flat dorsal valve and geniculate trail, but the ventral valve of the former is little inflated in profile and, has more densely-spaced spines and non-acute ears.

Family SPIRIFERIDAE King, 1846 Subfamily NEOSPIRIFERINAE Waterhouse, 1968

Genus PONDOSPIRIFER Waterhouse, 1978 PONDOSPIRIFER cf. MAGNIFICUS Waterhouse, 1978

Figure 5.1-5.3 cf. Pondospirifer magnificus WATERHOUSE, 1978, p. 93, pl. 16, figs. 1,

3, 4, 6, 7.

Material examined.-Two incomplete ventral valves with sur- face exfoliated (NMV P303121, NMV P303122).

Discussion.-The two available ventral valves closely resemble P. magnificus described by Waterhouse (1978) from the Nisal Member of the Senja Formation in northwest Nepal in all external features of ventral valve except the relatively small size. This species is also somewhat similar to Spirifer bambadhurensis de- scribed by Diener (1903, p. 108, pl. 4, figs. 8-10) in its outline, inconspicuous plications, swollen cardinal extremities and low and broad umbo, but S. bambadhurensis has finer costellae, a less prominent sulcus and a higher and more prominent umbo than the present species.

Genus COSTATISPIRIFER Archbold and Thomas, 1985 COSTATISPIRIFER? Sp.

Figure 5.4-5.5

Material examined.-Two fragments of external molds of ven- tral valves (NMV P303123, NMV P303124).

Discussion.-The two fragmentary external molds potentially represent a new species of Costatispirifer or an allied genus in terms of their extremely fine costellae on the ventral valve. The only known species of Costatispirifer, C. gracilis, from the Ufi- mian Lightjack Formation of the Canning Basin, Western Austra- lia, differs from the present Tibetan species in having about 20 costellae in 1 cm and a more or less weakly plicate surface.

Genus FUSISPIRIFER Waterhouse, 1966 FUSISPIRIFER sp. Figure 5.8-5.11

Material examined.-Two incomplete internal molds of im- mature dorsal valves (NMV P303125, NMV P303126); an exter- nal mold of an immature ventral valve (NMV P303127); and a huge incomplete internal mold of a ventral valve (NMV P303128).

Discussion.-The small immature specimens are probably con- specific with those described as Fusispirifer nitiensis by Diener (1897, p. 41, pl. 4, figs. 4, 5; 1899, p. 65, pl. 5, fig. 9) from Kiunglung near Niti Pass and from Nepal by Waterhouse (1966, p. 44, pl. 9, fig. 5; pl. 11, figs. 1, 3, 4; pl. 12, fig. 1; 1978, p. 94, pl. 17, figs. 1, 6) in terms of their similar outline and costation. The huge ventral valve (Fig. 5.11) is comparable with F. mar- couiformis Jin (=Ching) in Zhang and Jin (1976, p. 209, pl. 12, figs. 7, 9-11; pl. 13, figs. 1, 2, 21-23) from the Selong Formation in southern Xizang (Tibet) in terms of its large size.

ACKNOWLEDGMENTS

The senior author especially thanks Ge Yan who contributed to the success of the expedition and gave S. Shen much help during collection of fossils. The authors also greatly appreciate the indis- pensable logistic support including transport of fossils from China to Melbourne, given by Prof. Jin Yugan of the Nanjing Institute of Geology and Paleontology. Zhang Binggao of the Nanjing Institute of Geology and Paleontology of the Chinese Academy of Sciences has very extensive knowledge of the Qomolangma region, and pro- vided much advice before our expedition. J. B. Waterhouse provided Shuzhong Shen much helpful information for correlating the Permian System in Nepal with those in the Salt Range, Pakistan and Kashmir. R. Pierson of Deakin University carefully revised the first draft of this paper. We acknowledge J. A. Talent and B. R. Wardlaw for reviewing this paper. The work was supported by Major Basic Re- search Projects of MST (G2000077700) of People's Republic of Chi- na, CAS Hundred Talents Program in 1999 and the Australian Re- search Council.

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