Chonetoidea (Brachiopoda) from the Lopingian (Late Permian) of South China

Chonetoidea (Brachiopoda) from the Lopingian (Late Permian) of South China

SHUZHONG SHEN & NElL W. ARCHBOLD

SHEN, S.-Z. & ARCHBOLD, N. W., 2002.01.31. Chonetoidea (Brachiopoda) from the Lopingian (Late Permian) of South China. AIcheringa 25. 327-349. ISSN 0311-5518.

Ten species of the superfamily Chonetoidea fiom the Lopingian (Late Permian) of South China are described or revised. A review of all recorded Chonetoidea species from the Lopingian (Late Permian) of South China indicates that some 22 species of five genera can be recognised. Species of Tethyochonetes and Neochonetes are characteristic in the lithofacies dominated by unudstone, siltstone or siliceous rocks in the Lopingian and some argillaceous limestone and clay rock facies near the Permian-Triassic boundary. New taxa are Neochonetes (Zhongyingia) subgen, nov., Neochonetes (Eluangichonetes) subgen, nov. and Tethyochonetes flatus sp. nov.

S.-Z Shen*, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 Eastern Beijing Road, Nanjing 210008, P.R. China; N.W. Archbold, School of Ecology and EnvironmenL Deakin University, Rusden Campus, 662 Blackburn Road, Clayton, Victoria 3168, Australia [[email protected]]; received 24 March, 2000: accepted 27 November, 2000: *corresponding author [[email protected]]

Keywords: Chonetoidea, Brachiopoda, Lopingian, Late Permian, South China

SPECIES of the superfamily Chonetoidea (Brachiopoda) are common in the Lopingian (Late Permian) faunas of South China. Their abundance in terms of number of specimens as well as survivorship into the earliest Triassic has stimulated the interest of many geologists. Despite sporadic reports and brief descriptions (e.g., Huang 1932; Chao 1928; Liao 1979, 1980a, 1980b, 1984; Zhan in Hou et al. 1979; Xu in Yang et al. 1987; Xu & Grant 1994) and the establishment of Tethyochonetes Chen et al.,

2000, study of the South Chinese Lopingian Chonetoidea as a group has been virtually neglected and is out of date. In this paper, we describe the chonetoideans from three sections (Fig. 1) in South China. The geographic distribution, stratigraphic distribution, and a complete taxonomic revision of all species of Chonetoidea recorded from the Lopingian of South China are also provided.

The repository of all specimens used in this study is the Nanjing Institute of Geology and 0311/5518/2001/03327-23 $3.00 © AAP

Palaeontology of Chinese Academy of Sciences, Nanjing, P.R. China with the prefix NIGP.

Stratigraphy According to the international standard of the Permian System (Jin et al. 1997), the uppermost Permian series, i.e., the Lopingian Series, is constrained by the base of the conodont Clarkina postbitteri Zone at the base and the base of the conodont Hindeodus parvus Zone at the top. The Lopingian is subdivided into Wuchiapingian and Changhsingian Stages in ascending order, of which each consists of two substages (see Jin et al. 1999, table 2). The Lopingian Series in South China varies considerably in lithofacies. The fossils described in this paper come from three sequences with different lithofacies (Fig. 2). At the Beifengjing section in Zhongliang Hill near Chongqing City (Fig. 1A), the Lopingian includes the Longtan Formation in the lower and the Changhsing Formation in the upper. The Longtan Formation

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328 SHUZHONG SHEN & NElL W. ARCHBOLD AL CHERINGA

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is a coal-bearing sequence mainly consisting of clastic sedimentary rocks with abundant plant fossils but rare marine fossils. The Changhsing Formation is mostly of limestone with over 180 species ofbrachiopods listed by Shen & He (1991) and Shen et al. (1992). Chonetoidean fossils are found from three horizons (Fig. 2).

At the Wenjiangsi section about 5 km northwest of Guiding, Guizhou (Fig. 1B), the Lopingian includes the Wuchiaping Formation of Wuchiapingian age, and the Changhsing and Talung Formations of Changhsingian age. Brachiopods are extremely abundant in the Changhsing and Talung Formations.

measured sections described in this study. A- Beifengjing

Chonetoidean fossils studied herein are from Bed 45 of the Changhsing Formation and Bed 55 of the Talung Formation (Fig. 2).

The Lopingian at the Xiaoyuanchong section in Jiahe, Hunan (Fig. 1 C) includes coal-bearing deposits in the Douling Formation followed by siliceous rocks in the Talung Formation (Fig. 2). The lower part of the Talung Formation yields the ammonoids Konglingites, Anderssonoceras and Prototoceras (Zhou 1987), which apparently suggest a late Wuchiapingian age. The upper part of the Talung Formation is probably of Changhsingian age as constrained by the overlying Griesbachian (Earliest Triassic) bivalve


ALCHERINGA LATE PERMIAN CHONETOIDEA FROM SOUTH CHINA 329

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Pseudoclaraia wangi Zone and the underlying late Wuchiapingian ammonoids. However, careful biostratigraphic analysis is required from which the precise ages may be determined. The Douling Coal Series (Douling Formation) originally referred to a suite of clastic sedimentary rocks of Late Permian age intervening between the siliceous

rocks o f the Talung Formation and the Dangchong Formation (Tien 1929). Three lithostratigraphic members are recognised within the Douling Formation, of which the uppermost member is characterised by normal continental shelf deposits consisting of calcareous mudstone with bioclastic limestone lens. The base of this


330 SHUZHONG SHEN & NEIL W. ARCHBOLD ALCHERINGA

Wuchiapingian

Fusichonetes nayongensis Neochonetes (Z.) zhongyingensis

Tethyochonetes soochowensis Neochonetes (H.) substrophomenoides

Tethyochonetes flatus Neochonetes (S.) strophomenoides

Tethyochonetes chaoi Tethyochonetes liaoi

Tethyochonetes quadrata Tethyochonetes wongiana

Tethyochonetes guizhouensis Tethyochonetes sp. A

Tethyochonetes sp. Tethyochonetes dissulcata

Fanichonetes campigia Neochonetes (H.) cursothornia

?Fanichonetes striata ?Pygmochonetes sp.

Tethyochonetes pigmaea ?Fanichonetes. sp.

Tethyochonetes ?longtanensis Tethyochonetes sp.

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member is usually marked by a coal seam (Fig. 2). This member contains abundant marine invertebrate fossils including ammonoids and bivalves. A conodont assemblage, including Clarkina postbit teri , Hindeodus sp. and Xaniognathus sp. has been found from the limestone lens in Bed 13 of the upper member (Mei, pers. comm. 1998), hence indicating that the upper member of the Douling Formation is of early Wuchiapingian because the base of the Clarkina postbitteri Zone has been considered as the boundary between the Late Permian Lopingian and the Middle Permian Guadalupian (Jin et al. 1997). Chonetoidean fossils from this section are all found from Bed 13 of the Douling Format ion (Fig. 2). The coeval strata corresponding to this member are absent at the Beifengjing and Wenjiangsi sections due to the well-known uplift Dongwu Movement in these regions.

Discussion

A complete list of previously and herein recorded chonetoidean species from the Lopingian of South China is given in Appendix 1. All belong to the Rugosochonetidae except Pygmochonetes sp., which has been assigned to the Anopliidae by Racheboeuf(2000). Some 22 species belonging to five genera are recognised. Lopingian chonetoideans are characteristically dominated by species of Tethyochonetes and Neochonetes. With the exception of Pygmochonetes sp., no species has been recorded from horizons older than the Lopingian. Five species in total persist into the earliest Triassic (Fig. 3, Appendix) because they occur in the horizons above the conodont Hindeodus parvus Zone (Liao 1979, Zhao et al. 1981). As shown in Figure 3, no restricted assemblages can be recognised based only on chonetoidean species (Fig. 3), but a few species appear to be useful in local and regional correlation. Tethyochonetespigmaea (Liao, 1979) is a useful index fossil for the Changhsingian and earliest Triassic. Neochonetes (Sommeriella)



strophomenoides (Waagen, 1884) may serve as an index fossil for the Lopingian (Late Permian) in South China and the Peri-Gondwanan Region. This species has been documented from the Wuchiapingian Kalabagh Member of the Wargal Formation in the Salt Range, Pakistan (Waagen 1884) and the Lopingian Qubuerga Formation at Shengmi in the Qomolangma region (Shen et al. 2001). Neochonetes (Huangichonetes) substrophomenoides and Tethyochonetes chaoi also range from Wuchiapingian to Changhsingian in South China. Fanichonetes species are restricted to Changhsingian and earliest Triassic.

Within all three Lopingian sequences examined herein, chonetoideans mainly occur in mudstone, siltstone and siliceous rocks in the Lopingian and argillaceous limestone and clay rocks near the Permian-Triassic boundary. They are typically associated with bivalves and lingulids near the Permian-Triassic boundary. Other associated brachiopods are commonly small and thin-shelled, which are mostly euryhaline taxa and, hence apparently indicating that their habits are adapted to the muddy, silty or siliceous substrate and were usually near a source of sediment adjacent to an old landmass or a site of volcanic activity (Xu & Grant 1994).

Brachiopod assemblages dominated by small thin-shelled chonetoideans associated with lingulids and Crurithyris appear to characterise proximity to the Permian-Triassic boundary in South China. Similar assemblages have also been documented from the Selong Xishan section in South Tibet (Shen & Jin 1999), the Salt Range, Pakistan (Grant 1970); Kashmir (Nakazawa et al. 1975) and southern Alps (Broglio-Loriga et al. 1986). This assemblage is usually found in finely laminated lithologies that are interpreted to have accumulated in low-oxygen conditions. Evidence of this is the large amount of framboidal pyrite preserved within the rock. Brachiopods of this assemblage have been considered as Palaeozoic dysaerobic taxa that were able to tolerate the deteriorating sea-floor oxygen conditions that appear to have occurred globally at the very end of Permian through to earliest Triassic time (Hallam & Wignall 1997). Interpreta t ion of this

assemblage may fit the scenario preferred by Hallam & Wignall (1999) of a rapid transgression at the very end of the Permian that produced a worldwide anoxic environment, causing the end- Permian mass extinction of many marine faunas.

Systematic palaeontology

Occurrences of species in South China have been shown in Appendix 1, so are not repeated in the description below. The systematic study follows the classification of Racheboeuf (2000). All specimens were whitened before photographed.

Suborder CHONETIDINA Muir-Wood, 1955 Superfamily CHONETOIDEA Bronn, 1862 Family RUGOSOCHONETIDAE Muir-Wood, 1962 Subfamily RUGOSOCHONETINAE Muir-Wood, 1962

Neochonetes Muir-Wood, 1962

Type species. Chonetes dominus King, 1938, p. 259, pl. 36, figs 1-7. Late Carboniferous, West Texas (USA).

Remarks. Medium to large rugosochonetids with a feeble to distinct sulcus are usually referred to Neochonetes (Muir-Wood, 1962). Neochonetes was a highly diverse genus during the Late Palaeozoic and numerous species are recognised (Archbold 1981). This genus ranges from Late Carboniferous to Permian and several groups or stocks are recognised (Archbold 1981). One dist inct ive group o f Neochonetes with a conspicuous sulcus, gentle dorsal fold and maximum width in mature shells usually anterior o f the hinge was named as Neochonetes (Sommeriella) with Chonetes prattii (Davidson, 1859) as type species (Archbold 1981, 1982). Two other subgenera, N. (Nongtaia) and N. (Zechiella), have subsequently been recognised by Archbold (1999) on the basis of material from the Rati Buri Group (Roadian/Wordian) of southern Thailand and the Zechstein


332 SHUZHONG SHEN & NEIL W. ARCHBOED ALCHERINGA



(Wuchiapingian) of Germany.

Distribution. Late Carboniferous to Late Permian; cosmopolitan.

Neochonetes (Zhongyingia) subgen, nov.

Type species. Neochonetes zhongyingensis Liao, 1980a, p. 257, pl. 5, figs 10-13. Changhsingian (Late Permian), South China.

Etymology. Zhongying, a village in Guizhou Province of South China.

Diagnosis. Neochonetes species with reverse trapezoidal outline and acute ears; greatest width always at hinge; cardinal extremities extended; hinge spines projecting posterolaterally with angle less than 45°; ventral valve with only slight convexity; sulcus very shallow and broad; surface finely costellate. Dorsal interior with long lateral ridges parallel to hinge.

Remarks. Archbold (1981) recognised several groups within Neochonetes. The Neochonetes granulifer (Owen, 1852) group includes N. granulifer Owen, 1852 and N. dominus King, 1938. Both are subquadrate shells with small ears, moderately convex profile, and appear to be quite different from Neochonetes (Zhongyingia). However, N. granulifer var. emaciatus King, 1938, N. fragilis King, 1938 and N. puebloensis King, 1938 are closer to N. (Zhongyingia) in that they all have reverse trapezoidal outline, a very gently convex ventral valve with a shallow, broad sulcus and extended cardinal extremities. The only representative of Neochonetes from the Permian of New Zealand, N. beatusi Waterhouse, 1964,

may also belong to N. (Zhongyingia) on the basis o f its outline and ventral sulcus, but it is a large species. One o f the specimens figured as N. (Sommeria) prattii (Davidson, 1859) by Archbold (1981, fig. 6J, K, non A-I, L-T) also recalls N. (Zhongyingia) in view of its extended ears and broad sulcus, but it is an extremely transverse individual and hence an aberrant representative o f Davidson's species. A third group, based on N. carboniferus (Keyserling, 1846) (see Archbold 1981, p. 111, table 2), is closely allied to N. (Zhongyingia) with its hinge spines set at a low angle and a broad, shallow sulcus, but it is less transverse, somewhat subquadrate in outline and more convex in profile.

Distribution. Late Carboniferous to Late Permian; South China, USA.

Neochonetes (Zhongyingia) zhongyingensis Liao, 1980a (Fig. 4A-Q)

1980a Neochonetes zhongyingensis Liao, p. 257, pl. 5, figs 10-13.

Material. Four internal moulds of ventral valves (NIGP 132460-132463), two external moulds o f ventral valves (NIGP132464, 132465), eight external moulds of dorsal valves (NIGP 132466- 132473) and an internal mould of a dorsal valve (NIGP132474) from Bed 13 of the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan.

Description. Shell large, reverse trapezoidal in outline; widest at hinge; cardinal extremities extended; ears large, acute; anterior broadly rounded; lateral margins almost straight or

Fig. 4. Neochonetes (Zhongyingia) zhongyingensis Liao. A, B, internal mould of a ventral valve, latex of the same specimen, NIGP132460, x2. C, F, internal mould of a ventral valve, latex of the same specimen, NIGP132462, x2, x2.3+ D, latex of an external mould of ventral valve, NIGPI32465, x2.5. E, latex of an external mould of ventral valve, NIGPI32464, x3.8. G, latex of an internal mould of ventral valve, NIGPI32461, xl.7. H, latex of external mould of dorsal valve, showing the ventral interarea, N1GPI32466, × 2. I, latex of an external mould of dorsal valve, showing the interareas of both valves and the quadrilobate cardinal process, NIGPI32467, x2.5. J, N, latex of an external mould of dorsal valve and external mould of the dorsal valve, N1GPI32473, x2.8. L, latex of an external mould of dorsal valve, N1GPI32469, x2. K, external mould of dorsal valve, NIGPI32472, x2. M, external mould of dorsal valve, NIGPI32471, x2.5. O, latex of an external mould of dorsal valve, NIGP132470, x3.8. P, Q, internal mould of dorsal valve, latex of the same specimen, NIGP132474, x3.



slightly concave. Ventral valve gently to moderately convex in profile; beak low and broad; interarea linear; sulcus broad and shallow, beginning from anterior to umbo, rapidly widening anteriorly. Dorsal valve gently concave; fold very low; ears fiat. Ventral hinge with 4-6 pairs of spines; spines projecting posterolaterally at an angle of 30-60°; costellae fine, increasing by bifurcation and intercalation, numbering about 40 near margin. Ventral interior with median septum; median septum extending about one- third of shell length and higher to posterior; at anterior two short parallel vascular trunks form weak ridges adjacent to septum. Dorsal interior with externally quadrilobate cardinal process; lateral septa short, divergent, arising anterior of alveolus; median septum arising anterior of lateral septa, extending two-thirds of shell length; sockets deep; inner socket ridges prominent, not united with base of cardinal process, extending into lateral ridges until ears; remaining inner surface radially papillose. Dimensions: adult ventral length, 9.4-11.2 mm; dorsal length, 9.1- 13,8 mm; maximum width, 16.1-21.1 ram; thickness, 2.0-2,6mm.

Remarks . Neochone tes (Sommeriel la) strophomenoides (Waagen) described below is readily distinguished from the present species by its more convex profile, subquadrate outline and usually greatest width anterior to hinge. Neochonetes (Zhongyingia) species from the Late Carboniferous of USA (King 1938) differ from the present species in their finer costellation. Specimens described as Neochonetes granulifer (Owen, 1852) by Spencer (1970) from the Late Carboniferous of USA may include several species of Neochonetes. The specimens from the Falls City Limestone, the Americus Limestone Member and the Havensville Shale Member (Spencer 1970,pl. 136, figs 7-10, 13, 14) are close to Neochonetes (Zhongyingia).

Occurrence. This species has been recorded from a number of formations of Changhsingian age within Guizhou as well as the Douling Formation (Bed 13, Wuchiapingian, see Fig. 2) at the Xiaoyuanchong section in Jiahe, Hunan.

Neochonetes (Sommeriella) Archbold, 1982

Type species. Chonetes prattii Davidson, 1859, p. 116, pl. 4, figs 9-12. Sterlitamakian (Sakmarian), Western Australia.

Diagnosis. As for Neochonetes (Sommeria) Archbold (1981, p. 113).

Remarks . N. (Nongtaia) is similar to N. (Sommeriella) in its subquadrate outline, but differs from it by having distinctive coarser ornament, a distinct and normally narrow ventral sulcus and corresponding dorsal fold (Archbold 1999). N. (Zechiella) consists of a distinctive group of Late Permian species from the Zechstein Basin in Germany, England and possibly Armenia and is readily distinguished from other subgenera of Neochonetes in having obsolescent radial capillae on the shell surface and the absence of a ventral sulcus.

Distribution. Permian; Australia, South China, southern Tibet, European Russia.

Neochonetes (Sommeriella) strophomenoides (Waagen, 1884) (Fig. 5A-D)

1884 Chonetes strophomenoides Waagen, p. 628, pl. 58, figs 10a-f.

191t Chonetes strophomenoides Waagen; Frech, 191 I, p. 164, pl. 23, figs 5a, b.

1980a Neochonetes convexa Liao, p. 257, pl. 5, figs 18- 22.

1982 Neochonetes convexa Liao~ Wang et aL, p. 200, pl. 95, figs 12, 13;

1987 Neochonetes convexa Liao; Xu in Yang et al., 1987, p. 221, pl. 9, figs 13, 25, 28; pl. 10, figs 4- 10.

Material. Two ventral valves (NIGP132475, 132476) and a dorsal interior (NIGP 132477) from Bed 20 of the Changhsing Formation at the Bei fengj ing section in Zhongl iang Hill, Chongqing City and an internal mould of a ventral valve (NIGP132478) from Bed 55 of the Talung Formation at the Wenjiangsi section in Guiding, Guizhou.

Description, Shell large, subquadrate in outline,



widest at midvalve; hinge shorter or nearly equal to the greatest width; cardinal extremities nearly quadrate with an angle of about 90°; ears small, nearly flat. Ventral valve moderately strongly convex in profile; beak slightly over hinge line; sulcus arising at umbo, moderately deep and fairly broad near front margin. Surface finely costellate, numbering about 40 near front margin; costellae increasing by bifurcation and occasionally intercalation. Ventral median septum about one quarter of shell length. Dorsal alveolus deep; inner socket ridges about 2 mm long, united with cardinal process; two lateral septa low and short, about 1.5 mm long; median septum arising from anterior to the lateral septa, extending to two- thirds of shell length; internal surface finely radially papillose. Dimensions are: length, 7.7- 11.1 ram; maximnm width, 12.7-15.4 ram; thickness, about 3.6 mm.

Remarks. The present species was placed into Chonetes Fischer von Waldheim, 1830 by Waagen (1884) and Frech (1911), Neochonetes Muir- Wood, 1962 by Shen & He (1994) or listed as a species of Rugaria Cooper & Grant, 1969 by Waterhouse & Gupta (1983) and Waterhouseiella by Archbold (1983). According to Ramsbottom (1952), Imbrie (1959), Cooper & Grant (1975) and Racheboeuf(1978), the type species of Chonetes is a Devonian form with totally different structure from Neochonetes. Chonetes has a very long median septum in the ventral valve and several accessory septa in the dorsal valve. The median septum in the dorsal valve of Chonetes commences immediately under the base of the cardinal process, whereas in Neochonetes it usually arises anteriorly (Racheboeuf 1978, p. 349, fig. 1). Therefore, the present species from the Late Permian of the Salt Range, Pakistan (Waagen 1884) cannot be assigned to Chonetes. The present species possesses a moderately deep sulcus and fine costellae, and hence differs from Rugaria species. The present species is somewhat close to Waterhouseiella in outline and sulcus, but is larger and possesses finer costellae and costellate ears. Thus, Neochonetes (Sommeriella) is the most appropriate assignment

for the present species in view of its subquadrate outline, moderately deep sulcus and fine costellation. Nevertheless, the internal structure of Chonetes strophomenoides (Waagen) from the Salt Range is poorly known.

N. convexa Liao, 1980a is morphologically and chronostratigraphically consistent with the Salt Range species, therefore is herein considered to be a synonym of N. (Sommeriella) strophomenoides.

Occurrence. N. ( Sommeriella) strophomenoides (Waagen, 1884) has been also recorded from the Chhidru Formation of the Salt Range, Pakistan; the lower or middle part of the Qubuerga Formation at Shengmi in the Mt. Qomolangma (Mt. Everest) region, southern Tibet (Shen et al. 2001).

Neochonetes (Huangichonetes) subgen, nov.

Type species. Chonetes substrophomenoides Huang, 1932, p. 3, pl. 1, figs 3-7. Lopingian, South China.

Etymology. Huang-, named in honour of Huang Jiqing (Huang, T. K.), a pioneering geologist of China; Chonetes, a genus of Brachiopoda.

Diagnosis. Small Neochonetes, shell reverse trapezoidal, with small but prominent and acute ears, strongly convex visceral disc and conspicuous and moderately wide sulcus; 4 pairs of hinge spines, posterolaterally projecting at an angle of 30-40 ° to hinge line. Costellae fine, numbering 30-50 near margin. Ventral interior with very short median septum.

Remarks. N. (Huangichonetes) can be readily distinguished from N. (Sommeriella) Archbold, 1982 by its small size, maximum width at hinge with small prominent acute ears and strong convexity. N. (Nongtaia) Archbold, 1999 is also small, but differs from the present subgenus by its subquadrate outline, narrower but deeper sulcus and coarser costellae. N. (Zechiella) Archbold, 1999 differs fi'om the present subgenus



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in its obsolescent radial capillae, absence o f su lcus and also subquad ra t e out l ine . Neochone tes (Zhongyingia) differs from N. (Huangichonetes) by its larger size, less convex ventral valve and shallow, broad sulcus.

Species o f N. (Huang ichone tes ) appear somewhat transitional between Chonetinetta Ramsbottom, 1952 and Neochonetes Muir-Wood, 1962 in its strong convexity and prominent sulcus, but is closer to Neochone tes because o f its r e l a t i ve ly sha l low sulcus . The sulcus o f Chonetinella is so deep as to give the shell a bilobate appearance.

Chonetinella victoriana (Girty, 1929), C. costellata Cooper & Grant, 1975 and C. magna Cooper & Grant, 1975 from the Leonardian Bone Spring Formation of West Texas are closer to N. (Huangichonetes) in terms of their shallow sulcus, strong convexity and acute ears. Chonetinella cursothornia Xu & Grant (1994, p. 27, fig. 15.4, 6, 8-16) clearly has a shallower sulcus than any other Chonet inel la species, and is therefore also assigned to N. (Huangichonetes).

Distr ibut ion. Artinskian to Lopingian (Late Permian); South China and West Texas (USA).

Neochonetes (Huangichonetes) substrophomenoides (Huang, 1932) (Fig. 5E-M)

1932 Chonetes substrophomenoides Huang, p. 3, pl. 1, figs 3-7.

1964 Chonetinella substrophomenoides (Huang); Wang et al., p, 243, pl. 37, fig. 31.

1977 Neochonetes substrophomenoides (Huang); Yang et at., p. 331, pl. 135, fig, 20.

1978 Chonetinella substrophomenoides (Huang); Feng & Jiang, p. 242, pl. 88, fig. 1.

1979 Neochonetes sublatisinuata Zhan in Hou et al., p. 70, pl. 11, figs 5, 6, 8.

1982 Neochonetes substrophomenoides (Huang); Wang et al., p. 200, pl. 96, figs 10-11.

1982 Neochonetes substrophomenoides (Huang); Xu in Yang et al., p. 221, pl. 9, figs 14-20, 26-27.

1987 Neochonetes volitanliopsis Xu in Yang et aL, p. 222, pl. 10, fig. I.

1989 Neochonetes cf. substrophomenoides (Huang); Zhan in Li et aL, pl. 25, fig. 16.

1994 Chonetinella votitanliopsis (Xu); Xu & Grant, p. 28, figs 15. I-3, 5, 7.

Material. A ventral valve (NIGP132479), two dorsal interiors (NIGP132480, 132481), two external moulds of dorsal valves (NIGP132482, 132483) from Bed 20 at the Beifengiing section in Zhongliang Hill, Chongqing City; four internal moulds of ventral valves (NIGP 132484-132487) from Bed 55 at the Wenjiangsi section in Guiding, Guizhou and an internal mould o f a ventral valve (NIGP132488) from Bed 13 o f the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan.

Descript ion. Shell small, transverse reverse trapezoidal in outline, widest at hinge. Ventral valve strongly convex in profile; beak low and swollen, slightly beyond hinge line; cardinal extremities acute with an angle less than 70°; ears nearly fiat, well demarcated from visceral region; umbonal region highly convex with moderately deep sulcus; sulcus beginning from umbo, widening and deepening anteriorly, occupying about one-third of shell width near anterior margin; costellae very fine, usually bifurcating once or twice anteriorly, numbering about 40 near anterior margin; 4 pairs o f hinge spines, posterolaterally projecting at an angle o f about 30-40 ° to hinge line. Dimensions: length, 5.0-6.4 mm; maximum width 7.9-11.1 mm, thickness, 1.8-2.0 mm.

Ventral interior radially papillose; median

Fig. 5. A-D, Neochonetes (Sommeriella)strophomenoides (Waagen). A, ventral valve, NIGPI32475, x2.6. B, internal mould of ventral valve, NIGP132478, x3.5. C, dorsal interior, NIGPI32477, x3.4. D, ventral valve, NIGPI32476, x3.7. E-M, Neochonetes (Huangichonetes) substrophomenoides (Huang). E, internal mould of ventral valve, NIGPI32488, x6. F, internal mould of ventral valve, NIGP 132487, x5. G, internal mould of ventral valve, NIGPI32485, x4. H, dorsal interior, NIGPI32481, x4.5. I, external mould of dorsal valve, NIGPI32482, x4. J, ventral valve, NIGPI32479, x5.8. K, internal mould of ventral valve, NIGPI32484, x4.5. L, dorsal interior, NIGPI32480, x3.5. M, external mould of dorsal valve, NIGP132483, x4.5. N, ?Pygmochonetes sp. ventral valve, NIGPI32510, x4.5. O-R, Tethyochonetes soochowensis (Chao). O, external mould of dorsal valve, NIGPI32490, x5. P, external mould of dorsal valve, NIGP132491, x5. Q, external mould of dorsal valve, NIGP132492, x4. R, ventral valve, NIGPI32489, x4.



septum short, only present in apical cavity. Dorsal interior strongly radially papillose; median septum unknown; two lateral septa short, divergent anteriorly at an angle of about 30°; sockets deep with pronounced inner socket ridges and feeble short outer socket ridges.

Remarks. Specimens used by Huang (1932) to designate the present species are from two localities. The holotype and paratype (Huang 1932, pl. 1, figs 3, 4) from Tongzi, Guizhou display a transverse trapezoidal outline with acute ears, whereas specimens from Bijie, Guizhou (Huang 1932, pl. 1, figs 5, 6) have a more subquadrate outline and more subquadrate cardinal extremities, and hence may be different from the types. The specimen from the Late Permian of Nanjing, China, described as Chonetes strophomenoides Waagen by Frech (1911, p. 164, pl. 23, figs 5a, b) was assigned to the present species by Huang (1932), but is clearly larger and possesses a subquadrate outline, which is characteristic of N. (Sommeriella) strophomenoides (Waagen) (see discussion above). Specimens from the early Changhsingian of Guangdong, South China, described as Neochonetes sublatisinuata Zhan (in Hou et al. 1979) are synonymous with the present species based on their similar size, outline, sulcus and costellation (also see Xu in Yang et al. 1987). Specimens described as Chonetinella volitanliopsis by Xu in Yang et al. (1987) and Xu & Grant (1994) are most likely conspecific with the present species based on their convexity, acute ears and relatively deep sulcus.

Tethyochonetes Chen, Shi, Shen & Archbold, 2000

Type species. Waagenites soochowensis quadrata Zhan in Hou et al., 1979, p. 70, pl. 4, figs 16-19. Wuchiapingian (Late Permian); South China.

Remarks. Chen et al. (2000) indicated that species from the Lopingian of South China previously referred to Waagenites Paeckelmann, 1930 are actually different from those species from the Salt

Range, Pakistan both internally and externally; therefore Chen et al. (2000) proposed Tethyochonetes for the South Chinese "Waagenites" species. Tethyochonetes differs from Waagenites (s.s.) in its finer costellae, less distinct sulcus, longer ventral median septum, and shallow alveolus, stout median septum and lateral septa in the dorsal valve (Chen et al. 2000). Few other Permian genera are closely allied to Tethyochonetes. Fusichonetes was designated by Liao in Zhao et al. (1981) with Plicochonetes nayongensis Liao (1980a, p. 256, pl. 4, figs 7-9) as type species. This genus is poorly known in terms of its internal structures, but externally is distinguishable from Tethyochonetes by its extremely transverse outline and simple, non- bifurcate costellae (see Fig. 6Q). Waterhouseiella Archbold, 1983 with Waagenites speciosus Waterhouse & Piyasin (1970, p. 1 t 2, pl. 17, figs 9- 30; pl. 18, figs 1-21; pl. 19, figs 1-8; text-figs 6, 18, 21) from the Kungurian of Thailand as type species is similar to Tethyochonetes in size, outline, costellation and the nature of the ventral sulcus. However, Waterhouseiella appears to have more hinge spines (7-8 pairs), whereas Tethyochonetes usually possesses only 2-4 pairs of hinge spines. Sulcirugaria Waterhouse, 1983 also closely resembles Tethyochonetes in the nature of ventral sulcus and costellation, but does not possess lateral septa according to the description of Waterhouse (1983). Fanichonetes Xu & Grant (1994, p. 29) differs from Tethyochonetes by its hinge spines that are mostly inclined towards the midline.

Distribution. Lopingian (Late Permian); South China, Qinghai ofnorthwestem China and eastem Tibet.

Tethyochonetes soochowensis (Chao, 1928) (Fig. 50-R)

1928 Chonetes soochowensis Chao, p. 31, pl. 1, figs 14- 16.

1932 Chonetes soochowensis Chao; Huang, p. 5, pl. 1, figs 8a, 8b, ?9.

1964 Chonetes soochowensis Chao; Wang et al., p. 241, pl. 37, figs 20, 21.



1977 Waagenites soochowensis (Chao); Yang et al., p. 332, pl. 135, fig. 22.

1978 Waagenites soochowensis (Chao); Feng & Jiang, p. 243, pl. 88, fig. 4.

1979 Waagenites soochowensis (Chao); Zhan in Hou et al., p. 72, pl. 11, fig. 7.

1980a Waagenites cf. soochowensis (Chao); Liao, pl. 1, fig. 2.

1980a Waagenites soochowensis (Chao); Liao, pl. 5, fig. 4.

1982 Waagenites soochowensis (Chao); Wang et al., p. 197, pl. 91, figs 3-4.

1984 Waagenites soochowensis (Chao); Liao, pl. 1, fig. 10.

1987 Waagenites soochowensis (Chao); Xu in Yang et aL, pl. 8, figs 15, 16.

1990 Waagenites soochowensis (Chao); Zhu, p. 64, pl. 18, figs 1, 2.

1998 Waagenites soochowensis (Chao); Shi & Shen, p. 509, figs 4-6.

Material. A ventral valve (NIGP 132489) and an external mould of a dorsal valve (NIGP132490) from Bed 13 of the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan; an external mould of a dorsal valve (NIGP 13249 I) from Bed 45 of the Changhsing Formation and an external mould of a dorsal valve (NIGP 132492) from Bed 55 of the Talung Formation at the Wenjiangsi section in Guiding, Guizhou.

Descript ion. Shell medium sized for genus, trapezoidal in outline; widest at hinge; ears large, smooth, nearly flat, well demarcated from visceral region; cardinal extremities acute with an angle of about 60-80 °. Ventral valve moderately convex; beak broad and low, slightly beyond hinge; sulcus shallow, beginning from anterior to umbo, about one-third of shell width near anterior margin. Dorsal valve gently concave; correspondingly with weak fold. Ventral costellae numbering about 16 near umbo, increasing anteriorly by bifurcation and intercalation, numbering about 20-30 near anterior margin; four pairs of hinge spines. Dimensions: length, 4.7-6.9 rnm; maximum width, 9.8-11.3 mm; thickness, about 2 mm.

Remarks. A specimen figured as Waagenites

soochowensis (Chao) by Huang (1932, pl. 1, fig. 9) may not be a representative of Tethyochonetes in view of its costellate ears. Specimens from

South China previously described as Waagenites

barusiensis (Davidson, 1866), and renamed as Tethyochonetes chaoi and T. liaoi by Chen et al. 2000), were considered by Xu in Yang et al. (1987) to be probable juveniles of 7". soochowensis in view of small numbers ofcostellae, characteristic of early ontogeny. However, most specimens assigned to T. chaoi and T. liaoi by Chen et al. (2000) are readily distinguishable from T. soochowensis in having fewer costellae (usually 13) and being smaller and less transverse. T. soochowensis differs from T. quadrata (Zhan in Hou et al., 1979) by its relatively large size, more transverse outline and greater number of costellae. T. wongiana (Chao, 1928) resembles the present species in the number ofcostellae, but it appears less transverse, more convex and possesses more simple costellae.

A specimen from the Longtan Formation in Sichuan Province of South China figured as W. barusiensis (Davidson, 1866) by Jin et al. (1974, p. 311, pl. 164, fig. 8) is very close to the present species in view of its t ransverse (length:width=2:l) outline with acute cardinal extremities and 18 costellae on the ventral valve. However, the specimen ofJin et al. (1974) was recently designated as the holotype of T. chaoi

by Chen et al. (2000).

Occurrence . This species has been widely recorded from the Lopingian of South China and the early Changhsingian Yenduyet Formation, northwestern Vietnam.

Tethyochonetes quadrata (Zhan in Hou et al.,

1979) (Fig. 6A-E)

1979 Waagenites soochowensis quadrata Zhan in Hou et al., p. 70, pl. 4, figs 16-19.

2000 Tethyochonetes quadrata (Zhan); Chen et aL, p. 5, figs 4A-D, G.

Material. An internal mould of a ventral valve (NIGP132493), two external moulds of dorsal valves (NIGP 132494, 132495) and two external moulds of dorsal and ventral valves (NIGP 132496, 132497) from Bed 55 of the Talung Formation and an external mould of a dorsal valve (NIGP 132498)


340 SHUZHONG SHEN & NEIL W. ARCHBOLD ALCHER1NGA



from Bed 45 of the Changhsing Formation at the Wenjiangsi section in Guiding, Guizhou.

Descr ip t i on . Shell smal l , t r ansve r se ly subquadrate in outline, widest at hinge; ears acute, smooth and flat, well demarcated from visceral disc. Ventral valve strongly convex; sulcus beginning from umbo, distinct, moderately deep, about one-third of shell width near front margin. Dorsal valve correspondingly concave with distinct fold; ears flat. Surface with simple costellae, numbering about 20 near anterior margin; interspaces as wide as costellae. Ventral interior with two strong teeth; median septum short, beginning under delthyrium, extending about one-third o f shell length; parallel vascular trunks forming prominent ridges adjacent to the median septum, extending to two-thirds o f shell length. Dimensions: length, 5.0-5.8 mm; maximum width, 8.6-11.0 mm; thickness, 1.4-1.8 mm.

Remarks. This species was first proposed as a subspecies of Waagenites soochowensis by Zhan in Hou et al. (1979) and subsequently raised to species rank by Chen et al. (2000) on the basis o f its less transverse outline and more convex prof i le . T. quadra ta is c o m p a r a b l e to T. soochowens i s in details o f costat ion; both species have about 20-25 costellae on both valves. However, T. quadrata possesses a distinctly less transverse outline, more convex ventral valve and more prominent sulcus than T. soochowensis.

Specimens from the Late Permian in Qinghai Province o f northwestern China described as Waagenites convexa by Fan in Yang et al. (1962, p. 48, pl. 14, figs 11-14) have been reassigned to Tethyochonetes by Chen et al. (2000). T. convexa is closely similar to the present species in its

outline, convexity and costellation, but may be distinguishable by its deeper and more distinct sulcus.

Specimens from the Longtan Formation in Guizhou Province o f South China previously described as Waagenites barusiensis (Davidson, 1866) by Liao (1980a) are very close to T. quadrata in outline and convexity, but differ in having about 13-15 costellae. Chen etal. (2000) proposed a new species, T. liaoi, for the specimens figured by Liao (1980a). However, some other specimens from the lowest Triassic Yinkeng Formation at the Huangzhishan section in Zhejiang Province o f southeastern China assigned to T. liaoi by Chen et al. (2000) are ques t ionable . The Huangzhishan specimens (see Chen et al. 2000, fig. 5E, H-I) possess more than 25 costellae (twice as many as those o f L i ao ' s specimen f rom Guizhou) and a less convex ventral valve, and therefore may be different from the specimens o f Guizhou. These Huangzhishan specimens need to be re-investigated in terms of their hinge spine pattern because Xu & Grant (1994) proposed F a n i c h o n e t e s for the spec imens from the Changhsingian and earliest Triassic of some loca l i t ies in South China , i nc lud ing the Huangzhishan section. Fusichonetes differs from Tethyochonetes in having hinge spines inclined towards the midline.

Tethyochonetes pigmaea (Liao, 1979) (Fig. 6F, G)

1979 Fusichonetes pigmaea Liao, pl. 1, fig. 14. 1980a Plicochonetes pigmaea Liao, p. 257, pl. 4, figs 4-

6. 1980b Fusichonetes pigmaea Liao, pl. 1, figs 5, 6. 1981 Fusichonetes pigmaea Liao in Zhao et al., pl. 8,

figs 7, 8. 1982 Fusichonetes pigmaea Liao; Wang et al., p. 200,

pl. 96, figs 8, 9.

Fig. 6. A-E, Tethyochonetes quadrata (Zhan). A, external moulds of dorsal and ventral valves, NIGPI32497, x5. B, external mould of dorsal valve, NIGP132494, x5. C, latex of external mould of dorsal valve, N1GPI32495, x5. D, external mould of dorsal and ventral valves, NIGP132496, x5. E, external mould of dorsal valve, NIGPI32498, x5. F, G, Tethyochonetes pigmaea (Liao). F, ventral valve, NIGP132499, x6. G, ventral valve, N1GP132500, x5. H-N, Tethyochonetesflatus sp. nov. H, internal mould of ventral valve, paratype, NIGPI32505, x5. 1, external mould of dorsal valve, paratype, NIGPI32501, x8. J, internal mould of ventral valve, paratype, NIGPI32506, x5. K, latex of external mould of dorsal valve, paratype, NIGPI32503, x8. L, latex of external mould of ventral valve, holotype, NIGPI32511, x8. M, latex of external mould of dorsal valve, paratype, NIGPI32502, x8. N, latex of internal mould of dorsal valve, NIGPI32504, x8. O, P, Tethyochonetes sp. O, ventral valve, NIGP132508, x5. P, ventral valve, NIGPI32507, x3. Q, Fusichonetes nayongensis (Liao). External mould of dorsal valve, NIGP132509, xS.



1984 Waagenites pigmaea Liao, p. 279, pl. 1, fig. 7. 1987 Waagenites pigmaea Liao, p. 100. pl. 3, fig. 24.

Material. A ventral valve (NIGP 132499) from Bed 28 and another ventral valve NIGP 132500) from Bed 32 of the Changhsing Formation at the Bei fengj ing section in Zhongliang Hill, Chongqing City.

Description. Shell very small for genus; moderately transversely trapezoidal in outline, widest at hinge; ears small, acute and smooth, well demarcated from visceral disc. Ventral valve moderately convex; sulcus commencing anterior to umbo; widening anteriorly, moderately deep on anterior portion of shell, about one third wide near anterior margin; flanks strongly inclined. Costellae numbering about 20, simple, beginning from umbo.

Remarks. Fus&honetespigmaea was erected for specimens from the Permian-Triassic boundary beds in southeastem China by Liao (1979), but he did not give a description for the species. Subsequently, Liao (1980a, p. 257) provided a brief description of the present species, but assigned it to Plicochonetes Paeckelmann, 1930. Based on observations on more specimens from the Permian-Triass ic boundary beds o f southeastern China, Liao (1984) reassigned the present species to Waagenites. The present species is referred to Tethyochonetes by Chen et al. (2000, table 2).

Fusichonetes nayongensis (Liao, 1980a, p. 256, pl. 4, figs 7-9) differs from Tethyochonetes pigmaea by its extremely transverse outline and coarser, simpler costellae. T. flatus sp. nov., described below, differs from the present species in its gently convex profile and fewer costellae.

Tethyochonetes flatus sp. nov. (Fig. 6H-N)

Material. Holotype, an external mould of a ventral valve (NIGP 132511) from Bed 13 of the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan. Paratypes: three external moulds of dorsal valves (NIGP132501-132503), an internal mould of dorsal valve (NIGP 132504) and two internal

moulds of ventral valves (NIGP 132505, 132506) from Bed 13 of the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan.

Etymology. Flat, refer to its nearly flat ventral valve.

Diagnosis. Very small Tethyochonetes with nearly flat ventral valve; flat and smooth ears and 14-17 simple and coarse costellae; ventral sulcus absent or slightly flattened medially.

Description. Shell very small for genus, transversely subquadrate to trapezoidal in outline, widest at hinge, cardinal extremities acute; corpus cavity thin. Ventral visceral disc nearly flat; ears flat and smooth, not well differentiated from flanks of ventral disc; sulcus nearly absent or ventral valve slightly flattened medially. Dorsal valve nearly flat or slightly concave; fold absent. Dorsal alveolus small; sockets shallow; inner socket ridges coalescing with cardinal process, extending laterally at an angle less than 10 ° with hinge line; inner surface radially papillose. Ventral surface with 14-17 simple, coarse costellae; costellae beginning from umbo, not bifurcating anteriorly; crests rounded; interspaces narrower than costellae; 3-4 pairs of hinge spines, projecting posterolaterally at an angle of about 45 ° with hinge line. Dimensions: length 2.3-3.4 mm; maximum width, 4.2-7.1 mm.

Remarks. The specimen designated by Jin et al. (1974, pl. 164, fig. 8) as holotype of Tethyochonetes chaoi by Chen et al. (2000) is larger and possesses more costellae than the present species. However, some other specimens assigned to T. chaoi by Chen et al. (2000) are similar to the present species in size, outline and costellation. The specimen figured as Chonetes cf. barusiensis (Davidson, 1866) from the Lopingian of Jiangsi Province by Chao (1928, p. 29, pl. 1, fig. 18) is also very small, but possesses a more distinct sulcus and has only 13 costellae. The specimen figured as Waagenites barusiensis (Davidson, 1866) from the Changhsing Formation in Hubei Province by Yang et al. (1977, p. 332. pl. 135, fig. 4) is also close to the present species in



size and outline, but has a more prominent sulcus and more convex ventral valve. Plicochonetes pigmaea (Liao, 1980a, p. 257, pl. 4, figs 4-6) from the Changhsingian of Guizhou Province is also very small, but possesses a distinct sulcus and is more convex than the present species. Plicochonetes dissulcata Liao (1980a, p. 257, pi. 5, figs 14-17) may be also referable to Tethyochonetes, but is a nomen nudum with no formal description. This species closely resembles T. flatus in its inconspicuous sulcus, size and outline, but appears more convex in profile.

Tethyochonetes sp. (Fig. 60, P)

Material. A complete ventral valve (NIGP 132507) from Bed 20 of the Changhsing Formation at the Bei fengj ing section in Zhongl iang Hill, Chongqing City and another ventral valve (NIGP 132508) from Bed 55 of the Talung Formation at the Wenjiangsi section in Guiding, Guizhou.

Description. Shell medium for genus; somewhat semi-circular in outline; widest at hinge; strongly convex; ears small, smooth, acute and slightly attenuated; well demarcated from visceral disc; beak slightly over hinge; sulcus beginning from beak, slightly widening anteriorly, remaining narrow and shallow near anterior margin. Surface with about 30 costellae; costellae increasing by bifurcation and intercalation; four pairs of hinge spines, posterolaterally projecting.

Remarks . This species is most like Tethyochonetes wongiana (Chao, 1928, p. 28, pl. 1, fig. 17) in its size, outline and strongly convex, sulcate ventral valve. However, T. wongiana possesses a deeper sulcus and its costellae appear simpler and fewer than those of the present species. T. soochowensis (Chao) differs from the present species in its much more transverse outline and fewer costellae.

Fusichonetes Liao in Zhao et al., 1981

Type species. Plicochonetes nayongensis Liao, 1980a, p. 256, pl. 4, figs 7-9. Changhsingian;

Guizhou, South China.

Remarks. Fusichonetes was first proposed by Liao (1979), without a description. Subsequently, Liao (in Zhao et al. 1981) gave a brief diagnosis for this genus and a brief description of the type species Plicochonetes nayongensis (Liao, 1980a). The internal structures of Fusichonetes are poorly known except for a long median septum in the dorsal valve and a typical rugosochonetid cardinal process (Liao in Zhao et al. 1981, p. 52). Externally, Fusichonetes is closely similar to Plicochonetes Paeckelmann, 1930 except for the s t rongly transverse outl ine. Addit ional specimens are required to clearly define Fusichonetes . However , Plicochonetes is essentially a Carboniferous genus, whereas Fusichonetes is only known from the Lopingian of South China. Tethyochonetes Chen et al., 2000 may be congeneric with Fusichonetes in that both share similar size, sulcus, costellation and distribution.

Distribution. Lopingian (Late Permian); South China.

Fusichonetes nayongensis (Liao, 1979) (Fig. 6Q)

1979 1980a

1980b 1981

Fusichonetes nayongensis Liao, pl. I, fig. 12, 13. Plicochonetes nayongensis Liao, p. 256, pl. 4, figs 7-9. Fusichonetes nayongensis Liao, pl. 2, figs 5, 6. Fusichonetes nayongensis Liao in Zhao et al., pl. 8, fig. 13.

Material. An extemal mould of a dorsal valve (NIGP132509) from Bed 13 of the Douling Formation at the Xiaoyuanchong section in Jiahe, Hunan.

Description. Shell 3.6 mm long, 10.3 mm wide; transversely semielliptical in outline; widest at hinge; cardinal extremities alate; ears large, flat and smooth. Surface with 16 costellae; costellae simple, not bifurcating; interspaces narrower than costellae; fold absent.

Remarks. The present specimen agrees with the



holotype (Liao 1980a, pl. 4, fig. 8) in all external characters. F. nayongensis externally differs from all Tethyochonetes species from the Lopingian of South China by its extremely transverse (usually length:width<1:3) outline and simple, non-bifurcated costellae.

Family ANOPLIIDAE Muir-Wood, 1962 Subfamily CAENANOPLIINAE Archbold, 1980

Pygmochonetes Jin & Hu, 1978

Type species. Pygmochonetes jingxianensis Jin & Hu, 1978, p. t11, pl. 1, figs 28-31. Late Maokouan (Late Guadalupian); Anhui Province, southeastern China.

Remarks. Costachonetes Waterhouse, 1975 with Chonetes uralica var. pygmaea Loczy (1897, p. 64, figs 13a-f, 14a-b) from the Late Carboniferous of North China as the type species is closely similar to Pygmochonetes in its highly convex visceral region, but as Jin & Hu (1978) noted, it clearly differs in having a shorter median septum and no accessory septa in the dorsal valve. Externally, Costachonetes is somewhat triangular in outline with fewer costellae on the umbo and a narrow sulcus on the highly elevated visceral disc.

Neochonetes (Zechiella) Archbold, 1999 also lacks a sulcus on the ventral valve, but represents a group of thin-shelled Neochonetes with obsolescent radial capillae and usually has a subquadrate outline (Archbold 1999).

Distribution. Guadalupian-Lopingian (Middle- Late Permian); South China, North China and Tibet.

?Pygmochonetes sp. (Fig. 5N)

Material. A complete ventral valve (NIGP 132510) from Bed 45 of the Changhsing Formation at the Wenjiangsi section in Guiding, Guizhou.

Description. Shell 6.5 mm long, 10.4 mm wide and about 2.6 mm thick; semicircular in outline; widest at hinge. Ventral visceral disc highly elevated;

lateral slopes strongly inclined; ears gently convex; slightly attenuated; cardinal extremities acute, with a cardinal angle of about 60°; anterior margin regularly rounded; sulcus totally absent. Surface finely costellate; costellae numbering about 20 on umbo, increasing by intercalation and bifurcation anteriorly; 4 pairs of spines along hinge projecting posterolaterally at an angle about 30 ° with hinge line. Ventral interior with prominent median septum extending anteriorly to about one third of shell length.

Remarks. Although the character of the dorsal valve is unknown, the present single ventral valve most l ikely represents a species of Pygmochonetes based on the absence of a ventral sulcus, the distinct, highly elevated visceral disc and the costellation pattern. This species differs from P. jingxianensis Jin & Hu, 1978 from the Kuhfengian (Late Guadalupian) of South China in its less transverse outline, the slightly attenuated ears and bifurcated costellation. P. taniantawengica Jin & Shi, in Jin et al. (1985) from the Middle Permian at Mangkang of eastern Tibet differs from the present species in its larger and flat ears and finer costellae. Specimens described as Plicochonetes minor by Ting (1965, p. 266, pl. 1, figs 9-11) may be a Pygmochonetes in view of the absence of a ventral sulcus, but are much smaller than the present species and posses a hinge line that is shorter than the greatest width which is at midvalve. Neochonetes (Sommeriella) robustus Archbold, 1981 from the Artinskian Mingenew Formation in the Perth Basin of Western Australia also resembles the present species in the absence of ventral sulcus, but the Western Australian species has a longer median septum, two strong lateral septa and no accessory septa.

Acknowledgments

Shuzhong Shen expresses his thanks to Prof. He Xilin of the China University of Mining & Technology in Xuzhou of Jiangsu province for supervision of field studies in South China and an initial study of the specimens described in


A L C H E R I N G A L A T E P E R M I A N C H O N E T O I D E A F R O M S O U T H c H I N A 345

t h i s p a p e r . T h e s p e c i m e n s f r o m the

X i a o y u a n c h o n g section in J iahe County, Hunan

( local i ty C in this paper) were contr ibuted by

Zhang Zhipei o f Xian Geo log ica l College. Mei

Shi long provided the conodont information f rom

the upper m e m b e r o f the Dou l ing Format ion in

southern Hunan. We are also grateful to P.R.

R a c h e b o e u f and H. Campbe l l for their careful

r ev i ew and comments on the paper. This study is

suppor ted by Major Basic Research Projects o f

M S T (G2000077700) o f Peop le ' s Republ ic o f

China, CAS Hundred Talents Program and Deakin

Unive r s i ty o f Australia.

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Appendix. Species of Chonetoidea recorded from the Lopingian of South China.

(see pages 348-349)



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Chonetoidea (Brachiopoda) from the Lopingian (Late Permian) of South China

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