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Earliest Wuchiapingian (Lopingian, Late Permian) Brachiopods in Southern Hunan, South China: Implications for the Pre-Lopingian Crisis and Onset of Lopingian Recovery/Radiation

September 30, 2008

By Shen, Shu-Zhong Zhang, Yi-Chun

ABSTRACT- The uppermost 5-15 m of the Douling Formation in the southern Hunan area. South China, yields a diverse fauna comprised of ammonoids, bivalves, and brachiopods. The brachiopods reported in this paper consist of 51 species in 34 genera and are dominated by the Lopingian (Late Permian) species associated with a few species persisting from the underlying Maokouan (Late Guadalupian). This fauna is of earliest Wuchiapingian in age as precisely constrained by the associated conodont Clarkina postbitteri postbitteri and the Guadalupian-type ammonoid fauna of the Roadoceras-Doulingoceras Zone in the brachiopod horizon. The discovery of the Lopingian species- dominated brachiopod fauna in the earliest Wuchiapingian in southern Hunan suggests a much less pronounced effect of the pre-Lopingian crisis (end-Guadalupian mass extinction) than the end-Changhsingian mass extinction in terms of brachiopods, a contemporaneous onset of the Lopingian recovery/radiation during the pre-Lopingian crisis period, and taxonomic selectivity of the pre-Lopingian crisis in terms of different fossil groups. New taxa are Echinauris doulingensis n. sp., Pararigbyella quadrilobata n. gen. and n. sp. and P. doulingensis n. gen. and n. sp.

INTRODUCTION

IT HAS been well known that the end-Permian mass extinction in the sense of Sepkoski (1981) actually consists of two phases, the pre-Lopingian crisis (end-Guadalupian mass extinction) and the end- Changhsingian mass extinction (Jin, 1993; Jin et al., 1994; Stanley and Yang, 1994; Shen and Shi, 1996, 2002; Shi et al., 1999; Wang and Sugiyama, 2000; Rong and Shen, 2002; Chen et al., 2005). To date, there are numerous studies on the PermianTriassic boundary and end- Changhsingian mass extinction in China (e.g., Yin et al., 1996; Jin et al., 2000; Shen et al., 2006a; Isozaki et al., 2007a). Relatively less has been studied in terms of the pre-Lopingian crisis. One of the reasons for the circumstance of the pre-Lopingian crisis is that a great global regression happened at the end-Guadalupian. This regression resulted in the withdraw of seawater from the supercontinent Pangea, which is stratigraphically recorded by the marine deposits of the Guadalupian overlain by Lopingian continental/ evaporite deposits (e.g., Russian Platform, Europe, Australia, West Texas, etc.) (Scotese and Langford, 1995). In the Paleotethyan region such as central Iran, South China, Pamir, Japan, etc., this regression is marked by an unconformity between the Guadalupian and Lopingian Series, although both the Guadalupian and Lopingian are marine deposits. Only the central part of the Jiangnan Basin in South China (e.g., the Laibin area in Guangxi of southwest China) possesses continuous marine deposition across the Guadalupian/ Lopingian boundary (Jin et al., 1998, 2006; Mei et al., 1998; Shen et al., 2007). Therefore, the biotas in the continuous sections between the Guadalupian and Lopingian are critical to understanding the end-Guadalupian mass extinction and the recovery/radiation after the pre-Lopingian crisis.

In this paper, a brachiopod fauna from the conodont Clarkina postbitteri postbitteri-beanng horizon is documented from the uppermost 5-15 m of the Douling Formation in southern Hunan, South China. This fauna, together with the bivalve (Fang, 1987) and ammonoid faunas (Zhou, 1987), provides significant data for understanding the pre-Lopingian crisis and the recovery/radiation of the Lopingian faunas after the pre-Lopingian crisis. The collection is from five localities in southern Hunan: Yejiwo, Yansha, Yanhu, Sanhe, and Xiaoyuanchong (Fig. 1; Table 1).

STRATIGRAPHY

The Upper and Middle Permian (Lopingian and Guadalupian) in southern Hunan are composed of chert and coal-bearing deposits of 415-1165 m in thickness. It is divided into four formations: Talung Formation, Douling Formation, Dangchong Formation, and Chihsia Formation, in descending order.

The Chihsia Formation is correlative with the units outcropping in other regions of South China and consists of dark gray limestone with cherty bands and nodules. This formation is largely Kungurian in terms of the fusulinids, ammonoids (Zhou, 1987), and conodonts (Mei et al., 2002), and is overlain by the Dangchong Formation, which is about 20-40 m thick and is composed of chert, siliceous limestone, and siliceous mudstone. In the Dangchong Formation, two early Maokouan ammonoid zones, the Altudoceras-Paraceltites Zone in the lower and the Quiyangoceras Zone in the upper, have been documented (Zhou, 1987).

The Douling Coal Series (Douling Formation in this paper) was first proposed by Tien (1929) and originally referred to a suite of clastic sedimentary rocks of Late Permian age intercalating with siliceous rocks of the Talung Formation in the upper and the Dangchong Formation in the lower. Sheng (1962) suggested using the Wuchiapingian Lungtan Formation to refer to the coal-bearing clastic deposits of Lopingian age in South China. Therefore, the Douling Formation was considered to be a synonym of the Lungtan Formation (Sheng, 1962). However, Meng (1980) discovered some Late Guadalupian (Maokouan) ammonoids including Altudoceras Ruzhentsev, 1940, Paraceltites Gemmellaro, 1887 and Cibolites Plummer and Scott, 1937, and considered the Douling Formation to be of late Maokouan age. On the basis of lithology and the presence/absence of coal seams, three lithostratigraphic members were subdivided. The lower member is about 150-300 m thick and composed of fine clastic sediments dominated by sandy shale interbedded with thin-bedded fine sandstone and siltstone; coal seams are absent. The middle member of the Douling Formation is about 70-400 m thick and consists mainly of feldspathic sandstone, sandy mudstone, and mudstone interbedded with several workable coal seams yielding a diverse flora. The uppermost member (5-15 m) of the Douling Formation is characterized by normal continental shelf deposits consisting of calcareous mudstone interbedded with thin-bedded bioclastic limestone. The base of this member is marked by a limestone bed immediately overlying a coal seam (Fig. 2). This member contains abundant marine invertebrate fossils, notably including ammonoids [e.g., Doulingoceras Zhou, 1985, Roadoceras Zhou, 1985, Paraceltites Gemmellaro, 1887 etc.) and bivalves [e.g., "Lima" Bruguiere, 1797, Paradoxipecten Zhang, 1981, Permoperna Nakazawa and Newell, 1968, Aviculopecten McCoy, 1851, Etheripecten Waterhouse, 1963 etc.]. The age of this fauna was regarded as Capitanian by Zhou (1987) and Fang (1987). However, as noted by Fang (1987), the bivalve assemblage clearly shows some mixed characters between the Lopingian and the Maokouan (Late Guadalupian) faunas. Brachiopods are abundant in the upper member. Meng (1980) listed six species, including Transennatia gratiosa (Waagen, 1884), Spinomarginifera lopingensis (Kayser, 1883), Haydenella kiangsiensis (Kayser, 1883), Edriosteges poyangensis (Kayser, 1883), Cathaysia chonetoides (Chao, 1927) and Leptodus nobilis (Waagen, 1883). Wang and Jin (1991) figured a specimen of Dicystoconcha lapparenti Termier et al., 1974 of Permianellidae He and Zhu, 1979 from the upper member of the Douling Formation. Shen and Archbold (2001) described five chonetid species from the upper member of the Douling Formation (Table 1). However, a complete systematic treatment of these brachiopods has not been published.

FIGURE 1-Map of the southern Hunan area showing the fossil localities studied in this paper.

The Talung Formation in the uppermost Permian in this area is about 50-120 m thick and dominated by gray-black siliceous shale and siliceous limestone. In the Talung Formation, an ammonoid succession, including Pleuronodoceras Chao, 1965, Pseudotirolites Sun, 1939, and Tapashanites Chao, 1965 in the upper and Konglingites Chao and Liang in Chao, 1966, Anderssonoceras Grabau, 1924, and Prototoceras Spath, 1930 in the lower, has been documented (Zhou, 1987), therefore suggesting that it ranges from Wuchiapingian to Changhsingian in age. Typical Changhsingian brachiopods in this area have been described by Shen and Shi (2007).

AGE AND CORRELATION OF THE BRACHIOPOD FAUNA

Together with the six previously-described species by Wang and Jin (1991) and Shen and Archbold (2001), a total of 51 species belonging to 34 brachiopod genera have been recognized from the upper member of the Douling Formation in the southern Hunan area (Table 1). The brachiopod fauna as a whole shows some mixed characters between the Maokouan (Late Guadalupian) and the Lopingian faunas but dominated by the Lopingian species, as indicated by the presence of many characteristic species such as Echinauris opuntia (Waagen, 1884), Edriosteges poyangensis, Neochonetes (Huangichonetes) substrophomenoides (Huang, 1932), Neochonetes (Zhongyingia) zhongyingensis Liao, 1980, Permophricodothyris squamularioides (Huang, 1933), Spinomarginifera lopingensis, Tethyochonetes soochowensis (Chao, 1928), Transennatia gratiosa (Waagen, 1884), and Tyloplecta yangtzeensis (Chao, 1927). Among them, Cathaysia chonetoides, Haydenella kiangsiensis, Spinomarginifera lopingensis, Tethyochonetes soochowensis, Transennatia gratiosa, and Tyloplecta yangtzeensis are the most common species in the Lopingian of South China. Paramarginifera kwangtungensis Zhan in Hou et al., 1979 and Haydenoides orientalis Zhan in Hou et al., 1979 have been reported from the Shuizhutang Formation in the neighboring Lianyang area in northern Guangdong Province. The Shuizhutang Formation is of late Wuchiapingian age as suggested by the presence of the ammonoids Prototoceras, Araxoceras Ruzhentsev, 1959, and Konglingites. Neoplicatifera huangi Jin et al, 1974 has been extensively documented from the Maokou Formation and its equivalents in South China. N. costata Ni in Yang et al., 1977 was known from the Maokou Formation in Songchi of Hubei Province (Jin et al., 1974). Spinomarginiferal sp. is most likely conspecific with those figured as S. qiaotingensis Jin et al., 1974, which was recorded from the Maokou Formation in Nanjiang, Sichuan Province. So far, these are the youngest record of the genus Neoplicatifera Jin et al., 1974.

The age of the present brachiopod fauna is of earliest Wuchiapingian age as well constrained by the association of the conodont Clarkina postbitten postbitten Mei and Wardlaw in Mei et al., 1994 in the limestone bed. According to Mei et al. (personal commun.), a conodont assemblage, including Clarkina postbitteri postbitten, Hindeodus sp. and Xaniognathus sp., has been found from a limestone bed in the upper member of the Douling Formation (see Fig. 2). The species Clarkina cf. bitten (Kozur, 1975) previously reported by Zhou and Gong (1994) from the Xiaoyuanchong section and the Matian section in the southern Hunan area can be re-assigned to Clarkina postbitteri postbitteri from the earliest Wuchiapingian at the Penglaitan section in Laibin and the Fengshan section in Liuzhou, Guangxi, South China (detailed locality position see Mei and Wardlaw, 1996 and Jin et al., 2006). The FAD of Clarkina postbitteri postbitteri at the Penglaitan section in Laibin, Guangxi, China has been ratified as the GSSP of the base of the Lopingian (Jin et al., 2006). Associated with this brachiopod fauna, an ammonoid zone, namely the Roadoceras-Doulingoceras Zone, has been recognized and previously assigned to Capitanian by Zhou (1987) and Zhou and Gong (1994). However, this ammonoid zone has been placed in the earliest Wuchiapingian based on the latest definition of the Lopingianbase GSSP (Jin et al., 2006). In addition, a bivalve “Lima”- Paradoxipecten Assemblage with 50 species was also documented by Fang (1987). This bivalve assemblage, as discussed by Fang (1987), clearly exhibits some mixed characters between the Maokouan and the Lopingian faunas. About 60% of species persist from the underlying assemblage in the middle member of the Douling Formation, but many Late Permian species such as Etheripecten sichuanensis Liu in Chen et al., 1974, Crenipecten exilis Liu in Chen et al., 1974, and Pernopecten symmetricus curtus Liu in Chen et al., 1974 also occur in the “Lima”-Paradoxipecten Assemblage.

TABLE 1-Brachiopod species from the upper part of the Douling Formation in five localities of southern Hunan. YS-Yansha, YH- Yanhu, XYC-Xiaoyuanchong, SH-Sanhe, YJW-Yejiwo. *-newcomers and very common species in the Lopingian in South China; #-common species in the Late Guadalupian in South China. The last six species are from Wang and Jin (1991) and Shen and Archbold (2001).

IMPLICATIONS OF THE PRE-LOPINGIAN CRISIS AND ONSET OF LOPINGIAN RECOVERY/RADIATION

The pre-Lopingian crisis (Jin, 1993) or the end-Guadalupian mass extinction (Jin et al., 1994; Stanley and Yang, 1994; Shen and Shi, 1996, 2002; Shi et al., 1999; Yang et al., 1999; Wang and Sugiyama, 2000; Shen et al., 2006b) was accompanied by a major regression, confined primarily to continental Pangea (Scotese and Langford, 1995). Like most other regions in the world, South China also experienced an uplift movement (the Dongwu Movement) at end- Maokouan (Hu, 1994). The end-Guadalupian mass extinction recorded at the Guadalupian/Lopingian unconformity resulted in the disappearances of many large fusulinids such as verbeekids and neoschwagerinids (Stanley and Yang, 1994; Shen and Shi, 2002; Isozaki et al., 2007b) and rugose corals (Wang and Sugiyama, 2000; Wang et al., 2006). These two fossil groups never recovered after the end-Guadalupian extinction. In contrast to fusulinids and corals, brachiopods and gastropods suffered less pronounced obliteration and immediately recovered during the early Wuchiapingian (Zhan in Hou et al., 1979; Liao, 1980, 1987; Pan and Erwin, 1994; Shen et al, 1992, 2006b; Erwin, 1996; Shen and Shi, 2002). Thus, a detailed comparison in taxonomic composition of each group between the Maokouan and the Lopingian is essential for recognizing the end- or Late Guadalupian changeover pattern. However, this has been long hampered for brachiopods due to the lack of the early Wuchiapingian data. The discovery of the present brachiopod fauna from the earliest Wuchiapingian in the southern Hunan area provides critical data for recognizing the diversity during the earliest Wuchiapingian after the end-Guadalupian extinction in South China and onset of the Lopingian recovery/ radiation within the pre-Lopingian crisis interval.

FIGURE 2-Permian stratigraphical column in the southern Hunan area indicating the brachiopod horizon and other associated fossil assemblages.

As listed in Table 1, 51 species in 34 genera have been found from the early Wuchiapingian upper member of the Douling Formation. Among them, 33 species are newcomers and become very common in the Lopingian in South China (Table 1). The discovery of this highly diverse fauna undermines the end-Guadalupian mass extinction in terms of brachiopod diversity, implying the contemporaneous recovery in the Lopingian crisis interval and indicating that the pre- Lopingian crisis (end-Guadalupian extinction) is probably not as severe as documented previously in South China (Shen and Shi, 1996, 2002; Shen et al, 2006b) and much less pronounced than the end- Changhsingian mass extinction, which eliminated the brachiopod orders Productida, Orthotetida, and Spiriferida, as well as many other fossil groups (Shen et al., 2006b; Wang et al., 2006). Nearly all common brachiopod families of Lopingian age were already present during the early Wuchiapingian. The sharp drop of early Wuchiapingian brachiopod species/genera number in South China previously recognized (Shen and Shi, 1996) may be partly derived from preservation bias or species area effect (Sepkoski, 1976). According to Raup (1976) and Sepkoski (1979), the estimated number of fossil species in each geological interval correlates strongly with measures of the amount of rock available for study. The end- Guadalupian regression event resulted in the late Wuchiapingian Lungtan Coal Measures directly and unconformably overlying the Late Guadalupian Maokou/Lengwu formations, with very restricted distribution of early Wuchiapingian marine strata. In the Laibin area, the early Wuchiapingian deposits are dominated by chert or siliceous limestone with rare brachiopods. Therefore, the Douling fauna is the ideal fauna to provide the diversity data of the early Wuchiapingian brachiopods in the world. Chen et al. (2005) reported brachiopods from the basal part of the Lungtan Formation near Chongqing City and these correlated with the earliest Wuchiapingian faunas. The contact between the Lungtan and the underlying Maokou formations at the Daijiagou section near Chongqing City is distinctly marked by a paleoweathering crust-type limonite bed followed by a bauxite bed, which apparently indicates an unconformity between the Guadalupian and Lopingian like that of most other areas in South China (Zeng et al., 1995). Chen et al. (2005) misidentified the occurrence of Roadoceras sp. of Sheng and Jin (1994), which was from southern Hunan, not the Huayingshan area in Chongqing City. Thus, no reliable brachiopod fauna has been reported from the Clarkina postbitteri positbitteri-beanng horizon of South China except for the Laibin (Shen in Jin et al., 2006) and southern Hunan areas reported in this paper.

A brachiopod changeover did occur in the Late Guadalupian in South China. Detailed vertical ranges of brachiopod genera (Shen and Shi, 1996; Rong and Shen, 2002, fig. 4; Shen et al., 2006b) indicate that the Maokouan brachiopod faunas are characterized by abundant Monticulifera Muir-Wood and Cooper, 1960, Cryptospirifer Grabau, 1931, Vediproductus Sarytcheva in Ruzhentsev and Sarytcheva, 1965, Urushtenoidea Jin and Hu, 1978 and Neoplicatifera. According to Shen and Shi (1996, 2002) and Shen et al. (2006b), the Maokouan/ Wuchiapingian boundary marks a distinct discontinuity with the above typically Maokouan genera terminated near the boundary except for Neoplicatifera and Urushtenoidea. The discovery of the Douling brachiopod fauna indicates that the pre-Lopingian crisis is only reflected at genus and species levels, and the Lopingian recovery/ radiation began in the earliest Wuchiapingian, as with the gastropods (Pan and Erwin, 1994; Erwin, 1996) and bivalves (Fang, 1987). It has been well documented that rugose corals and fusulinids suffered a pronounced extinction (Stanley and Yang, 1994; Yang et al., 1999; Wang and Sugiyama, 2000), and therefore the pre- Lopingian crisis (end-Guadalupian mass extinction) was taxonomically selective (Shen and Shi, 2002).

SYSTEMATIC PALAEONTOLOGY

For the sake of keeping the paper within a reasonable length, only new taxa are fully described below and in some cases descriptions and/or comments are also provided for some species and/ or genera where sufficient material is available and revision of these taxa is necessary. Some species in South China have been described in detail by Shen and Archbold (2001), Shen et al. (2004), Chen et al. (2005) and Shen and Shi (2007), and therefore are not repeated in this paper. All reported species are figured. All species described herein are from the upper member of the Douling Formation in the southern Hunan area and all specimens are housed in the Nanjing Institute of Geology and Palaeontology with prefix NIGP (Figs. 3-6). Order PRODUCTIDA Sarytcheva and Sokolskaja, 1959

Suborder CHONETIDINA Muir-Wood, 1955

Superfamily CHONETOIDEA Bronn, 1862

Family RUGOSOCHONETIDAE Muir-Wood, 1962

Subfamily QUINQUENELLINAE Archbold, 1981

Genus QUINQUENELLA Waterhouse, 1975

QUINQUENELLA? sp.

Figure 3.19

Discussion.-A possible species of Quinquenella is indicated by a complete internal mold of the ventral valve (NIGP130740). This mold is very small, measuring 5.0 mm long, 6.7 mm wide and 2.2 mm thick; nearly quadrate in outline, greatest width at hinge; ventral valve strongly convex in lateral profile, somewhat globose; beak wide, curved; ears small, acute and slightly convex; sulcus completely absent; external ornamentation not preserved, radially papillose; ventral interior with a very long median septum.

This single ventral valve is like some species of Quinquenella in terms of its strong convexity with no sulcus and a long median septum in the ventral valve. However, the dorsal valve is unknown and the absence of external ornamentation makes this assignment questionable. The species Quinquenella kuwaensis Waterhouse, 1978 from the Kuwa Member of the Senja Formation in northwest Nepal is similar to the present species in outline, but the Nepalese species is generally larger on average and further comparison is hampered because of the lack of dorsal valve.

Suborder PRODUCTIDINA Waagen, 1883

Superfamily PRODUCTOIDEA Gray, 1840

Family PRODUCTELLIDAE Schuchert in Schuchert and LeVene, 1929

Subfamily PRODUCTELLINAE Schuchert in Schuchert and LeVene, 1929

Tribe PARAMARGINIFERINI Lazarev, 1986

Genus PARAMARGINIFERA Fredericks, 1916

PARAMARGINIFERA? KWANGTUNGENSIS Zhan in Hou et al., 1979

Figure 4.7

Paramarginifera kwangtungensis ZHAN in Hou et al., 1979, p. 79, pl. 6, figs. 1-6; pl. 9, fig. 17.

Description.-Shell 16.0-22.3 mm long, and 26.0-31.7 mm wide, subquadrate in outline; widest at hinge; cardinal extremities nearly rectangular; ventral valve gently convex in lateral profile, sharply geniculated anteriorly; beak incurved, slightly beyond hinge line; ears small, gently convex, marked by several wrinkles, well demarcated from umbonal slope by a groove; sulcus narrow and shallow, originating from visceral region, not widening anteriorly, becoming deeper on trail; dorsal valve slightly concave; fold inconspicuous; surface of both valve with variable number of costae increasing by bifurcation; costae coarser anteriorly, about 5 per 5 mm; concentric lines well developed on visceral region and forming delicate reticulated ornamentation with costae, but not developed on trail; two coarse spines symmetrically on the anterolateral part of the flanks; ventral external mold with a prominent cincture along the geniculation, probably corresponding to some internal ridge.

Material examined.-Four ventral valves. Registered specimen (NIGP130754).

Discussion.-This species could represent a new genus allied to Paramarginifera or Retimarginifera. It was originally referred to Paramarginifera Fredericks by Zhan in Hou et al. (1979), probably in view of the cincture along the geniculation, which clearly suggests that some internal ridges are present. However, Paramarginifera has a strong tendency to form a tube on the anterior part of the trail, which is totally absent in the present species. This species appears to have some affinities with Retimarginifera in terms of its reticulation, the prominent internal ridge along the geniculation in the ventral valve and the absence of a tube on anterior part of the trail, but has much finer reticulation on the visceral disc.

Subfamily MARGINIFERINAE Stehli, 1954

Tribe MARGINIFERINI Stehli, 1954

Genus HAYDENOIDES Zhan in Yang et al., 1977

Diagnosis.-Externally like Haydenella, but interior of ventral valve with narrow ridges, extending inside ears to lateral or anterior margins; interior of dorsal valve with broad ridges crossing inside ears and extending around visceral disc.

Type species.-Haydenoides orientalis Zhan in Yang et al., 1977, p. 352, pl. 140, fig. 13; Lungtan Formation of the Lopingian; Lianxian, Guangdong, South China.

Distribution.-Wuchiapingian; Hunan and Guangdong, South China.

Discussion.-Haydenoides closely resembles Haydenella in outline, profile, ornamentation and spinosity, but differs in having ridges around the lateral and anterior margins of the visceral disc in both valves. Therefore, this genus may not be related to the genus Haydenella, which lacks ridges in both valves, although Zhan in Hou et al., (1979) assigned both to the same family. The presence of ridges in both valves seems to recall some genera of the subfamily Marginiferinae Stehli, 1954. Haydenoides was considered to be a synonym of Spinomarginifera Huang, 1932 by Brunton et al. in Williams et al. (2000), but can be readily distinguished by its rugae on its ears, absence of reticulate ornamentation and different spinosity. Haydenoides is somewhat similar to Productus Sowerby, 1814 in Sowerby (1812-1815) in its broad ridge in the dorsal valve, but the latter has much stronger costae and a very long trail.

FIGURE 3-1,-Lingula sp., ventral view of an external mold, x 6, NIGP130724, Yanhu; 2-5, Acosarina minuta (Abich); 2, internal mold of a ventral valve showing a long median septum to anterior margin, x 5.5, NIGP130725; 3, internal mold of a dorsal valve, x 5.5, NIGP130726; 4, ventral view of a ventral valve showing tubular costellation, x 3, NIGP130727; 5, internal mold of a ventral valve showing a median septum to midvalve, x 3, NIGP130728, Xiaoyuanchong; 6-7, Enteletes hemiplicata (Hall), ventral view showing the two dental plates and median septum and anterior view of a complete articulated shell, x 1.5, NIGP130730, Yansha; 8, Enteletes retardata Huang, internal mold of a ventral valve showing two dental plates and a median septum, x 3, NIGP130729, Yansha; 9- 10, Orthothetina regularis (Huang); 9, internal mold of a ventral valve showing two dental plates, x 3, NIGP130731; 10, ventral view of a ventral valve showing two slightly divergent dental plates, x 2, NIGP130732, Yansha; 11-12, Streptorhynchus sp. ventral and dorsal views of a ventral internal mold, x 2, NIGP130734, Yansha. 13, Streptorhynchus pelargonatus (Schlotheim), ventral view of a ventral internal mold, x 2, NIGP130733, Yansha; 14, Derbyia schellwieni Frech, ventral view of a ventral internal mold showing a median septum, x 1.5, NIGP130735, Yansha; 15-18, Neochonetes (Zhongyingia) zhongyingensis Liao; 15, ventral view of a complete ventral valve, x 2, NIGP130736; 16, external mold of a dorsal valve, x 3, NIGP130737; 17, external mold of a ventral valve, x 5, NIGP130739; 18, external mold of a dorsal valve, x 5, NIGP130738, Yansha; 19, Quinquenella? sp., internal mold of a ventral valve, x 5, NIGP130740, Yansha; 20-21, Edriosteges poyangensis (Kayser); 20, ventral view of an incomplete ventral valve, showing spines and concentric rugae on visceral region, x 1.5, NIGP130742, Yansha; 21, external mold of a ventral valve showing strong rugae on ears, x 1.5, NIGP130741, Yansha; 22, Cathaysia chonetoides (Chao), ventral view of a complete ventral valve, x 1.5, NIGP130743, Sanhe; 25-27, Haydenella kiangsiensis (Kayser); 23, ventral view of a complete ventral valve, x 2, NIGP130744, Xiaoyuanchong; 24, ventral view of a complete ventral valve, x 1, NIGP130745, Yansha; 25, ventral view of an articulated shell with broken umbo, x 2, NIGP130747, Yansha; 26-27, ventral and side views of a complete specimen, x 2, NIGP130746. 28, Haydenoides orientalis Zhan, internal mold of a ventral valve showing broad ridges acrossing inside ears and extending around visceral disc, x 1.5, NIGP130748, Yansha. (d.p.- dental plates; m.s.-median septum; l.r.-lateral ridge)

FIGURE 4-1-3, Neoplicatifera huangi Jin, Liao and Fang; 1, ventral view of a complete ventral valve, x 1.5, NIGP130751; 2, external mold of a ventral valve, x 1.5, NIGP130750; 3, external mold of a dorsal valve, x 1.5, NIGP130749, Yansha; 4-6, Neoplicatifera costata Ni; 4-5, ventral and anterior views of an external mold, x 3, NIGP130752; 6, internal mold of a complete ventral valve, x 3, NIGP130753, Yansha; 7, Paramarginiferal kwangtungensis (Zhan), internal mold of a ventral valve showing the prominent cincture along the geniculation, x 1.5, NIGP130754, Yejiwo; 8-12, Echinauris doulingensis n. sp.; 8-9, ventral and anterior views of an articulated shell, x 1.5, NIGP130759, Yejiwo; 70-72, anterior, dorsal and side views of an articulated shell, x 1, NIGP130758, holotype, Yejiwo; 13-19, Spinomarginifera lopingensis (Kayser); 13-15, anterior, side and ventral views of an articulated shell, x 1.5, NIGP130755; 16-19, anterior ventral, dorsal and side views of an articulated shell, x 1.5, NIGP130756, Xiaoyuanchong; 20- 22, Transennatia gratiosa (Waagen); 20, ventral view of a ventral internal mold, x 2, NIGP130762; 27-22, ventral and anterior views of a ventral external mold, x 3, NIGP130763, Yansha; 23-25, Echinauris opuntia (Waagen), ventral, side and anterior views of an articulated shell, x 2, NIGP130757, Xiaoyuanchong; 26-27, Spinomarginifera? sp.; 26, internal mold of a dorsal valve showing the mold of marginal ridge, x 3, NIGP130761; 27, external mold of a dorsal valve, x 3, NIGP130760, Yansha; 28-31, Tyloplecta? sp., anterior, ventral, side and dorsal views of a complete articulated shell, x 1.5, NIGP130766, Yansha. (m.p.-musclar platform; m.r.- marginal ridge)

HAYDENOIDES ORIENTALIS Zhan in Hou et al., 1979

Figure 3.28

Haydenoides orientalis ZHAN in Hou et al., 1979, p. 83, pl. 5, figs. 6-9, 11-12.

Description.-Shell 15.0 mm long, 23.9 mm wide and 6.5 mm thick; transversely semielliptical in outline; hinge slightly shorter than greatest width at middle shell length; ears small, flat and marked by several concentric rugae; cardinal extremities nearly rectangular; ventral valve strongly convex in lateral profile, maximum convexity at visceral disc; beak small, inconspicuous and not turning over hinge line; trail moderately geniculated; sulcus absent; dorsal visceral region uniformly concave; trail strongly geniculated; ventral surface with low and rounded costae; costae usually irregular, not developed on apical region, increasing by bifurcation, numbering 6 in 5 mm on trail; interspaces shallow and narrow; trail with several large spines; dorsal interior with a marginal plate. Material examined.-An internal mold of a ventral valve (NIGP130748).

Discussion.-The single mold is completely comparable with the types figured by Zhan in Hou et al. (1979) in internal details. This species is easily confused with Haydenella kiangsiensis in external characters if the internal marginal ridges are not revealed. However, H. kiangsiensis does not have a marginal ridge in either valve.

Subfamily OVERTONIINAE Muir-Wood and Cooper, 1960

Tribe COSTISPINIFERINI Muir-Wood and Cooper, 1960

Genus NEOPLICATIFERA Jin et al., 1974

NEOPLICATIFERA HUANGI Jin et al., 1974

Figure 4.1-4.3

Plicatifera minor (Schellwien); HUANG, 1932, p. 38, pl. 3, figs. 1-4.

Plicatifera hungi USTRITSKY ET AL., 1960, p. 25, pl. 3, figs. 7- 9; GRUNT AND DMITRIEV, 1973, p. 95.

Neoplicatifera huangi (Ustritsky); JIN ET AL., 1974, p. 309, pl. 162, figs. 4-7; YANG ET AL., 1977, p. 345, pl. 138, fig. 7; FENG AND JIANG, 1978, p. 251, pl. 89, fig. 15; JIN AND HU, 1978, p. 118, pl. 4, figs. 1-14; TONG, 1978, p. 221, pl. 78, fig. 24; WANG ET AL., 1982, p. 220, pl. 87, figs. 1-3; pl. 90, figs. 14-16; HU, 1983, pl. 3, figs. 1a-1b; ZHAO AND TAN, 1984, p. 26, pl. 1, figs. 12-14; YANG, 1984, p. 217, pl. 32, fig. 17.

Description.-Shell 17.0-18.0 mm long, 24.6-29.4 mm wide and about 6.5 mm thick; semicircular in outline; hinge slightly narrower than greatest width at shell midlength; ears small, flat, marked by rugae, demarcated by 4-5 spines from visceral region; cardinal extremities nearly rectangular; lateral sides broadly rounded; anterior margin evenly rounded; ventral valve moderately convex in lateral profile; maximum convexity at apical region; anterior slope gently convex; sulcus absent; dorsal valve slightly concave, maximum concavity at umbonal region; strongly geniculated anterior to visceral region; surface of both valves with concentric rugae and densely-spaced, erect and fine spines; rugae not developed on trail.

Material examined.-An external mold of a ventral valve (NIGP130749) and two external molds of dorsal valves (NIGP130750, 130751).

Discussion.-The type specimen of Plicatifera minor (Schellwien, 1903) from the Upper Carboniferous possesses regular and strong rugae and a few coarse spines on trail, and is very different from those figured by Huang (1932) from the Middle Permian Maokou Formation of South China. Therefore, Ustritsky et al. (1960) renamed Huang’s specimens as Plicatifera huangi Ustritsky et al., 1960. This species was proposed as the genus Neoplicatifera in terms of its irregular and weak rugae and fine spines on trail by Jin et al. (1974). The present species differs from N. sintanensis (Chao, 1927) from the Wushan Limestone (Maokou Formation) of Zigui, Hubei, in its larger size, presence of more rugae on the visceral region and indistinct costae on the venter.

NEOPLICATIFERA COSTATA Ni in Yang et al., 1977

Figure 4.4-4.6

Neoplicatifera costata NI in Yang et al., 1977, p. 346, pl. 139, figs. 4a-4c.

Description.-Shell small, about 10 mm long, 11-13 mm wide and 6- 9 mm thick; somewhat cylindrical; greatest width at hinge; cardinal extremities acute, forming an angle of about 70[degrees]; ventral valve strongly convex in lateral profile; longitudinal greatest curvature at posterior region; trail nearly flat and slightly geniculated; lateral slopes very steep; ears small, flat, demarcated by grooves from visceral region; beak pointed, strongly incurved, beyond hinge-line; sulcus absent; surface marked by rugae and erect spines; trail ornamented with wide and rounded costae and scattered spines; ventral interior with elongate and strongly elevated adductor.

Material examined.-Two internal molds of ventral valves (NIGP130752, 130753).

Discussion.-The shape, costae and spines of the present specimens are identical with the type figured by Ni in Yang et al. (1977). However, the surface of the visceral region is too poorly preserved to be compared with other species.

Genus ECHINAURIS Muir-Wood and Cooper, 1960

ECHINAURIS OPUNTIA Waagen, 1884

Figure 4.23-4.25

Productus opuntia WAAGEN, 1884, p. 707, pl. 79, figs. 1, 2; BROILI, 1916, p. 17, pl. 117, figs. 9, 10; HAMLET, 1928, p. 28, pl. 2, fig. 6.

Productus (Pustula) opuntia (Waagen); LICHAREW, 1937, p. 114, pl. 7, fig. 14.

Productus (Avonia) opuntia (Waagen); REED, 1944, p. 74.

Krotovia opuntia (Waagen); WATERHOUSE, 1966, p. 15, pl. 2, figs. 1, 5.

Echinauris opuntia (Waagen); GRANT, 1968, p. 27, pl. 8, figs. 1- 8; pl. 9, figs. 1-8; ZHANG AND JIN, 1976, p. 173, pl. 7, fig. 9.

?Echinauris sp., SHI AND SHEN, 1997, p. 44, fig. 3D.

Description.-Shell about 15 mm long, 18 mm wide and 7 mm thick, subquadrate to somewhat triangular in outline; hinge nearly equal to the greatest width; ventral valve moderately convex in profile; visceral region slightly flattened; beak thick and low; strongly incurved; umbonal and lateral slopes strongly inclined; ears small, commonly broken; demarcated from the visceral region by a groove with a row of halteroid spines; visceral region with numerous small spines; rugae and costae absent; trail also with small spines, spines curved forward; spinal bases slightly elongated; dorsal valve moderately concave; trail geniculated dorsally; dorsal interior with inconspicuous marginal ridge.

Material examined.-Six articulated shells: Registered specimen: NIGP130757.

Discussion.-The inconspicuous marginal ridge in the dorsal valve and the absence of rugae and costae on collected materials strongly suggest that they are referable to Echinauris opuntia rather than Spinomarginifera species. This species is externally somewhat similar to S. lopingensis in its outline and size, but S. lopingensis usually has more strongly incurved profile and prominent costae on trail surface.

ECHINAURIS DOULINGENSIS new species

Figure 4.8-4.12

Diagnosis.-Large Echinauris with numerous fine spines, no rugae and very weak marginal ridge.

Description.-Shell large for genus, 22.0-36.0 mm long, 24.3-33.4 mm wide and 12.6-20.2 mm thick; cylindrical; hinge nearly equal to greatest width; ears small, convex, usually marked by fairly long rhizoid spines and demarcated from umbonal regions; ventral valve strongly curved longitudinally; greatest curvature at posterior region; trail gently and evenly curved; visceral region gently and evenly convex and lateral slopes sharply inclined; beak pointed and strongly incurved, turning over hingeline; sulcus absent; dorsal valve deeply and evenly concave; moderately geniculated; ears small and flat; surface of both valve ornamented with densely spaced and fine spines; spines on visceral region and ears nearly erect; spine bases not inflated but elongated and forming intermittent costale appearance, numbering 4 in 5 mm; dorsal surface with numerous spines in quincunx; dorsal interior with weak marginal ridge; median ridge long; adductor flabellate and elevated; visceral region with numerous small endospines.

FIGURE 5-1-2, Tyloplecta yangtzeensis (Chao); 1, anterior view of an articulated shell, x 1, NIGP130764; 2, interior of a dorsal valve showing the cardinal process, cardinal ridges along hinge line, median ridge and brachial scars, x 1, NIGP130765, Yejiwo; 3, Gubleria huangi Wang, Jin and Fang, internal mold of a ventral valve showing bead-like median septum, x 1, NIGP130767, Yansha; 4, Leptodus sp., external mold of a dorsal valve, x 1, NIGP130768, Yansha, x 5-11, Pararigbyella quadrilobata n. sp.; 5, ventral view of internal mold of a ventral valve, x 2, NIGP130769; 6, dorsal view of external mold of same specimen, x 2; 7, ventral view of a complete internal mold of a ventral valve, x 2, NIGP130771; 8, dorsal view of internal mold of a dorsal valve, x 2, NIGP130772; 9, dorsal view of external mold of a ventral valve, x 2, NIGP130770; 10, internal mold of a dorsal valve, x 2, NIGP130773; 11, latex cast of the specimen NIGP130773, x 2, Yansha; 12-14, Pararigbyella doulingensis n. sp.; 12, internal mold of a ventral valve showing six lobes and five wider septa, x 2, NIGP130775; 13, enlargement of NIGP130775, showing pseudopunctatae, x 8; 14, internal mold of a complete ventral valve, x 2, NIGP130774, Yansha; 15-17, Uncinunellina timorensis (Beyrich), ventral, anterior and side views of a complete internal mold, x 3, NIGP130776, Yansha; 18-21, Terebratuloidea minor Waagen, ventral, dorsal, anterior and side views of a complete internal mold, x 2, NIGP130777, Yansha; 22-23, Permophricodothyris sp., ventral and side views of a complete internal mold, x 1, NIGP130778, Yansha; 24-30, Permophricodothyris squamularioides (Huang); 24-26, ventral, dorsal and side views of a complete internal mold, x 1.5, NIGP130779; 27-30, dorsal, ventral, anterior and side views of a complete internal mold, x 1.5, NIGP130780, Yansha; 31-32, Phricodothyris asiaticus Chao; 31, ventral view of an internal mold, x 3, NIGP130781; 32, dorsal view of a complete specimen, x 1.5, NIGP130782, Yejiwo; 33-34, Martinia orbicularis Gemmellaro, dorsal and anterior views of an internal mold, x 1.5, NIGP130783, Yansha. (a.s.-adductor scar; b.s.- brachial scar; c.p.-cardinal process; 11-16-lobes; l.r.-lateral ridge; m.r.-median ridge; m.sl.-median slit; l.sl.-lateral slit)

Etymology.-Referring to the type locality of the Douling Formation.

Type.-A complete articulated shell (NIGP130758).

Other material examined.-Six articulated shells and a dorsal valve. Registered specimen: a complete articulated shell (NIGP130759). Discussion.-The present specimens are suggestive of Spinomarginifera kueichowensis Huang, 1933 which commonly occurred in the Lopingian of South China, but the fine spines, uninflated spine bases, weaker concentric rugae and fine costae on the trail can easily distinguish it from the later. The new species is also somewhat similar to Echinauris opuntia in its outline and longitudinal curvature, but the former is larger and has more densely spaced spines and finer costae on its trail. The specimens figured as Spinomarginifera janus (Huang, 1933) by Zhu (1990) from the Tongziyan Formation in Fujian Province is largely comparable with this species in their spinosity, but they have a distinct marginal ridge.

Suborder LYTTONIIDINA Williams, Happer and Grant in Williams et al., 2000

Superfamily LYTONIOIDEA Waagen, 1883

Family LYTONIIDAE Waagen, 1883

Subfamily LYTONIINAE Waagen, 1883

Genus PARARIGBYELLA new genus

Type species.-Pararigbyella quadrilobata n. sp. from the upper member of the Douling Formation at Yansha, Hunan, South China.

Diagnosis.-Rigbyella-like lyttoniids with elongate triangular outline, moderately convex ventral valve, distinct median septum beginning from umbonal cavity, and symmetrically-distributed lateral septa in the ventral valve and correspondingly bifurcated dorsal lobes; septa much wider than sinues.

Discussion.-Terminology defined by Williams et al. (1965), Rudwick (1968), and Grant (1976) is used for the present genus. The raised ridges of the ventral valve are called septa. The spaces between the septa are called sinues. The extensions that cover the ventral sinues are lobes, and the indentations that fit around the septa are slits (Fig. 5). The present new genus can be readily distinguished from any other genera among lyttoniids by its distinct wide median septum within the ventral valve, the symmetrically- distributed lateral septa beginning much more anteriorly than the median septum, and bifurcated dorsal lobe system. Rigbyella Stehli, 1956 is most like Pararigbyella, but usually with a few radically- extended lobes which are rarely forking and subequal in strength and length to the median septum. Rigbyella tends to have a septum along midline, but never as distinct as in Pararigbyella. In addition, the septa in Rigbyella are nearly as wide as the sinues between them. The elongate triangular outline of Pararigbyella indicates that it is highly likely different from the cup-like shape of Rigbyella with a very strong cicatrix on the beak.

Paralyttonia Wanner in Wanner and Sieverts, 1935 is similar to Pararigbyella in its septal system but differs in that it also has a few radically-extended subequal septa, usually lacks a distinct median septum, and the crests of the septa are flattened to rather deeply depressed.

Pararigbyella also resembles some permianellids in view of their symmetrically bifurcated lobes, but permianellids are always bilobated, both valves are incised; their ventral valve usually has a thin knife-edged median septum or a central median platform.

Distribution.-This new genus is only found from the early Wuchiapingian upper member of the Douling Formation in southern Hunan, South China.

PARARIGBYELLA QUADRILOBATA new species

Figure 5.5-5.11

Diagnosis.-Small-sized Pararigbyella with symmetrically quadrilobated dorsal valve, elongate triangular in outline; lateral septa beginning at about midvalve; dorsal lobes bifurcated anteriorly.

Description.-Shell 5.5-11.3 mm long and 3.5-8.0 mm wide, elongately triangular in outline, greatest width always at anterior margin; attached by beak region; ventral valve moderately convex in lateral profile; beak acute; median septum distinct, bounded by two narrower symmetrical sinues; sinues even in width, bifurcated once at midvalve; crests of septa rounded; numbering 3 in total separating 4 lobes; dorsal valve thin, correspondingly consisting of two lobes only at posterior and bifurcated once at about midvalve.

Etymology.-Referring to the dorsal valve with four lobes.

Types.-Holotype, an internal mold of a ventral valve (NIGP130771, Fig. 5.7); paratype, an internal mold of a dorsal valve (NIGP130772, Fig. 5.8).

Other material examined.-An internal mold of a dorsal valve (NIGP130773) and two internal molds of ventral valves (NIGP130769130770).

Discussion.-This species differs from Pararigbyella doulin- gensis n. sp. described below by its small size and simpler septal system in the ventral valve.

PARARIGBYELLA DOULINGENSIS new species

Figure 5.12-5.14

Diagnosis.-Medium-sized Rigbyella with symmetrical hexalobated dorsal valve; elongately triangular in outline; dorsal lobes bifurcated twice anteriorly.

Description.-Shell 16.5-19.0 mm long and 11.2-13.6 mm wide; greatest width always at anterior; ventral valve gently convex; median septum distinct, bounded by two sinues in posterior one- third; two symmetrical pairs of lateral septa beginning from one- third of shell length; crests of all septa rounded; sinues bifurcated twice anteriorly, even in width, much narrower than septa; dorsal valve thin, consisting six narrow lobes; lobes extending forward regularly.

FIGURE 6-1-2, Juxathyris bisulcata (Liao), ventral and anterior views of an internal mold, X3, NIGP130791, Yansha; 3, Paraspiriferina multiplicata (Sowerby), posterior view of a complete internal mold, x 3, NIGP130786, Yansha; 4, Crenispirifer alpheus (Huang), ventral view of a complete internal mold, x 1.5, NIGP130786, Yansha; 5, Spiriferellinasp., ventral view of an internal mold, x 1.5, NIGP130787, Yansha; 6-7, Martinia semiplana Waagen, anterior and ventral views of a complete internal mold, x 3, NIGP130784, Yejiwo; 8, Crurithyris sp., dorsal view of an internal mold showing the slightly divergent brachial plates, x 5, NIGP130785, Yansha; 9-14, Juxathyris guizhouensis (Liao); 9-11, anterior, side and ventral views of an articulated shell, x 1.5, NIGP130789; 12-14, side, anterior, and ventral views of an articulated shell, x 1.5, MGP130788, Yansha; 75, Juxathyris? sp., ventral view of an articulated shell, x 1.5, NIGP130790, Xiaoyuanchong; 16-21, Hustedia remota (Eichwald); 16-18, ventral, dorsal and side views of a complete internal mold, x 3, NIGP130792; 19-21, dorsal, ventral and anterior views of a complete internal mold, x 3, NIGP130793, Yansha; 22-28, Hustedia lata Grabau; 22-25, ventral, dorsal, side and anterior views of an incomplete specimen, x 5, NIGP130795; 26-25, dorsal, ventral and anterior views of a complete internal mold, x 5, NIGP130794, Yansha; 26-29, Notothyris crassa Reed, ventral, dorsal, side, and anterior views of an articulated shell, x 5, NIGP130796, Yansha. (b.p.-brachial plates)

Etymology.-Referring to the type locality of the Douling Formation.

Types.-Holotype, an internal mold of a ventral valve (NIGP130775, Fig. 5.12).

Other material examined.-An internal mold of ventral valve (NIGP130774).

Discussion.-See Pararigbyella quadrilobata n. sp.

ACKNOWLEDGMENTS

The specimens described in this paper were collected by Meng Feng- Yuan, originally employed in the Fifth Team of the Coalfield Geology and Exploration of Hunan Province during the 1980s, and are at the late Prof. Jin Yu-Gan’s disposal. We appreciate Prof. Jin Yu-Gan, who generously gave us the collection to study and provided many constructive discussions. We thank Guang R. Shi and Susan Butts for their very careful review and kind comments and suggestions. This work is supported by the 973 Project of MST (2006CB806400) of China, the CAS/SAFEA International Partnership Program for Creative Research Teams and NSFC.

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