A New Brachiopod Fauna From the Early to Middle Permian of Southern Qinghai Province, Northwest China
By He, Weihong Shi, G R; Bu, Jianjun; Niu, Zhijun
ABSTRACT- A new brachiopod fauna is described from the Early and Middle Permian of Zadoi and Zhidoi counties, southern Qinghai (Changdu block), northwest China. This fauna includes 13 species in nine genera with Spinomarginifera concentrica n. sp. and Transennatia waterhousei n. sp. The Early to Middle Permian brachiopod fauna from southern Qinghai is very similar to the contemporary Cathaysian faunas of South China with which the new fauna shares 70 per cent of its species. On the other hand, the Qinghai brachiopods also demonstrate a significant link with the Permian brachiopod fauna of the Sino-Mongolian-Japanese Province in northeast China, as suggested by Marginifera septentrionalis and Attenuatella. In particular, the occurrence of the bipolar brachiopod genus Attenuatella suggests that southern Qinghai may have played an important role as a biogeographic stepping stone in the marine faunal interchanges between the northern and southern hemispheres during the Early and Middle Permian.
THE QINGHAI-TIBETAN Plateau is a complex tectonic entity and contains important geological information for understanding the evolution of the Palaeo-Tethys (Huang et al., 1984; Pan et al., 1997; Acharyya, 2005). The Permian brachiopod faunas and their palaeobiogeography from the southern part of the plateau have received attention recently (Shen et al., 2001 a, 200Ib, 2003a, 2003b, 2003c; Shi et al., 2003). However, the Permian brachiopod fauna from northern Qinghai-Tibetan Plateau, particularly in southern Qinghai, is less studied despite some previous reports on brachiopod taxonomy and biostratigraphy (Ching and Ye, 1979; Jin and Sun, 1981; Hu, 1983; Liu, 1991; Liao and Xu, 2002; Niu et al., 2003).
The aims of this paper are to describe a brachiopod fauna from the Early to Middle Permian of Zadoi and Zhidoi counties, southern Qinghai, and to discuss its palaeobiogeographical significance.
GEOGRAPHICAL AND GEOLOGICAL SETTING
The study area is situated 100 km east of Tanggula City, and 660 km southwest of Xining, the capital of Qinghai Province (latitude: 33[degrees]00′-34[degrees]00′, longitude: 93[degrees]00′- 94[degrees]30′) (Fig. 1). Tectonically, this area belongs to the Changdu terrane of Pan et al. (1997), which in turn is an integral part of the larger SangpanGarez Ecotope of Yin (1988). The studied sections include the Garizaren (GZ) and Danglangzhai (D) sections (Fig. 1).
Most brachiopod fossils were obtained from the Suojia and Garizaren formations of the Garizaren section (GZ), and the Gadikao Formation of the Danglangzhai section (D) (Fig. 2). Other fossils were collected from the Middle Permian of Mariadazhou (MA) and Xiqiasaisuqu (0155) (Fig. 1).
The Suojia and Garizaren formations in the area were named by Niu et al. (2006a). The Suojia Formation of the Garizaren section consists mainly of bioclastic micrite (Fig. 2) and contains brachiopods Haydenella kiangsiensis (Kayser, 1883) and Transennatia waterhousei Shi and He n. sp., corals and foraminifers. Additionally, the Suojia Formation contains the fusulinid Sumatrina annae assemblage, indicating a Capitanian age (Middle Permian) (Niu et al., 2006a).
The Garizaren Formation of the Garizaren section consists of mudstone, quartz arenite intercalated with andesite and micrite, and contains brachiopods Spinomarginifera lopingensis (Kayser, 1883), Acosarina minuta (Abich, 1878), Haydenella kiangsiensis and Transennatia waterhousei in the micrite (Fig. 2). The Garizaren Formation overlies the Wordian Jiushidaoban Formation (which contains a fusulinid Afghanella schencki-Neoschwagerina craticulafera assemblage), and is in turn overlain by the Capitanian Suojia Formation with a parallel disconformity (Niu et al., 2006a). Consequently, the Garizaren Formation is roughly assignable to the late Wordian or early Capitanian of the Middle Permian (Niu et al., 2006a).
The Gadikao Formation at the Danglangzhai section consists of quartz arenite and volcanic rocks (Fig. 2). Brachiopods Marginifera septentrionalis Tschernyschew, 1902, Hustedia remota (Eichwald, 1860), Spinomarginifera concentrica He and Shi n. sp., Marginifera cf. M. helicus (Frech, 1911), Tyloplecta yangtzeensis (Chao, 1927), and Perigeyerella sp. were mainly collected from the clastic rock of Member 1 (Fig. 2). The fusulinid Misellina-Schwagerina assemblage of Kungurian age has been recognized from the Gadikao Formation (Niu et al., 2006b).
The classification of Brachiopoda adopted herein follows Brunton et al. (2000) for Productida, Williams and Harper (2000) for Orthida, Williams et al. (2000) for Orthotetida, and Alvarez and Rong (2002) for Athyridida. All the specimens described in this paper are preserved in the Micropaleontology Laboratory, Faculty of Earth Sciences, China University of Geosciences, Wuhan, People’s Republic of China, with prefixes D for specimens of the Danglangzhai section, GZ for specimens from the Garizaren section, MA for specimens of Mariadazhou, and 0155 for specimens of Xiqiasaisuqu.
Phylum BRACHIOPODA Dumeril, 1806
Subphylum LINGULIFORMEA Williams et al., 1996
Order PRODUCTIDA Sarytcheva and Sokolskaya, 1959
Family PRODUCTELLIDAE Schuchert in Schuchert and LeVene, 1929
Subfamily MARGINIFERINAE Stehli, 1954
Tribe MARGINIFERINI Stehli, 1954
Genus SPINOMARGINIFERA Huang, 1932
Discussion.-Spinomarginifera is similar to Neoplicatifera Jin et al., 1974 in its irregular rugae over the umbo, but the former has a marginal ridge extending from posterior to the anterior parts of the visceral disk, while in the latter the marginal ridge does not extend anteriorly. Spinomarginifera is similar to Marginifera Waagen, 1884 in having wide and low costae on trail, but the former has irregular rugae on the umbo, and the latter has reticulated ornamentation.
SPINOMARGINIFERA CONCENTRICA He and Shi sp. nov.
Diagnosis.-Fine concentric lines, and low and wide costae on trail; rows of spines up flanks, no spine on hinge.
FIGURE 1-Location of studied area. GZ: Garizaren section; D: Danglangzhai section; MA: Mariadazhou Village; 0155: Xiqiasaisuqu Village.
Description.-Shell 10-13 mm long and 16-20 mm wide, subquadrate in outline, widest at hinge. Ventral valve strongly convex near geniculation; beak slightly protruding over hinge line; umbo slightly convex; ear small, slightly convex or cylindrically enrolled; flanks steep; geniculation posterior to venter; marginal ridge prominent on well-preserved trail; trail long; irregular, prominent concentric rugae on visceral disk, fine concentric lines on trail; costae low and wide on trail, numbering 4 in 5 mm, obscure on visceral disk; spines in prominent rows along flanks and others widely scattered on venter, none at hinge. Dorsal valve moderately concave and geniculate anteriorly; rugae uneven, interrupted by fine concentric lines anteriorly.
Etymology.-Referring to the fine concentric lines on trail.
Types.-Holotype an incomplete dorsal external mold (D-lf-4), paratypes complete ventral valve (D-lf-7) and a conjoined shell (D- If-I).
Other material.-Twenty-six specimens. Registered material: three external moulds of three dorsal valves (D-lf-100, D-lf-8, D-lf-5); a deformed ventral valve (D-16f-10); two incomplete ventral valves (D- If-11, D-16f-3).
Occurrence.-Southern Qinghai, northwest China; the Kungurian of the Early Permian.
Discussion.-Prominent and irregular rugae on visceral disks and the prominent marginal ridge suggests a species of Spinomarginifera. The present species is similar to Spinomarginifera kueichowensis (Huang, 1932) from the Permian coal-bearing beds of Chengyaoyen, Tatinghsien, Kueichow, West China in terms of its transverse outline, convexity, and irregular rugae on the visceral disk but differs from the latter in having a posteriorly narrow umbo, absence of costae and presence of elongate spine bases. Fine concentric lines on trail also distinguish the present species from others of this genus.
SPINOMARGINIFERA LOPINGENSIS (Kayser, 1883)
Productus nystianus KON, var. lopingensis KAYSER, 1883, p. 187, pl. 28, figs. 1-5.
Marginifera lopingensis (Kayser) CHAD, 1927, p. 153-156, pl. 16, figs. 8-12.
Spinomarginifera lopingensis (Kayser) ZHANO AND CHINO, 1961, p. 412, pl. 4, figs. 26-33; WANG, JIN, AND FANG, 1964, p. 312-313, pl. 49, figs. 21-23; LIAO, 1987, pl. 5, figs. 5, 8, 9; ZHAN, 1989, pl. 26, figs. 1-8.
Material.-Eight specimens. Registered material: a conjoined shell (GZ4f-14); an incomplete ventral valve (GZ-4f-15); an interior of a dorsal valve (GZ-4f-13); an external mould of a dorsal valve (GZ-4f- 12); an interior of a dorsal valve and a dorsal view of an incomplete dorsal valve (D-37f-20).
Occurrence.-South China; Permian.
Discussion.-The present species is consistent with Spinomarginifera lopingensis (Kayser, 1883) from the Upper Carboniferous of Leping, Jiangsi Province, China, having moderate size, a width slightly larger than length, concavo-convex profile, numerous coarse spines on valves, and brachial interior. The present species differs from Spinomarginifera concentrica from the Gadikao Formation of Danglangsai, Zadoi County, Qinghai Province (this paper) in terms of its narrow umbo and coarser costae and spines.
Genus MARGINIFERA Waagen, 1884
Discussion.-Marginifera is similar to Spinomarginifera in its subquadrate outline, numerous spines on ventral valve, prominent marginal ridge, and dorsal endospines in rows, but Marginifera has stronger, more prominent costae and reticulated ornamentation; Spinomarginifera has weak costae or none, and irregular rugae on the umbo. Marginifera differs from Probolionia Cooper, 1957 in possessing more prominent costae, especially on trail, and more than one row of spines along the flanks. Marginifera differs from Marginoproductus Tan, 1986 in having fewer but stronger costae. MARGINIFERA CF. M. HELICUS (Frech, 1911)
Description.-Shell 14 mm long and 10 mm wide, elongate and ovate in outline. Ventral valve strongly and evenly inflated; beak strongly enrolled; umbo strongly incurved; flanks steep; sulcus absent; costae even in width and height, originating from umbo, extending anteriorly, numbering 6 in 5 mm; spines in two rows along each flank, quincunxial and rounded spines scattered on venter, spines on anterior part of venter becoming elongate; marginal ridge prominent.
Material.-One complete ventral valve (D-lf-24).
Occurrence.-Southern Qinghai, northwest China; the Kungurian of the Early Permian.
Discussion.-The present species is similar to Marginifera helicus (Frech, 1911) from the Permian of Leping, Jiangxi, South China as both have an elongate outline, prominent and regular costae on venter, quincunxially arranged spines, and a prominent marginal ridge, but the former differs inits small size and umbo narrowing more promptly towards the beak. The present species is similar to Spinomarginifera chengyaoensis Huang, 1932 from the Coal Series of the Permian at Chengyaoyen, Tatinghsien, Guizhou Province, China in its elongate outline, strongly convex profile, and spines, but the present species has no wrinkles on the umbonal flanks and prominent and regular costae on the venter.
MARGINIFERA SEPTENTRIONALIS Tschernyschew, 1902
Marginifera septentrional is TSCHERNYSCHEW, 1902, p. 646-649, pl. 36, figs. 10-12; TINO, XIA, DUAN, Li, LIU, AND LIANG, 1985, p. 111, pl. 45, figs. 11-15; CHAO, 1927, p. 163, pl. 16, figs. 34-37.
FIGURE 2-Stratigraphic columns of the two studied sections (Garizaren and Danglangzhai sections) and brachiopod species ranges in these sections.
Marginifera sp. Gu AND ZHU, 1985, pl. 1, figs. 22-24.
Material.-Fifteen specimens. Registered material: five incomplete ventral valves (D-lf-18, D-lf-19, D-lf-22, D-37M7, D-lf-23); a ventral view of an incomplete ventral valve (D0 -37I0 -40).
Occurrence.-Permian; Lindong and Zhesi of Neimong, Shenzha of Tibet, Jisu Honguer of Mongolia, and south Urals of Russia.
Discussion.-The present species is consistent with Marginifera septentrionaux Tschernyschew, 1902 from the Lower Permian of the Urals, as both are characterized by a small size, transversly subquadrate outline, wide and shallow sulcus, weak costellae, a row of spines between the ears and visceral disk, and a strong spine near the end of each ear. This species also compares with Marginifera septentrionalis of Chao, 1927 from the Permian Jisuhonguer limestone (now Zhesi Formation) of Jisu Honguer, Inner Mongolia in its external morphology including the small shell size, shell outline and ornamentation of numerous fine, forward- projecting, pustular spines and pustule-pits, and fine radiating costae. Marginifera septentrionalis differs from Marginifera typica Waagen, 1883 from the Permian of the Salt Range by its strong spine near the end of each ear, and from M. drastica Grant, 1976 from the Rat Buri Limestone, Phangnga, southern Thailand in possessing a more transverse outline and less convex lateral profile.
FIGURE 3-1-13, Spinomarginifera concentrica He and Shi sp. nov.; 1, D-16f-10, posterior view of a ventral valve, x 3; 2, D-1f-100, posterior view of external mould of a dorsal valve, x 3.5; 3, D-1f- 11, ventral view of a ventral valve, m represents marginal ridge, x 2; 4, D-1f-8, posterior view of external mould of a dorsal valve, x 3.2; 5, D-1f-5, posterior view of external mould of a dorsal valve, x 3.2; 6, D-1f-7, paratype, ventral view of a ventral valve, m represents marginal ridge, x 2; 7-8, D-1f-4, holotype, dorsal and posterior views of external mould of a dorsal valve, cl represents concentrate lines, x 3; 9-12, D-1f-1, paratype, posterior, lateral, dorsal and ventral views of a conjoined shell, cl represents concentrate lines, x 2.5. x 2, x 2, x 2.5; 13, D-16f-3, ventral view of a ventral valve, cl represents concentrate lines, x 3. 14- 15, Tyloplecta yangtzeensis (Chao, 1927); D-1f-43, posterior and ventral views of a ventral valve, x 1. 16-18, Marginifera cf. M. helicus (Frech, 1911); D-1f-24, ventral, lateral and posterior views of a ventral valve, x 3.
MARGINIFERA ELONGATA (Huang, 1932)
Productus elongata HUANG, 1932, p. 23. pl. 1, fig. 14.
Marginifera elongata (Huang) TONG, 1978, p. 221-222, pl. 79, fig. 4.
Material.-Two specimens. Registered material: an incomplete ventral valve (0155-f-41).
Occurrence.-Middle and Late Permian; southwestern China.
Discussion.-The present species is consistent with Marginifera elongata (Huang, 1932) from the Middle Permian, Lopotung, Hsifenghsien, Guizhou Province, South China in terms of its elongated outline, strongly convex ventral valve, parallel flanks, geniculation located at posterior of the venter, and coarse costae and spines. This species is similar to M. drastica Grant, 1976 from the Rat Buri Limestone, Phangnga, Southern Thailand in an elongate outline, and low and wide radial ribbing on the ventral valve surface, but differs in a more inflated umbo.
Subfamily PRODUCTININAE Muir-Wood and Cooper, 1960
Tribe CHONETELLINI Licharew in Sarytcheva et al., 1960
Genus HAYDENELLA Reed, 1944
HAYDENELLA KIANGSIENSIS (Kayser, 1883)
Productus kiangsiensis KAVSER, 1883, p. 185, pl. 26, figs. 6-11.
Haydenella kiangsiensis (Kayser) MUIR-WOOD AND COOPER, 1960, p. 224-225, figs. 1-14; ZHAN, 1979, p. 81, pl. 5, figs. 3, 4; LIAO, 1980, pi. 6, fig. 27; YANG, YIN, WU, YANO, DING, AND XU, 1987, pl. 10, figs. 26, 27; pl. 11, figs. 1, 2, 5, 6.
Material.-Three ventral specimens. Registered material: an incomplete ventral valve (GZ-13f-45).
Occurrence.-Late Permian; South China and Pakistan.
Discussion.-The present species is totally comparable with H. kiangsiensis (Kayser, 1883) from the Lower Permian of Leping, Jiangxi, South China in size, the rounded outline, highly convex ventral valve, weak capillae, and rugae on the ears. This species is similar to Haydenella qinglongensis Liao, 1980 in its highly convex ventral valve, weak capillae, and spinose ornamentation, but differs in its larger size and rounded outline. H. kiangsiensis may be distinguished from H. buravasi Grant, 1976 by its more globose outline and a more inflated umbo.
Subfamily MARGINIFERINAE Stehli, 1954
Tribe PAUCISPINIFERINI Muir-Wood and Cooper, 1960
Genus TRANSENNATIA Waterhouse, 1975
Discussion.-Transennatia is similar to Retimarginifera Waterhouse, 1970 in the reticulated ornamention, but Transennatia has stronger costae with a semicircular cross section and smaller ears, while Retimarginifera has subdued ornamention and more extended ears. Transennatia is similar to Caricula Grant, 1976 as both are characterized by reticulated ornamention, but Caricula has strong rugae and weak costae that smooth down within the sulcus, and more extended ears.
TRANSENNATIA WATERHOUSEI He and Shi new species
Diagnosis.-Fine and dense reticulation on dorsal visceral disk, obscure reticulation on ventral disk, fewer costae on anterior part of valves, and intensive anterior thickening of costae.
Description.-Shell 12-15 mm long and 17-24 mm wide, transverse in outline, concavo-convex in profile, widest at hinge. Ventral valve strongly convex; geniculate at posterior to median length; beak slightly protruding over hinge; umbo flat; ears moderate, convex, triangular; sulcus prominent, widest at median sulcus; strong costae crossed by obscure rugae on visceral disk, costae numbering 3 in 2 mm in the posterior part, and intensively furcated anteriorly, all numbering about 20. Dorsal valve moderately concave; geniculate at median length; umbo flat; ears moderate, triangular; fold prominent; strong semicircular costae crossed by fine and sharp rugae, forming even and dense reticulation on visceral disk, and those costae nearby fold converged onto fold, and those far from fold continuing to anterior and increasing in height and width anteriorly, all numbering about 14-18 on anterior part.
Etymology.-For Dr. J. B. Waterhouse, in honor of his contributions to the studies of Permian brachiopods.
Types.-Holotype a complete dorsal external mould (GZ0 -20f-25), paratype an incomplete ventral valve (GZ-20f-27).
Other material.-Eight specimens. Registered material: a ventral valve (GZ-20f-29); an incomplete ventral valve (GZ-20f-26); two incomplete dorsal external moulds (GZ-20f -28, GZ-20f-101).
Occurrence.-Southern Qinghai, northwest China; the Capitanian of the Middle Permian.
Discussion.-This species is similar to Transennatia gratiosus (Waagen, 1884) from the Wargal and Chhidru Formation of the Salt Range, Pakistan in moderate size, transverse outline, and sharply reticulated ornamention on the dorsal disk, but it differs in having larger ears, a more concave dorsal valve, a dorsal fold which is wider at midlength and narrower at ends, costae thickening anteriorly, fewer costae on the trail (more than 30 costae on the trail of T. gratiosus), and obscure reticulation on the ventral disk. It is similar to Transennatia insculpta (Grant, 1976) from the Rat Buri Limestone of southern Thailand in transverse outline, a dorsal fold which is wider at midlength and narrower at ends, and reticulated ornamentation on the dorsal valve, but differs in a larger size, stronger costae than rugae (the latter having costae as thick as rugae), more intensively thickening of the costae anteriorly, fewer costae on the anterior part (more than 30 costae on anterior of T. insculpta), and obscure reticulation on the ventral disk. The present species is also somewhat similar to T. cf. insculpta (Grant, 1976) of Sone et al., 2003 from the Middle Permian of Johore, Peninsular Malaysia in outline and sculpture of the dorsal valve, but the latter has numerous prominent rugae on the ventral visceral disk. Family PRODUCTIDAE Gray, 1840
Subfamily LEIOPRODUCTINAE Muir-Wood and Cooper, 1960
Tribe TYLOPLECTINI Termier and Termier, 1970
Genus TYLOPLECTA Muir-Wood and Cooper, 1960
TYLOPLECTA YANGTZEENSIS (Chao, 1927)
Figure 3.14, 3.15
Productus yangtzeensis CHAO, 1927, p. 50-54, pl. 5, figs. 1-3.
Tyloplecta yangtzeensis (Chao), TING, 1965, p. 268-269, pl. 3, figs. 1, 2; LIAO, 1980, pl. 4, figs. 21-22; NAKAMURA, SHIMIZU, AND GOLSHANI, 1981, p. 44-45, pl. 1, figs. 1-3; pl. 3, figs. 1, 2; LIAO, 1987, p. 104, pl. 5, figs. 23-24.
Material.-One ventral valve (D-1f-43).
FIGURE 4-1-10, Spinomarginifera lopingensis (Kayser, 1883); 1-4, GZ-4f-14, ventral, dorsal, posterior and lateral views of a conjoined shell, x 2; 5-7, GZ-4M5, lateral, posterior and ventral views of a ventral valve, x 2; 8, GZ-4f-13, interior of a dorsal valve, x 3; 9, D-37f-20, 9a, interior of a dorsal valve, 9b, dorsal view of an incomplete dorsal valve, x 2; 10, GZ-4f-12, external mould of a dorsal valve, x 3. 11-16, Transennatia waterhousei He and Shi sp. nov.; 11, GZ-20f-27, paratype, ventral view of a ventral valve, x 2; 12, GZ-20f-26, posterior view of a ventral valve, x 3; 13, GZ-20f-29, posterior view of a dorsal valve, x 2; 14, GZ-20f- 25, holotype, dorsal view of a dorsal external mould, x 2; 15, GZ- 20f-28, dorsal view of an incomplete dorsal external mould, x 2; 16, GZ-20f-101, dorsal view of a dorsal external mould, x 2. 17- 19, Haydenella kiangsiensis (Kayser, 1883); GZ-13f-45, posterior, ventral and lateral views of an incomplete ventral valve, x 2. 20, Linoproductidae gen. and sp. indet.; MA-13f-48, ventral view of a ventral valve, x 1.0.
Occurrence.-Middle to Late Permian; Asia and Europe.
Discussion.-This species is consistent with Tyloplecta yangtzeensis (Chao, 1927) from the Permian of Fengcheng, Kiangsi Province, South China in subquadrate outline, strong costae narrowing upwards, fine radial ribs, and coarse pustules on the posterior shell surface. Compared to Tyloplecta richthofeni (Chao, 1927) from Wushan Formation, Hubei Province, this species has thicker costae and a wider umbo. It also differs from Tyloplecta liziyaensis Zeng et al., 1995 from the Chihsia Formation of Liziya, Huayin Mountains, Sichuan in its subquadrate outline and upward- narrowing costae. Both Tyloplecta nankingensis (Frech, 1911) from the Chihsia Formation, Anhui Province, South China and Tyloplecta subrichthofeni Zhao and Tan, 1984 from the Maokou Formation, Sangzhi, Hunan Province, compare with the present species in general aspects, but Tyloplecta nankingensis has more prominent reticulation on its posterior part, while Tyloplecta nankingensis is distinguished by having sparser costae. Compared with the present species, Tyloplecta minisulcata Zeng et al., 1995 from the Maokou Formation, Liziya, Huayin Mountains, Sichuan is smaller in size, has a deeper and broader sulcus, and costae that maintain a consistent thickness from base to crest.
Family MONTICULIFERIDAE Muir-Wood and Cooper, 1960
Subfamily MONTICULIFERINAE Muir-Wood and Cooper, 1960
Genus MONTICULIFERA Muir-Wood and Cooper, 1960
MONTICULIFERA SINENSIS (Frech, 1911)
Productus sinensis FRECH, 1911, p. 176, pl. 25, fig. 3.
Monticulifera sinensis (Frech), JIN, LIAO, AND FANG, 1974, p. 309- 310, pl. 162, figs. 12-13; ZHAO AND TAN, 1984, pl. 1, fig. 9.
Diagnosis.-Short, rounded, quincuncially arranged monticules on posterior part; fine and irregular capillae on valve; fine concentric lines overlapping capillae on trail and flanks.
Materials.-Three specimens. Registered material: one complete ventral valve (0155-f-44).
Occurrence.-Middle Permian; Sichuan and Hunan provinces.
Discussion.-This species is probably conspecific with Monticulifera sinensis (Frech, 1911) from the Schwagerina Bed of the Lower Permian, western Hubei in size, subquadrate outline, short and quincuncially arranged monticules, fine, irregular, and closely spaced capillae, and fine concentric lines. This species differs from Monticulifera convexa Zeng et al., 1995 from the Maokou Formation, Mulongdong, Huayin Mountains, Sichuan as it is smaller and has finer capillae and fine concentric lines on trail and flanks. Compared with Monticulifera baishagouensis Tong, 1978 from the Maokou Formation, Baishagou, Yuexi, Sichuan and Monticulifera minor Tong, 1978 from the Maokou Formation, Changjianggou, Shangsi, Guangyuan, Sichuan, Monticulifera sinensis also has finer costellae and more capillae on the venter, and finer concentric lines on the trail and flanks. The present species also differs from Monticulifera plicatiformis Ting, 1965 from the Lower Permian, Yangkang, Tianjun, Qinghai, western China in the absence of plicae in flanks, the presence of finer capillae, and fine concentric lines.
Family LINOPRODUCTIDAE Stehli, 1954
LINOPRODUCTIDAE gen. and sp. indet.
Description.-Shell 30 mm wide, subrounded in outline, corpus cavity deep. Ventral valve inflated at posterior part; umbo evenly convex, incurved posteriorly; rugae on ears and flanks; fine costellae bifurcated, costellae numbering 7 in 5 mm in middle length.
Material.-One ventral valve (MA-13f-48).
Occurrence.-Southern Qinghai, northwest China; the Middle Permian.
Discussion.-Medium size, subrounded outline, deep corpus cavity, costellae fine and bifurcated, and rugae on the ears of ventral valve basically characterize a species of Anidanthus Hill, 1950; however, this specimen is difficult to compare with any species of Anidanthus because of poor preservation of the ears and missing dorsal valve.
Order ORTHOTETIDA Waagen, 1884
Superfamily ORTHOTETOIDEA Waagen, 1884
Family MEEKELLIDAE Stehli, 1954
Subfamily MEEKELLINAE Stehli, 1954
Genus PERIGEYERELLA Wang, 1955
Discussion.-Perigeyerella differs from Geyerella Schellwien, 1900 in having one order of costellation, and from Orthothetina Schellwien, 1900 in having a conical umbo.
Description.-Shell 22 mm long and 22 mm wide, conical in outline. Beak twisted slightly and deformed by cicatrix of attachment; pseudodeltidium with prominent median crest; interarea high, slightly concave; one rank of finely costellae.
Material.-Two incomplete ventral valves (D-1f-39, D-1f-40).
Occurrence.-Southern Qinghai; the Kungurian of the Early Permian.
Discussion.-Moderate size, conical outline and one rank of costellae suggest a species of Perigeyerella Wang, 1955. This species differs from Geyerella Schellwien, 1900 in its less conical outline and only one order of costellae. Compared to Perigeyerella tricosa Grant, 1976 from the Rat Buri Limestone, Ko Muk, Southern Thailand, the present species has a more conical posterior part of the ventral valve.
Order ORTHIDA Schuchert and Cooper, 1932
Family SCHIZOPHORIIDAE Schuchert and LeVene, 1929
Genus ACOSARINA Cooper and Grant, 1969
ACOSARINA MINUTA (Abich, 1878)
Streptorhynchus peregrinus van minutus ABICH, 1878, p. 78, pl. 9, fig. 1a.
Orthis indica WAAGEN, 1884, p. 568, pl. 56, figs. 7, 8, 14-16.
Orthotichia indica (Waagen) ZHAN AND LI, 1962, p. 473, pl. 1, figs. 1, 2.
Schizophoria indica (Waagen) WANG, JIN, AND FANG, 1964, p. 134, pl. 16, figs. 24, 25, 28.
Acosarina dorsisulcata COOPER AND GRANT, 1969, p. 2, pl. 5, figs. 19-23; COOPER AND GRANT, 1976, p. 2621, pl. 667, figs. 1-26; FENG AND JIANG, 1978, p. 235, pl. 85. fig. 10.
Acosarina indica (Waagen) YANG, NI, CHANG, AND ZHAO, 1977, p. 311, pl. 130, fig. 3; ZHAO AND TAN, 1984, pl. 1, fig. 1.
Acosarina minuta (Abich) SHI AND SHEN, 1998, p. 506-507, fig. 3.7- 3.10.
Material.-Three specimens. Registered material: two conjoined shells (GZ-4f-46, GZ-4f-47).
Occurrence.-This species has been found in the Upper Permian of the Transcaucasus, the Wargal Formation of the Salt Range, the Yenduyet Formation of Son La, northwest Vietnam, and the Lower Permian to the Early Triassic of South China.
Discussion.-This species differs from Acosarina strophiria Xu and Grant, 1994 from the Changxing Formation, Huangzhishan, Wuxi, Zhejiang, and from the Changxing Formation, Huayin Mountains, Sichuan, in having a shorter hinge line, narrower interarea, and a lower width to length ratio, and differs from Acosarina antesulcata Waterhouse (1983) from the Huai Tak Formation of northwest Thailand as the latter has an anterior sulcus on both valves. Compared to Acosarina tumita Zeng et al., 1995 from the Chihsia Formation of Liziya, Huayin Mountains, Sichuan, the present species is characterized by a lower ratio of thickness to length and finer and denser costellae. Acosarina regularis Liao, 1980 from the Wuchiapingian of Jiaozishan, Anshun, Guizhou, China may be also compared with this species in general morphology, but the former has a dorsal fold, a greater length to width ratio (i.e., longer than wide) and spinose ornamentation on the anterior costellae.
FIGURE 5-1-7, Marginifera septentrionalis Tschernyschew, 1902; 1, D-1f-18, ventral view of an incomplete ventral valve, X2; 2, D-lf- 19, posterior view of an incomplete ventral valve, X2; 3, D-lf-22, ventral view of an incomplete valve, X2; 4, D-37f-17, posterior view of an incomplete ventral valve, X2; 5-6, D-1f-23, posterior and ventral views of an incomplete ventral valve, X2; 7, D-37f-40, ventral view of an incomplete ventral valve; s represents spine, X2. 8-10, Marginifera elongata (Huang, 1932); 01550 -I0 -41, ventral, posterior and lateral views of an incomplete ventral valve, X2. 11, Monticulifera sinensis (Frech, 1911); 01550 -f-44, ventral view of an incomplete ventral valve, X1. 12-14, Perigeyerella sp.; 12-13, D- lf-39, ventral and posterior views of an incomplete ventral valve, X2; 14, D-lf-40, posterior view of an incomplete ventral valve, X2. 15-18, Hustedia remota (Eichwald, 1860); 15-17, D-lf-62, ventral, lateral and dorsal views of a conjoined shell, X 2.8; 18, D-lf-63, dorsal view of a conjoined shell, X 2. 19-22, Acosarina minuta (Abich, 1878); 1920, GZ-4f-46, lateral and dorsal views of a conjoined shell, X3; 21-22, GZ-4I-47, ventral and dorsal views of a conjoined shell, X2. FIGURE 6-Schematic reconstruction of Middle Permian paleobiogeography and tectonic configuration of East Asia (adopted from Shi, 2006). Symbols for tectonic blocks are as follows: Bs, Beishan area of NW China; Ca, Jiangnan oldland (southeastern China); Hg, Hida Gaien Belt (Terrane); Ic, Indo-China Block; Ir, Iran; Jg, Junggar Block (massif); Kg, Kurosegawa Belt (Terrane); Ls, Lhasa Terrane; Me, Central Mongolia-Ergun Terrane; PNG, Papua New Guinea; Qd, Qaidam Terrane; Ql, Qilian Mountains; Qt, Qiangtang Terrane; Sg, Sanpang-Garze Block (fold belt); Sk, South Kitakami Belt; Sm, Simao Terrane; St, Shan-Thai Terrane; Tp, Turpan Terrane; Xk, Xingkai (Khanka) Terrane; Ws, West Sumatra. Numbered and highly generalized occurrences of fossil sites for Attenuatella and Marginifera septentrionalis are explained as follows: 1-North Thailand; 2-Southwest China; 3-Study area (this study); 4-South Kitakami block (northeast Japan); 5-Northeast China; 6-North Mongolia; 7-Kolyma-Omolon Massif; 8-Verkhoyan Mountains (fold belt); 9-northwest Russia (e.g., Taimyr Peninsula).
Order ATHYRIDIDA Boucot et al., 1964
Family NEORETZIIDAE Dagys, 1972
Subfamily HUSTEDIINAE Grunt, 1986
Genus HUSTEDIA Hall and Clarke, 1893
HUSTEDIA REMOTA (Eichwald, 1860)
Rhynchonella remota EICHWALD, 1860, p. 768, pl. 35, fig. 10.
Retzia radialis var. grandicosta DAVIDSON, 1862, p. 28, pl. l, fig. 5.
Eumetria grandicosta (Davidson) WAAGEN, 1883, p. 491, pl. 34, figs. 6-12.
Retzia (Hustedia) grandicosta (Davidson) FRECH, 1911, p. 117, pl. 16, figs. 7, 8.
Hustedia remota (Eichwald) HUANG, 1933, p. 79, pl. 11, figs. 4, 5; GRABAU, 1934, p. 103, pl. 7, fig. 3; SHEN, SHI, AND ARCHBOLD, 2003b, p. 250, pl. 5, figs. 22-23.
Hustedia grandicosta (Davidson) GRABAU, 1934, p. 105, pl. 7, fig. 5; LI AND Gu, 1976, p. 275, pl. 160, fig. 17; LIAO AND MENG, 1986, pl. 4, figs. 2-4.
Materials.-Three specimens. Registered material: two complete conjoined specimens (D-1f-62, D-1f-63).
Occurrence.-This species has been widely found in the Permian of Asia and Europe.
Discussion.-This species is similar to Hustedia orbicostata Xu and Grant, 1994 from the Changxing Formation of Yutangjiao section, Nantong, Sichuan Province, of Beipei section, Sichuan Province, and the Lower Yinkeng Formation of Huangzhishan, Wuxi, Zhejiang Province, in shell outline and strong costae, but differs by its slightly larger size, slightly shorter ventral beak, lower ventral interarea, quadrate foramen, and more costae. It is also similar to Hustedia deminuta Wang, 1956 from the Upper Carboniferous sequence of Yantai Colliery, Liaoning Province, China in outline, lateral profile, the shape of costae, and ventral beak, but differs in its larger size and fewer costae.
The southern Qinghai brachiopod fauna consists of 13 species, among which nine species are commonly found in South China, including Spinomarginfera lopingensis, Marginifera cf. M. helicus, Marginifera elongata, Haydenella kiangsiensis, Tyloplecta yangtzeensis, Monticulifera sinensis, Perigeyerella sp., Acosarina minuta, and Hustedia remota. This list accounts for 70 per cent of the total species diversity. This would suggest that the Qinghai brachiopod fauna has a strong affinity with the South Chinese fauna of the warm-water paleoequatoral Cathaysian Province (Shi et al., 1995) (see also Fig. 6). However, the presence of Marginifera septentrionales in the Qinghai brachiopod fauna raises a very interesting paleobiogeographical point. This species, from the Gadikou Formation of the present study, has hitherto been found mainly in the middle paleolatitudinal regions of the Northern Hemisphere (Fig. 6), including the Lower Permian of the Urals (Tschernyschew, 1902) and the Lower to Middle Permian of Inner Mongolia (Ting et al., 1985; Gu and Zhu, 1985). It has never been found from South China. According to Shi et al. (1995) and Shi (2006), much of the Inner Mongolia region of northeast China belongs to the Sino-Mongolian-Japanese Province, of a transitional character in that it is characterized by a paleobiogeographically mixed marine fauna with both paleoequatorial Cathaysian and high paleolatitude Boreal faunal elements. This means that despite its strong affinities with the Cathaysian Province of South China, the Qinghai brachiopod fauna also maintained some biogeographical links with the Sino-Mongolian-Japanese Province located in a meso-thermal or temperate zone between the Boreal and Paleoequatorial realms during the Permian (Shi, 2006).
The above suggestion is also corroborated by the occurrence of another brachiopod genus, Attenuatella. Although this genus has not been found in the present fauna, it has previously been reported by Ching and Ye (1979) from the Lower Permian of Jiuzhi County, southern Qinghai. Attenuatella is usually found in association with Permian faunas of middle to high latitudes in both hemispheres (Tazawa, 1987) (Fig. 6), although it has also been found from lower latitude regions during the Late Permian (Lopingian) (Waterhouse, 1983; He et al., 2007). As such, the occurrence of this genus in southern Qinghai also might suggest that the Qinghai brachiopod fauna maintained some biogeographical links with the Sino-Mongolian- Japanese Province during the Permian where this genus also occurs. Since Permian Attenuatella also occurs in northern Thailand, southern Tibet, eastern Australia, and New Zealand (see He et al., 2007 for the latest review of the spatio-temporal distribution of Attenuatella), it is thus reasonable to suggest that southern Qinghai may have played an important role as a biogeographic stepping stone in the marine faunal interchanges between the northern and southern hemispheres during the Permian.
The authors owe thanks to Duan Qifa, Wang Jianxiong, Bai Yunshan, Tu Bing, He Longqing, and Zeng Bofu for help collecting brachiopod fossils. We are grateful to Shen Shuzhong and Yang Deli for suggestions on brachiopod identification, and to Dr. A. Perez- Huerta and Dr. R Isaacson for their review of the initial manuscript. This paper has been supported by NSFC (Grant Nos 40502001, 40621002), a research grant from the Australian Research Council (DP0772161 to GRS), support from the Chinese Academy of Sciences (Grant 2006-1-16 to GRS), a grant from the Ministry of Education of China (Grant No. 200228), and Program for Innovative Research Team in University (Grant No. IRT0546).
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ACCEPTED 12 DECEMBER 2007
WEIHONG HE,1 G. R. SHI,2 JIANJUN BU,1,3 AND ZHIJUN NIU1,3
1 Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Wuhan 430074, PR China,
2 School of Life and Environmental Sciences, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood, VIC 3125, Australia,
Copyright Paleontological Society Jul 2008
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