early carboniferous athyridid brachiopods from the qaidam basin ...

J. Paleont., 77(5), 2003, pp. 844–862 Copyright q 2003, The Paleontological Society 0022-3360/03/0077-844$03.00

EARLY CARBONIFEROUS ATHYRIDID BRACHIOPODS FROM THE QAIDAM BASIN, NORTHWEST CHINA ZHONG-QIANG CHEN,1,2 G. R. SHI,1 1

AND

LI-PEI ZHAN3

School of Ecology and Environment, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood, Victoria 3125, Australia, 2 Institute of Geology and Paleontology, Tohoku University, Aoba, Aramaki, Sendai 980-8578, Japan, ,[email protected]., and 3 Institute of Geology, Chinese Academy of Geological Sciences, Beijing, P. R. China

ABSTRACT—Four new Early Carboniferous athyridid species in three genera, including one new genus, Bruntonathyris, are described from the Qaidam Basin, northwest China: Lamellosathyris qaidamensis, Bruntonathyris amunikeensis, Bruntonathyris? heijianshanensis, and Lochengia qinghaiensis. Based on the new material and also on published information, we also reviewed the taxonomic composition and the stratigraphic and paleogeographic distributions of the three genera. As a result, Lamellosathyris is considered to be indicative of late Famennian to Viseán age, originating in late Famennian in central North America and Armenia of Russia, respectively. Later, the genus appears to have two migratory directions: one branch rapidly dispersed over Mississippi Valley, Oklahoma, Texas and New Mexico of central North America in Tournaisian; alternatively, another branch from Armenia migrated westerly to Belgium, France, Spain, Britain, Ireland, via the Moscow Basin and Ural seaway, eastward to the Tienshan Mountains and Qaidam Basin of northwest China during the Tournaisian to Viseán, and easterly along the southern shelves of the Paleo-Tethys to Iran and western Yunnan of southwestern China in Tournaisian. Both Bruntonathyris and Lochengia are restrictedly Tournaisian to Viseán in age, and probably originated in the Qaidam Basin. Later, Bruntonathyris migrated easterly to South China and Japan, and westerly to Urals, Moscow Basin, Donetsk Basin and Britain; Lochengia migrated easterly to South China and westerly to the Urals seaway and the adjoined Russian Platform (i.e., both the Moscow and Donetsk Basins).

INTRODUCTION

brachiopod faunas of the Qaidam Basin in northwest China remain poorly known due to the area’s remoteness and poor accessibility. This paper represents the first attempt to describe systematically the rich Carboniferous brachiopod faunas of this region and focuses specifically on athyridides because they are comparatively better preserved than other associated brachiopods and also contain several new species and a new genus. In addition, new material of the athyridides also provides some information on the emendation of two existing genera, Lochengia Yoh, 1929, and Lamellosathyris Jin and Fang, 1983. Adding knowledge on distributions in time and space results in new insight into the centers of origin and migratory paths of these Early Carboniferous athyridides. All described specimens were collected from the Lower Carboniferous of the Chuanshangou and Heijianshan sections in the northern Qaidam Basin, Qinghai Province, northwest China (Fig. 1) by one of the authors (ZLP) and his colleagues from the Chinese Academy of Geological Sciences from 1976 to 1983. The described and figured specimens are housed in the Department of Invertebrate Paleontology, Museum of Victoria (NMV P).

E

ARLY CARBONIFEROUS

STRATIGRAPHY

Due to the area’s very poor accessibility, the stratigraphy of the northern Qaidam Basin, especially that of the Early Carboniferous sequences, was not unveiled until recently. A well-exposed Early Carboniferous outcrop section was discovered in the early 1960s at the Chuanshangou area of the Amunike Mountain, Wulan County, Qinghai Province (Section 1, Fig. 1). Since then, this section has been studied by several geological survey teams (Lu and Zhao, 1962; Yang et al., 1962; Mu et al., 1963a–c; Wang, 1981, 1987, 1990; Shi, 1983); however, considerable discrepancies exist among their published stratigraphic schemes (Table 1). In this paper, the scheme proposed by Wang (1987, 1990) is adopted. According to this scheme, the Early Carboniferous sequence is bracketed between the underlying Ordovician and overlying Upper Carboniferous, and comprises the Chuanshangou, Chengqianggou, and Huaitoutala Formations in ascending order (Table 1). Lithologically, the Chuanshangou Formation can be further subdivided into the lower, middle, and upper members,

typified respectively by conglomerate, mixed sandstone, mudstone, and argillaceous limestone (Fig. 2). The Chengqianggou Formation consists mainly of cherty limestone, bioclastic limestone, siltstone, and black shale. The Huaitoutala Formation is defined by mixed sandstone and bioclastic limestone in the lower part and cherty limestone in the upper (Fig. 2). Marine invertebrate fossils, especially brachiopods and rugose corals, occur throughout all of the three formations and are particularly abundant in the Chuanshangou section (Fig. 2). The coral faunas have been documented by Wang (1987, 1990), while the brachiopods remain undescribed. Among the athyridides described in this paper, both Bruntonathyris? sp. and Composita ambigua (Sowerby) were collected from the lowest limestone bed in the middle member of the Chuanshangou Formation (Fig. 2). They were found in association with several other brachiopods as well as corals. The associated brachiopods include Schuchertella sp., Spinocarinifera arcuata (Hall), Syringothyris subcuspidatus (Hall), S. halli Winchell, Tylothyris sp., and Eumetria verneuiliana (Hall). In the generic level, Syringothyris was considered to be confined to the Late Devonian to early Chesterian, and Tylothyris is not younger than Osagean in Mississippi Valley (Carter, 1990). In addition, Eumetria verneuiliana has been reported from the Keokuk, Warsaw, and Salem formations of the Osagean-Meramecian in the Mississippi Valley (Kammer et al., 1990). Spinocarinifera arcuata was originally described from the Kinderhookian of Iowa (Hall, 1858), and later has been frequently applied to the lower Mississippian specimens of the Mississippi Valley (e.g., Hall and Clarke, 1895; Weller, 1914; Carter and Carter, 1970). Syringothyris halli is characteristic of the Kinderhookian in Mississippi Valley (Weller, 1914), later was found in the Tournaisian in the Kutznesk Basin (Sarycheva et al., 1963), northwest China (Jin et al., 1979; Zhang et al., 1983) and western Yunnan (Jin and Fang, 1983). Syringothyris subcuspidatus is restrictedly Osagean (Kammer et al., 1990). In view of these correlations, we regard this brachiopod fauna to be early Tournaisian in age. This age determination is reinforced by the associated coral fauna, to which Wang (1987) referred as the Kassinella-Lophophyllum densum Zone of Tournaisian age. The majority of the athyridid specimens described in this paper came from the interbedded sandstone and limestone in the upper Chengqianggou Formation (Fig. 2). They include Lamellosathyris

844

CHEN ET AL.—CARBONIFEROUS ATHYRIDID BRACHIOPODS FROM CHINA

845

FIGURE 1—Generalized locality map of the Qaidam Basin, Qinghai Province, northwest China, showing the outcrop sections. 1, highway; 2, railway; 3, town; 4, city or county town; 5, salty lake; 6, outcrop sections including the Chuanshangou section (1) and the Heijianshan section (2).

qaidamensis n. sp., Athyris? sp., Bruntonathyris amunikeensis n. gen. and sp., B.? heijianshanensis n. gen. and sp., and Lochengia qinghaiensis n. sp. Other co-existing brachiopods are Schellwienella sp., Buxtonia cf. dengisi Nalivkin, Tylothyris laminosa (McCoy), Spirifer (Grandispirifer) mylkensis Yang, and Cyrtina hibernica Brunton. Of these, Tylothyris laminosa is known from the late Tournaisian in Belgium and from the late Tournaisian to middle Viseán in Ireland (Brunton, 1984). Cyrtina hibernica has been

TABLE 1—Changes of stratigraphic subdivision of the Lower Carboniferous at the northern Qaidam Basin.

described from the mid-Viseán rock in northwest Ireland (Brunton, 1984). Buxtonia cf. dengisi has been reported from the Tournaisian in Urals (Nalivkin, 1975). Spirifer (Grandispirifer) mylkensis is characteristic of the late Tournaisian to early Viseán in northwest China (Yang, 1964, 1990; Jin et al., 1979; Zhang et al., 1983). The associated coral Ekvasophyllum heijianshanensis-Cystocyathoclisia sinensis Assemblage has also been constrained as the late Tournaisian (Wang, 1987, 1990). As such, the present fauna can be referred to the late Tournaisian by means of faunal correlation and its stratigraphic position above the early Tournaisian fauna. In addition, Lamellosathyris qaidamensis n. sp., Bruntonathyris amunikeensis n. gen. and sp., and Lochengia qinghaiensis n. sp. incorporating with Echinoconchella elegans (McCoy) and Fluctuaria undata (Defrance) are present in the basal Huaitoulata Formation. Although these described athyridid species persist from the underlying Chengqianggou Formation of the Tournaisian, Echinoconchella elegans and Fluctuaria undata are both characteristic of the Viseán in West Europe, Urals-Russian Platform, Central Asia, northwest China, and South China (i.e., Sarycheva and Sokolskaya, 1952; Muir-Wood and Cooper, 1960; Litvinovitch, 1962; Nalivkin, 1975; Galitzkaja, 1977; Yang, 1978, 1990). Accordingly, the brachiopod fauna can be safely assigned to the Viseán, likely early Viseán due to its stratigraphic position just above the Tournaisian Chengqianggou Formation. Based on the brachiopod data, this conclusion agrees well with the age constraint derived from the associated coral species of Neoclisiophyllum, Cravenia, and Rylstonia (Wang, 1987).

846

JOURNAL OF PALEONTOLOGY, V. 77, NO. 5, 2003

FIGURE 2—Lithological features of the Lower Carboniferous at the Chuanshangou and Heijianshan sections, showing the fossil horizons. 1, conglomerate; 2, sandstone; 3, siltstone; 4, sandy mudstone; 5, black shale; 6, cherty limestone; 7, limestone; 8, argillaceous limestone; 9, bio-clastic limestone; 10, oolitic limestone; 11, coral; 12, brachiopod.

Outside the Haoluge area in the Qaidam Basin, a comparable Early Carboniferous sequence also is well exposed at the Heijianshan section of the Taerding area, about 200 km west of Germu City, Qinghai Province (Section 2, Fig. 1). Here, the correlative of the Huaitoutala Formation is known as the Xihansitegou

Formation, from which three athyridid species, namely Bruntonathyris? sp., B.? heijianshanensis n. sp., and Composita ambigua (Sowerby), are described from a bioclastic limestone, along with Gigantoproductus superba (Sarycheva), Striatifera striata (Fischer de Waldheim), Latiproductus latissima (Sowerby), and

Lamellosathyris sp. L. qaidamensis n. sp. Arize, France Qaidam Basin, China Viseán late Tournaisian to early Viseán Lamellosathyris sp. L. qaidamensis n. sp.

Revision Tectonic unit

Belgium Yorkshire, England Belgium Belgium Belgium Missouri, USA Missouri, USA Missouri, USA Mississippi Valley Oklahoma New Mexico Moscow Basin Urals, Russia Armenia, Russia Elburz, Iran Missouri, USA Missouri, USA Texas, USA Tienshan Mts, China west Yunnan, China Yorkshire, England Belgium Yorkshire, England Fermanagh, Ireland Sierra Morena, Spain

Age

late Tournaisian late Tournaisian late Tournaisian late Tournaisian late Tournaisian Tournaisian early Tournaisian Tournaisian Tournaisian late Tournaisian Tournaisian Tournaisian Tournaisian latest Famennian Tournaisian late Famennian late Tournaisian Tournaisian Tournaisian to early Viseán late Tournaisian Tournaisian late Tournaisian late Tournaisian middle Viseán Viseán

Le´veille´, 1835 Davidson, 1858 Koninck, 1887 Frech, 1916 Wolfart, 1965 Hall and Clarke, 1893, 1895 Weller, 1914 Williams, 1943 Boucot et al., 1965 Carter, 1999 Carter, personal comm. Janischewsky, 1954 Nalivkin, 1937 Abramyan, 1957 Gaetani, 1968 Weller, 1914, 1931 Williams, 1943 Carter, 1967 Yang, 1964 Jin and Fang, 1983 Phillips, 1836 Brunton, 1980, figs. 16–17 Brunton, 1980, fig. 15 Brunton, 1984 Martinez-Chacoń and Legrand-Blain, 1992 Legrand-Blain et al., 1996 This paper

Author Species

Spirifer lamellosus Le´veille´ Athyris lamellosa (Le´veille´) A.? lamellosa (Le´veille´) A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) Lamellosathyris lamellosus (Le´veille´) L. lamellosa (Le´veille´) Athyris? forschi Janischewsky A. lamellosa (Le´veille´) A. lamellosa (Le´veille´) A.? lamellosus (Le´veille´) A. hannibalensis (Swallow) A. hannibalensis (Swallow) A. bradyensis Carter Cleiothyridina lamellosa (Le´veille´) Lamellosathyris lamellosa (Le´veille´) Spirifer squamosa Phillips Actinoconchus lamellosus (Le´veille´) A. lamellosus (Le´veille´) A. lamellosus (Le´veille´) Lamellosathyris sp.

TABLE 2—Specific composition with revisions, stratigraphic and geographic distributions of Lamellosathyris Jin and Fang, 1983.

Lamellosathyris lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. qaidamensis n. sp. L. qaidamensis n. sp. L. qaidamensis n. sp. L. qaidamensis n. sp. L. qaidamensis n. sp. L. qaidamensis n. sp. L. forschi (Janischewsky) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. qaidamensis n. sp. L. hannibalensis (Swallow) L. hannibalensis (Swallow) L. bradyensis (Carter) L. qaidamensis n. sp. ?L. lamellosa (Le´veille´) L. lamellosus (Le´veille´) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) L. lamellosa (Le´veille´) Lamellosathyris sp.


847

FIGURE 3—Paleogeographic distributions and their original centers and migratory paths of Lamellosathyris Jin and Fang, 1983, Bruntonathyris n. gen. and Lochengia Yoh, 1929. Kaz., Kazakhstan Plate; Qai., Qaidam Basin; T., Tarim Basin; Sc., South China Block; Ic., Indochina Block. 1, Missouri; 2, Oklahoma; 3, Texas; 4, New Mexico; 5, Armenia; 6, Tournai of Belgium; 7, Arize of France; 8, Sierra Morena of Spain; 9, Yorkshire of England; 10, Fermanagh of Ireland; 11, Moscow Basin; 12, Urals; 13, Tienshan Mountains of northwest China; 14, Qaidam Basin of northwest China; 15, Hunan and Guangxi of South China; 16, Akiyoshi of Japan; 17, Donetsk Basin of Russia; 18, Elburz Mountains of Iran; 19, western Yunnan of southwestern China. The fine bold lines with arrows demonstrate the migratory directions and paths of Lamellosathyris; the dash lines with arrows exhibit the inferred migratory directions and paths of Bruntonathyris; the coarse bold lines and arrows show the migratory directions and paths of Lochengia. The paleogeographic map is modified from Scotese and McKerrow (1990) and Metcalfe (1996).

Echinoconchella sp. (Wang, 1990). Most of these brachiopod species are typical of the late Viseán to Serpukhovian in Asia, West Europe, and Urals-Russian Platform (Sarycheva and Sokolskaya, 1952; Muir-Wood and Cooper, 1960; Yang, 1964, 1990). Associated corals Palaeoslilia planum Gorsky, Dibunophyllum vaughani Salee, D. bipartitum (McCoy), Kueichouphyllum sinense Yu, Lonsdaleia stelcki (Nelson), and Hexaphyllum qinghaiensis Wang have been referred to the Viseán by Wang (1987, 1990). STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTIONS OF LAMELLOSATHYRIS, BRUNTONATHYRIS, AND LOCHENGIA

To date, Lamellosathyris, as revised on Table 2, encompasses five species and two undetermined species worldwide, all of which are restricted to the late Famennian to Viseán of the Early Carboniferous. Geographically, Lamellosathyris has been reported from Tournai of Belgium; Sierra Morena of Spain; Arize of France, Yorkshire of England; Fermanagh of Ireland; Moscow Basin, Urals, and Armenia of Russia; Elburz Mountains of Iran; Tienshan Mountains, western Yunnan, and Qaidam Basin of China; and Missouri, Oklahoma, Texas, and New Mexico of the US (Table 2). In view of the Early Carboniferous paleogeographic reconstruction provided by Scotese and McKerrow (1990) and Metcalfe (1996), these tectonic units (see Table 2) were situated near the tropical and subtropical zones during the Early Carboniferous (Fig. 3). This may imply that Lamellosathyris inhabited warm to warm-temperate climatic conditions. Also evident from the distribution data (Table 2) the genus first occurred in the late Famennian in Missouri of the US (Williams, 1943) and Armenia of Russia (Abramyan, 1957), respectively. Accordingly, Lamellosathyris appears to have two centers of origin, which were isolated by the Phoibic Ocean (sensu Scotese and McKerrow, 1990) (Fig. 3). Of these, in North America the oldest Lamellosathyris species is L. hannibalensis (Sowallow) from the late Famennian Louisiana Limestone of Missouri (Williams, 1943). Another Famennian species Athyris transversa

Qaidam Basin, China early Tourn. to Viseán

Revision Tectonic unit

Yorkshire, England Yorkshire, England Hunan, South China Moscow Basin, Russia Urals, Russia Urals, Russia Akiyoshi, Japan Donetsk Basin, Russia Moscow Basin, Russia Urals, Russia Moscow Basin, Russia Qaidam Basin, China Qaidam Basin, China early Viseán early Viseán late Viseán Viseán Viseán Viseán Viseán Viseán Viseán Viseán Viseán late Tourn. to early Viseán late Tourn. to Viseán

Age Author

This paper

Type species.Spirifer lamellosus Le´veille´, 1835, p. 39, pl. 2, figs. 21–23, with neotype selected by Brunton (1980, p. 225, fig. 16). Revised diagnosis.Medium to large Athyridinae, suboval to subpentagonal in outline, variously dorsi-biconvex, with conspicuous fold and sulcus, rounded parasulcate anterior commissure; lamellae broad, with variably developed radial striations. Teeth strong; dental plates well-developed (Fig. 4). Cardinal plate thick,

Bruntonathyris? sp.

Order ATHYRIDIDA Boucot, Johnson, and Staton, 1964 Suborder ATHYRIDIDINA Boucot, Johnson, and Staton, 1964 Superfamily ATHYRIDIDAE Davidson, 1881 Family ATHYRIDIDAE Davidson, 1881 Subfamily ATHYRIDINAE Davidson, 1881 Genus LAMELLOSATHYRIS Jin and Fang, 1983

Davidson, 1859 (pars) Brunton, 1980 Ozaki, 1939 Sarycheva and Sokolskaya, 1952 (pars) Janischewsky, 1954 Mo¨ller, 1862 Yanagida, 1962 Poletaev, 1975 Grunt, 1968 Nalivkin, 1975 Grunt, 1980 This paper This paper

The classification of the Athyridida follows Alvarez et al. (1998). All morphological terms are in current use, as in Brunton et al. (1996) and Williams and Brunton (1997).

Species

SYSTEMATIC PALEONTOLOGY

Athyris expansa (Phillips) Actinoconchus expansus patulus Brunton Athyris expansa (Phillips) Athyris expansa (Phillips) Athyris expansa (Phillips) Athyris expansa (Phillips) Cleiothyridina expansa (Phillips) Actinoconchus? expansus (Phillips) Actinoconchus expansus (Phillips) Actinoconchus expansus (Phillips) Leiothycridina expansa (Phillips) Bruntonathyris amunikeensis n. gen. and sp. Bruntonathyris? heijianshanensis n. gen. and sp.

(Stainbrook) from the Percha Shale of New Mexico also possesses the requisite frilly lamellar ornament, is slightly older than L. hannibalensis (see also Stainbrook, 1947, p. 237; Cooper and Dutro, 1982, p. 92), and thus is the possible ancestor of Lamellosathyris. During the Tournaisian, the genus appears to have rapidly dispersed over the Mississippi Valley, Oklahoma, Texas, and New Mexico of central North America. In Armenia, Lamellosathyris raised in the Etonungian (latest Famennian), later migrated northwestward to Belgium, from there it then appears to have spread in two directions: a westerly branch to France, Spain, Britain and Ireland in late Tournaisian to Viseán and a northeastern branch to the Moscow Basin through the Urals and Tienshan Mountains eastward to the Qaidam Basin of northwest China in late Tournaisian to early Viseán (Fig. 3). This inferred eastward migration route basically followed the northern shelves of the Paleo-Tethys (in sense of Metcalfe, 1996). Alternatively, migration of Lamellosathyris along the southern shelves of the Paleo-Tethys also seems to have taken place during the Tournaisian, as indicated by the occurrence of this genus in Iran and western Yunnan in southwest China (Fig. 3). Both Bruntonathyris and Lochengia demonstrate narrower stratigraphic and paleogeographic ranges than Lamellosathyris; both occur only in the Tournaisian to Viseán and are confined to the western, northern, and eastern parts of the Paleo-Tethys (Tables 3 and 4; Fig. 3). Despite being restricted to the TournaisianViseán, more specific age determination is not possible at present for most species of Bruntonathyris, which hampers the understanding of its center of origin and its migration pathways. However, according to the known data (Table 3), the confirmed oldest species could be Bruntonathyris amunikeensis n. gen. and sp. from the late Tournaisian in the Qaidam Basin, northwest China, suggesting that the Qaidam Basin was the possible center of origin for the genus in the Tournaisian. Later, the genus migrated easterly to South China and Japan in Viseán, and westerly to Urals, Moscow Basin, Donetsk Basin, and Britain in early Viseán. In contrast, the migration route for Lochengia could be inferred on the basis of the first stratigraphic occurrences of its constituent species. Apart from L. qinghaiensis n. sp., which occurs in the early Tournaisian to early Viseán of the Qaidam Basin, all the other remaining species of this genus (Table 4) are restricted to the Viseán, largely to the late Viseán. It is therefore reasonable to suggest that Lochengia originated at the Qaidam Basin and migrated easterly to South China and westerly to the Urals seaway and the adjoined Russian Platform (i.e., both the Moscow and Donetsk basins) (Fig. 3).

Bruntonathyris patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) B. patula (Brunton) Bruntonathyris amunikeensis n. gen. and sp. Bruntonathyris? heijianshanensis n. gen. and sp. Bruntonathyris? sp.


TABLE 3—Specific composition with revisions, stratigraphic and geographic distributions of Bruntonathyris new genus.

848

Guangxi, South China Guangxi, South China Guangxi, South China Guangxi, South China Guangxi, South China Urals, Russia Donetsk Basin, Russia Donetsk Basin, Russia Urals, Russia Moscow Basin, Russia Moscow Basin, Russia Qaidam Basin, China

Revision Tectonic unit Age

late Viseán late Viseán late Viseán late Viseán late Viseán Viseán Viseán Viseán Viseán Viseán Viseán late Tourn. to early Viseán

Author

Grabau, 1931b Ching and Yang in Yang et al., 1977 Jin, 1983 Ching and Yang in Yang et al., 1977 Jin, 1983 Mo¨ller, 1862 Rotai, 1941 Poletaev, 1975 Nalivkin, 1975 Grunt, 1980 Grunt, 1980 This paper

Species

Cryptospirifer lochengensis Grabau Lochengia lochengensis (Grabau) Lochengia lochengensis (Grabau) Lochengia enteletiformis (Grabau) Lochengia enteletiformis (Grabau) Athyris variabilis Mo¨ller A. (Actinoconcha) variabilis (Mo¨ller) Actinoconcha? variabilis (Mo¨ller) Actinoconcha variablis (Mo¨ller) Flexathyris variabilis (Mo¨ller) Flexathyris prokofjevi (Grunt) Lochengia qinghaiensis n. sp.

TABLE 4—Specific composition with revisions, stratigraphic and geographic distributions of Lochengia Yoh, 1929.

849

Lochengia lochengensis (Grabau) L. lochengensis (Grabau) L. lochengensis (Grabau) L. enteletiformis (Grabau) L. enteletiformis (Grabau) Lochengia variabilis (Mo¨ller) L. variabilis (Mo¨ller) L. variabilis (Mo¨ller) L. variabilis (Mo¨ller) L. variabilis (Mo¨ller) Lochengia prokofjevi (Grunt) Lochengia qinghaiensis n. sp.


FIGURE 4—Ventral interiors of Lamellosathyris qaidamensis n. sp. (based on latex replica of ventral interior of specimen NMV P309558) showing well-developed dental plates. Scale bar represents 30 mm. t, teeth; dp, dental plate.

subtriangular to subtrapezoidal in outline, perforate apically, confined laterally by high inner socket ridges; cardinal flanges well or less developed, projecting posteroventrally; sockets broad, distinct; median myophragm low; spiralia similar to that of Athyris. Other species.See Table 2. Discussion.The above diagnosis is amended substantially from the original definition of the genus by Jin and Fang (1983, p. 147). The amendment is necessitated because Jin and Fang (1983) defined the genus based on the supposed Spirifer lamellosus Le´veille´ from western Yunnan instead of the type material of the species from the Tournai region, Belgium. The western Yunnan material is apparently not comparable with the neotype of Spirifer lamellosus selected by Brunton (1980, fig. 16). As a result, Brunton (1984) disregarded Lamellosathyris as a valid genus. However, the type species Spirifer lamellosus has been validated and re-described by Brunton (1980), Lamellosathyris therefore is also valid in accordance with the International Commission on Zoological Nomenclature (4th edition, 1999, Article 70). Subsequently, we agree with Alvarez et al. (1998) and Carter (1999) that Lamellosathyris is a distinct genus, but it is essential to amend the generic concept on the basis of the valid type specimens of the type species. The revised diagnosis given above is elaborated from Gaetani (1968) and Brunton (1980, 1984), who studied the type species, L. lamellosus (Le´veille´). Lamellosathyris is allied to several athyridid genera, as reflected by the fact that its type species, Spirifer lamellosus Le´veille´, has hitherto been assigned to several genera. Of these, Athyris is perhaps the closest. But, a re-study by Alvarez et al. (1996) of Terebratula concentrica Buch, 1834, type species of Athyris, demonstrates that this genus is sculptured with dense growth lines, unlike the lamellae of Lamellosathyris. Cleiothyridina is also related in general aspects to Lamellosathyris, from which Cleiothyridina is distinguished by possessing very close growth lamellae that project anteriorly and anterolaterally as flat solid spines, as illustrated by Alvarez (1999) and Alvarez and Brunton (2001). Prominent lamellose flanges, on the other hand, also characterize Actinoconchus, but in this genus the lamellose flanges are usually narrower and more strongly radially striated than in Lamellosathyris. Moreover, like Cleiothyridina, Actinoconchus also lacks a well-developed fold and sulcus. LAMELLOSATHYRIS QAIDAMENSIS new species Figures 4, 5.1–5.6, 5.8–5.9, 5.11–5.12, 5.16–5.17, 6 Athyris lamellosa (LE´VEILLE´, 1835). HALL AND CLARKE, 1893, p. 90, pl. 46, figs. 16–20; HALL AND CLARKE, 1895, pl. 46, figs. 16–20; WELLER,

850


TABLE 5—Shell dimensions (mm) of Lamellosathyris qaidamensis new species. Specimens

Width

Length

Thickness

NMV P309558* NMV P309559

54.0 42.0

38.8 34.7

19.5 20.5

* holotype.

1914, p. 465, pl. 78, figs. 1–5, 15–20; WILLIAMS, 1943, p. 91, pl. 8, fig. 25; YANG, 1964, p. 144–145, pl. 22, figs. 1–2; BOUCOT, JOHNSON, AND STATON, 1965, fig. 537, fig. 4b–d. Athyris? lamellosus (LE´VEILLE´, 1835). GAETANI, 1968, p. 708–711, pl. 51, figs. 1–3. Lamellosathyris lamellosa (LE´VEILLE´, 1835). CARTER, 1999, p. 121, figs. 13A–H, 14.

Diagnosis.Large Lamellosathyris (see measurements below), with distinct dorsibiconvex profile; about 10–11 broad growth lamellae on each valve. Description.Transversely oval to subpentagonal in outline; hingeline narrower than maximum width at midvalve; cardinal extremities rounded. Ventral beak small, incurved, slightly overhanging hingeline; foramen round; median sulcus originating at umbo, broadening and deepening anteriorly, forming a broad, deep lingual extension at anterior margin. Dorsal umbo highly convex, lateral slopes steep; median fold commencing anterior to umbo, indistinct on umbonal region, highly raised at anterior margin. Growth lamellae broad, each about 5–6 mm wide at midvalve and bearing about 10 fine concentric growth lines at midvalve. Teeth stout; dental plates thick, slightly converging ventrally (Fig. 4), extending nearly to one-third valve length; shell thickening posteriorly. Cardinal plate thick, well developed, subtriangular to subtrapezoidal in outline, confined laterally by high inner socket ridges; dorsal foramen small; inner hinge plates united, concave ventrally (Fig. 6); outer hinge plates narrow; crural bases strong; medium myophragm low, extending anteriorly to almost half length of dorsal valve. Etymology.Named for the Qaidam Basin. Type.NMV P309558 is designated herein as holotype. From the Chuanshangou section of the Amunike Mountain, Qinghai Province, northwest China. Measurements (in mm).See Table 5. Other material examined.NMV P309559 and seven broken shells. Occurrence.Tournaisian of Missouri, Oklahoma, and New Mexico, the US and the Elburz Mountains of Iran; Tournaisian to early Viseán of the Tienshan Mountains and the Qaidam Basin, northwest China. Discussion.Jin and Fang (1983, p. 147–149, pl. 2, figs. 6– 11) assigned some Lower Carboniferous specimens of western Yunnan to L. lamellosa (Le´veille´). However, as noted by the authors, the western Yunnan material embraces smooth lamellae and lacks dental plates, hence in sharp contrast to the radially striated

lamella and the well-developed dental plates in the topotype material of L. lamellosa (Le´veille´), as described and figured by Brunton (1980, 1984). These differences suggest that the western Yunnan specimens are convertible to a different species or even a separate genus (Jin and Fang, 1983, p. 147). Specimens from the Tienshan Mountains of northwest China (Yang, 1964), and Elburz Mountains of Iran (Gaetani, 1968) could be conspecific with the described material because they share almost all of the observed features. When compared with the typical L. lamellosa re-diagnosed and illustrated by Brunton (1980, 1984), the new species clearly differs from the type species in being significantly larger; of these specimens of L. lamellosa studied by Brunton (1980, 1984) from Belgium, Britain and Ireland, the largest is about 25 mm in width (Brunton, 2002, personal commun.), whereas the Qaidam specimens are usually more than 42 mm in width. In addition, the cardinal plate of the new species is not clearly perforated apically, but the foramen is very distinct in the typical L. lamellosa (see also Brunton, 1980, fig. 17). It is also noteworthy that the European specimens (Brunton, 1980, 1984) possess a slightly greater number of lamellae and a proportionately narrower but deeper lingual extension at the ventral anterior margin. Brunton (1980, p. 225) also stated L. lamellosa may have a shallow dorsal sulcus, but this character is never present in our specimens. Comparable materials from the central North America also have long been assigned to the lamellosa species (e.g., Hall and Clarke, 1893, 1895; Weller, 1914; Williams, 1943; Carter, 1999). However, like the new species, the adult specimens are often much larger than the true L. lamellosa. Although the smaller specimens of ‘‘L. lamellosa’’ from the Mississippi Valley (see Weller, 1914, p. 467, pl. 78, figs. 10–14) resemble the Brunton’s neotype in size and overall appearance, they are much more circular in outline and possess 7–9 lamellae, whereas the true L. lamellosa is proportionately much more transverse and has more than ten lamellae. As consequence, the American specimens apparently are not suitable for L. lamellose. Alternatively, the mature individuals of the American specimens agree well with the new species. Lamellosathyris sp. from the Lower Carboniferous of the Los Santos de Maimona Basin, Sierra Morena, Spain (Martıńez-Chacoń and Legrand-Blain, 1992, p. 119–120, pl. 6, figs. 1–3) can be differentiated from L. qaidamensis by its smaller size, narrower, and denser lamellae. Legrand-Blain et al. (1996, p. 197, pl. 28, figs. 17–18) also described several specimens as Lamellosathyris sp. from the Viseán of the Arize Massif, France, but these French specimens, showing relatively denser lamellae, are too poorly preserved to allow detailed comparison. L. bradyensis (Carter, 1967, p. 336–340, pl. 31, figs. 1–11) from the Mississippian of central Texas may be distinguished from L. qaidamensis by its smaller size, less pronounced fold and sulcus, and presence of a groove on the dorsal fold. Genus ATHYRIS McCoy, 1844 Type species.Terebratula concentrica VON BUCH, 1834, p. 103–104. By subsequent designation of King (1850).

→ FIGURE 5—1–6, 8–9, 11–12, 16–17, Lamellosathyris qaidamensis n. sp. 1–5, 16–17, Holotype, NMV P309558, a specimen in dorsal, ventral, lateral, posterior and anterior views, 31.3, 31.3, 31.3, 31.5, 31.0, latex replica of ventral interior, 31.4, close-up of ornaments, showing lamellae and growth lines, 33.0, all from the Chengqianggou Formation of the Chuanshangou section; 6, 8–9, 11–12, NMV P309559, a complete specimen in dorsal, ventral, lateral, posterior and anterior views, from the Chuanshangou Formation of the Chuanshangou section, all 31.2. 7, 10, 13–15, Athyris? sp. NMV P309560, a complete specimen in dorsal, ventral, posterior, anterior and lateral views, from the Chengqianggou Formation of the Chuanshangou section, all 31.3.


851

852


FIGURE 6—Serial sections of Lamellosathyris qidamensis n. sp. (based on specimen NMV P309559). The numbers are distance from the ventral beak, the direction of the sections is with dorsal valve downward (hereafter same). dp, dental plates; hp, hinge plate.

Diagnosis.Equidimensional to slightly transverse shells, biconvex to dorsibiconvex profile, variably developed fold and sulcus; ornamentation consisting of regular, fine, slightly lamellose growth lines; dental plates long, thin, subparallel and slightly concave; cardinal plate triangular to subtrapezoidal, flat or slightly concave, and apically perforated; cardinal flanges poorly developed or absent; jugum projecting anteriorly as jugal saddle and posteroventrally as thin jugal stem; accessory jugal lamellae terminating slightly posterior to lateral branches of jugum (Alvarez et al., 1996). ATHYRIS? sp. Figure 5.7, 5.10, 5.13–5.15 Description.Shell about 32.5 mm long, 35.0 mm wide and 18.5 mm thick; equidicircular to subpentagonal in outline; strongly dorsibiconvex in profile; anterior commissure strongly parasulcate; anterior margins strongly bent ventrally. Ventral valve moderately convex; foramen round, large; beak pointed, strongly

incurved, overhanging hingeline; umbonal region gently convex; median sulcus originating anterior to umbo, deepening anteriorly, forming V-shaped anterior extension. Dorsal median fold indistinct when commencing anterior to umbo, abruptly raised anteriorly, forming a prominent fold near anterior margin. Growth lines fine, dense, regularly spaced; lamellae poorly developed, only visible near anterior margins. Interiors not observed. Material examined.NMV P309560. Occurrence.Late Tournaisian of the Qaidam Basin, northwest China. Discussion.Potentially, our specimen may belong to a distinct new species, characterized by its strongly dorsibiconvex profile, a prominent ventral anterior extension, and ventrally bent anterior margins. However, erection of a full new species is hampered because of insufficient material and lack of knowledge on its internal structures, the latter rendering even its generic status uncertain. Nevertheless, our specimen is close to the neotype of Athyris concentrica (see Alvarez et al., 1996, pl. 1, fig. 1) in


853

TABLE 6—Shell dimensions (mm) of Bruntonathyris amunikeensis new genus and species. Specimens NMV NMV NMV NMV NMV NMV

P309561 P309562 P309563* P309564 P309565 P309566

Width

Length

Thickness

40.0 38.6 36.0 25.0 24.8 33.0

27.5 24.5 24.0 16.5 16.0 22.4

8.9 11.0 9.6 6.5 6.9 7.5

* holotype.

FIGURE 7—Dorsal cardinalia of Bruntonathyris amunikeensis n. gen. and sp. (based on serial sections of specimen NMV P309564). cb, crural base; f, dorsal foramen; hp, united inner hinge plates; ohp 1 isr, junction of outer hinge plate and inner socket ridge; osr, outer socket ridge; so, socket; my, myophragm; ms, muscle scars. Scale bar represents 30 mm.

outline and size, but the latter has a smaller ventral foramen, a less conspicuous median fold coupled by a shallower median depression on the ventral valve. On the other hand, the presence of a strongly dorsibiconvex profile and a distinct sulcus and fold of the Qaidam specimen recalls the Middle Devonian Bruntonites mellingeni Struve (1992, p. 560–561; see also Alvarez et al., 1996, pl. 5, figs. 25–27; pl. 6, figs. 28–29), type species of the genus, but in the latter the dorsal median fold originates anterior to the beak, the dorsal umbo is more strongly convex, and the lamellae are more pronounced and regular. Subfamily CLEIOTHYRIDININAE Alvarez, Rong, and Boucot, 1998 Genus BRUNTONATHYRIS new genus Type species.Bruntonathyris amunikeensis new genus and species. Included species.See Table 3. Diagnosis.Transverse Cleiothyridininae, with variously dorsibiconvex to plano-convex profile, fine growth lines; valves extremely thin; exfoliated surfaces display a delicate radial pattern. Description.Varied sizes; much wider than long; anterior commissure rectimarginate to weakly parasulcate; sulcus and fold variably developed; growth lines fine, rarely preserved, occasionally weakly lamellose anteriorly; striae dense, numerous, distinct when outer layer removed. Dental plates low, partially buried by secondary shell thickening. Crural bases strong, divergent; inner hinge plate conjunct, strongly concave ventrally (Fig. 7); myophragms broad, short; accessory ridges low, numerous. Etymology.After Dr. C. H. C. Brunton of the British Natural History Museum, London. Discussion.The new genus is proposed to represent a group of species distinguished by a transverse outline, a dorsibiconvex to plano-convex profile, weakly developed lamellae but dense radial striations on exfoliated shells, and a lack of a dorsal median sulcus. Leiothyrcridina Grunt, 1980, is the closest ally to the new genus, from which the Russian genus is differentiated by possessing a more circular outline, an equibiconvex profile, denser and more conspicuous spinose lamellae, thinner crural bases bounding flattened conjunct inner hinge plates, a relatively higher

and thicker myophragm, and lacking a ventral median sulcus or depression. The presence of long delicate flanges of shell from rugae or strong growth lines on both valves has served as a key criterion to distinguish Actinoconchus McCoy, 1844, from other allies (Brunton, 1980, 1984). These distinctive ornaments are invisible on Bruntonathyris. Furthermore, the equibiconvex profile and a shallow but distinct median groove on both valves of Actinoconchus are also markedly different from the dorsibiconvex to planoconvex profile and usually well-developed sulcus and fold of the new genus. Cleiothyridina Buckman, 1906, is defined by a circular outline, an equibiconvex profile, conspicuous spinose lamellae, and absence of sulcus and fold, and thus can be easily separated from Bruntonathyris. The pronounced radial striae of Bruntonathyris also recall the radial ornament of Atrythyris Struve, 1965, but the radial lines of Atrythyris are present on the external surfaces, forming reticulation with regular concentric lamellae, unlike the striae of Bruntonathyris, which are usually visible only when the outer shell layer is removed. Athyris is also fairly similar to Bruntonathyris in many respects, but is distinct due to the former having a proportionately more elongate outline, a more pronounced sulcus and fold that usually form a strongly parasulcate anterior commissure, and a pair of long, high, and subparallel dental plates. BRUNTONATHYRIS AMUNIKEENSIS new species Figures 7, 8, 9 Diagnosis.Typical Bruntonathyris, but distinct by its transverse outline and strongly dorsiconvex profile; striae conspicuous when outer shell layer removed. Description.Medium-sized to large for genus (see measurements); anterior commissure rectimarginate in juveniles, weakly parasulcate at maturity. Ventral beak small, strongly incurved, suspending above dorsal beak; umbo moderately convex; lateral slopes gentle; median sulcus absent or served by a depression near anterior margin. Dorsal valve slightly flattened to moderately, longitudinally convex to form a low median fold; beak strongly incurved under ventral delthyrium. Growth lines occasionally lamellose anteriorly; anterior margins occasionally weakly plicate in some specimens. Dental plates thin, high, diverging at angles varying between 140 degrees and 150 degrees, extending to one-third shell length. Dental sockets deep with distinct socket ridges; dorsal foramen large, rounded behind triangular cardinal plate; crural bases strong, divergent at about 40 degrees; inner hinge plates conjunct, strongly concave ventrally, outer hinge plates narrow, slightly flattened; muscle scars striate; myophragm broad, short; numerous accessory ridges low, short, extending for a half length of muscle scar areas (Fig. 7). Etymology.Named for the Amunike Mountain, where the Chuanshangou section is situated. Type.NMV P309563 is selected as holotype. From the Chuanshangou section. Measurements.See Table 6.

854



855

FIGURE 9—Serial sections of Bruntonathyris amunikeensis n. gen. and sp. (based on specimen NMV P309564). The numbers are distance from the ventral beak; the direction of the sections is with ventral valve downward (hereafter same). dp, dental plates; hp, hinge plate.

Other material examined.NMV P309561–562, NMV P309564–566, ten broken specimens. Occurrence.Late Tournaisian to early Viseán of the Qaidam Basin, northwest China. Discussion.Athyris expansa (Phillips) has been applied to a wide variety of athyridid shells with broad morphological variations. A re-examination of the lectotype material of A. expansa (Phillips) by Brunton (1980, p. 222–223) exhibits that the species could be separated into two different subspecies convertible to Actinoconchus: A. expansa expansa (Phillips) is characterized by a transversely oval shell having a nearly equibiconvex profile, short delicate flanges originating from the weakly developed growth lines, and a shallow but distinct median groove on both valves; A. expansa patula Brunton, 1980, representing the specimens with flattened shells, a rather transverse outline, a slightly

ventri-biconvex to plano-convex profile, finer growth lamellae, and fine radial ribs when the outer shell layers removed. Here it is argued that the stated differences between the two subspecies may well justify a separation at a higher taxonomic level. We therefore raise the two subspecies to species rank and assign them to two different genera: retaining Actinoconchus expansa (Phillips), but altering A. patula Brunton to Bruntonathyris on account of its transverse shell outline, a nearly plano-convex profile and fine radial striae when the outer shell layers are shed. Bruntonathyris patula (Brunton) shares many features with B. amunikeensis, especially in terms of shell outline, ornamentation and development of sulcus and fold, but, when compared with B. amunikeensis, the former is much more transverse in shape and its sulcus and fold are less developed. In addition, adult specimens of the new species occasionally exhibit weakly plicate anterior

← FIGURE 8—Bruntonathyris amunikeensis n. gen. and sp. 1–4, 16, NMV P309561, a specimen in dorsal, ventral, posterior, and anterior views, 31.5, enlargement of the ventral umbonal region showing radial striae when outer layer of shell removed, 33.0; 5, 9–10, 12, NMV P309562, a specimen in lateral, ventral, dorsal and posterior views, 31.6; 6–8, 11, 13, Holotype, NMV P309563, a specimen in dorsal, posterior, ventral, anterior and lateral views, 31.4; 14–15, 19, NMV P309564, a specimen in ventral, posterior and dorsal views, 32.0; 17–18, 22, NMV P309565, a specimen in ventral and dorsal views, 31.7, enlargement of the internal mold of the dorsal umbonal region, 34.0; 20–21, NMV P309566, a specimen in ventral and dorsal views, 31.5. All from the Chengqianggou Formation of the Chuanshangou section.

856



857

TABLE 7—Shell dimensions (mm) of Bruntonathyris? heijianshanensis new genus and species. Specimens

Width

Length

Thickness

NMV P309567* NMV P309568

47.0 17.0

35.4 16.8

16.2 8.9

* holotype.

FIGURE 11—Serial sections of Bruntonathyris? heijianshanensis n. gen. and sp. (based on specimen NMV P309568). dp, dental plates; hp, hinge plate.

margins and a relatively more pronounced parasulcate anterior commissure, features unknown in B. patula. BRUNTONATHYRIS? HEIJIANSHANENSIS new species Figures 10.1–10.5, 10.12–10.14, 10.25, 11 Diagnosis.Slightly transversely oval Bruntonathyris, with a dorsibiconvex profile, and rounded, evenly convex dorsal umbo; sulcus distinct, originating from umbo; fold low, commencing well anterior to umbo; shells strongly striated when outer layer removed. Description.Medium-sized for genus; cardinal extremities rounded, slightly compressed; anterior commissure parasulcate; antero-lateral and anterior margins slightly bent ventrally. Ventral valve moderately convex; beak small, pointed, slightly incurved; foramen large, rounded, touching dorsal umbo; lateral slopes indistinct; median sulcus shallow, distinct, originating at beak. Dorsal valve most convex at umbo; beak small, strongly incurved, deeply buried under ventral delthyrium; median fold indistinct, commencing anterior to umbo. Lamellose growth lines dense, fine, about six in 10 mm near anterior margin; lamellae narrow, delicate; radial striation prominent on exfoliated specimens, striae fine, dense, occasionally bifurcate, about 11 in 10 mm at umbo. Internal features (Fig. 11) are similar to that of B. amunikeensis. Etymology.Named for the Heijianshan section. From the Heijianshan section of the Qaidam Basin. Type.NMV P309567 is selected herein as holotype. Measurements.See Table 7. Other material examined.NMV P309568 and three broken specimens. Occurrence.Viseán of the Heijianshan section, the Qaidam Basin, northwest China.

Discussion.Compared with B. amunikeensis, the present species is proportionately less transverse and more circular in outline, and has a relatively more biconvex profile and a more conspicuously convex dorsal umbo. The new species also recalls Atrythyris lacrimula Struve, 1965, also illustrated by Alvarez et al. (1996, pl. 4, fig. 22), in having an oval outline, strongly striated shells when outer layer removed, and distinctive lamellose growth lines; the latter however is generally subpentagonal, strongly striated on external surfaces, and bearing a prominent sulcus and fold. BRUNTONATHYRIS? sp. Figure 10.6–10.11 Diagnosis.Slightly oval to subcircular outline, with weakly uniplicate anterior commissure, broad and delicate lamellae. Description.Shell about 38.8 mm long, 40.5 mm wide and 14.5 mm thick; slightly dorsibiconvex in profile; cardinal extremities rounded. Ventral beak slightly incurved; foramen small, rounded; median sulcus weakly developed, originating anterior to umbo, forming broad, shallow extension at anterior margin. Dorsal valve convex; beak small, deeply buried under ventral delthyrium; umbonal region slightly convex to flattened; median fold indistinct, commencing anterior to umbo. Lamellae broad, weakly delicate. Teeth stout, dental plates short. Dorsal interior cardinal plate subquadrate; crural bases strong, divergent; jugal saddle well-developed. Material examined.NMV P309569 and NMV P309574. Occurrence.Tournaisian to Viseán of the Qaidam Basin, northwest China. Discussion.Two distinctive specimens are herein tentatively referred to Bruntonathyris because of their fairly dorsibiconvex profile, weakly parasulcate anterior commissure, slightly delicate lamellae, and internal features to that of Bruntonathyris. However, these specimens clearly differ from the type species of the genus in having a more circular outline, a relatively more strongly parasulcate anterior commissure, slightly broader lamellae, and absence of radial striation even on exfoliated specimens. Bruntonathyris patula also cannot include the present specimens as it is much more transverse, has more flattened shells with a planoconvex profile, and lacks broad lamellae. Bruntonathyris? heijianshanensis also differs from the present specimens in having a more strongly parasulcate commissure, narrower, more weakly delicate lamellae, and more conspicuous striae. The presence of a subcircular outline, a parasulcate commissure and broad, weakly delicate lamellae of our specimens also recalls

← FIGURE 10—1–5, 12–14, 25, Bruntonathyris? heijianshanensis n. gen. and sp. 1–5, 25, Holotype, NMV P309567, a specimen in ventral, dorsal, posterior, anterior and lateral views, 31.1, enlargement of radial striae when outer shell layer removed, 32.3, all from the Xihansitegou Formation of the Heijianshan section; 12–14, NMV P309568, a specimen in dorsal, ventral and anterior views, from the Chengqianggou Formation of the Chuanshangou section, all 31.2. 6–11, Bruntonathyris? sp. 6–10, NMV P309569, a specimen in dorsal, ventral, anterior, posterior and lateral views, from the Xihansitegou Formation of the Heijianshan section, all 31.2; 11, NMV P309574, a specimen in dorsal view, from the Chuanshangou Formation of the Chuanshangou section, 31.0. 15–24, Composita ambigua (Sowerby, 1822). 15–19, NMV P309570, a specimen in posterior, ventral, dorsal, lateral and anterior views, from the Chuanshangou Formation of the Chuanshangou section, all 32.0; 20–24, NMV P309571, a specimen in lateral, posterior, anterior, ventral and dorsal views, from the Xihansitegou Formation of the Heijianshan section, all 31.6.

858


FIGURE 12—Serial sections of Composita ambigua (based on specimen NMV P309570). dp, dental plates; hp, hinge plate.

Carteridina Alvarez, Rong, and Boucot, 1998. This genus, however, cannot include our specimens due to its more highly convex valves, strongly uniplicate anterior commissure, pronounced fold and sulcus, and absence of dental plates and a dorsal myophragm. Subfamily SPIRIGERELLINAE Grunt, 1965 Genus COMPOSITA Brown, 1849 Type species.Spirifer ambiguus SOWERBY, 1822, p. 105, pl. 376. COMPOSITA AMBIGUA (Sowerby, 1822) Figures 10.15–10.24, 12 Spirifer ambiguus SOWERBY, 1822, p. 105, pl. 376. Terebratula ambigua (SOWERBY, 1822). PHILLIPS, 1836, p. 221, pl. 11, fig. 21. Athyris ambigua (SOWERBY, 1822). DAVIDSON, 1858–1863, p. 77, pl. 15, figs. 16–22; pl. 17, figs. 11–14; KONINCK, 1887, p. 75, pl. 19, figs. 25– 27, 30–38; SARYCHEVA AND SOKOLSKAYA, 1952, p. 235, pl. 69, fig. 399. Athyris (Composita) ambigua (SOWERBY, 1822). ROTAI, 1931, p. 96, pl. 10, figs. 8–9; JANISCHEWSKY, 1935, p. 90, pl. 4, figs. 1–6. Composita ambigua (SOWERBY, 1822). GOBBETT, 1963, p. 166, pl. 22, figs. 14–17; NALIVKIN, 1973, p. 76, pl. 24, fig. 11; POLETAEV, 1975, p. 101–107, pl. 25, figs. 1–8; GRUNT, 1976, p. 72, pl. 6, figs. 13–22; GRUNT, 1980, p. 104–108, pl. 3, figs. 1–8; pl. 4, figs. 1–7; BRUNTON, 1980, p. 230, figs. 26–28.

Discussion.Our material appears identical with the lectotype of Composita ambigua selected by Brunton (1980, fig. 26), as both are characterized by a pentagonal outline, a large ventral foramen, a distinct sulcus and fold, and a shallow but distinct median groove on the fold. Internally, these specimens also possess a pair of short but high dental plates (Fig. 12), and a subquadrate cardinal plate with strong, united crural bases, and ventrally concave inner hinge plates. Early Carboniferous C. subtilita (Hall), figured by Boucot et al. (1965, fig. 537.2a–c), is distinct from the present species by the possession of more pronounced concentric growth lines, more weakly developed sulcus and fold, and absence of a median groove on the fold. Like C. ambigua, C. humerosa Ivanov (see also Grunt, 1980, pl. 4, figs. 8–16), apparently also has a distinct sulcus and fold, but the Russian species is more elongate, has a relatively smaller ventral foramen, and lacks a median groove on the fold. Composita glabrata Poletaev (1975, p. 110–112, pl. 26, figs. 5–9) from the Viseán of the Donets Basin, Russia, can be differentiated from C. ambigua by its less-pronounced sulcus and fold, and absence of a median groove on the fold. Carter (1987, 1988) described six species, five from the Early Carboniferous Banff Formation of western Alberta, Canada and one from the early Mississippian Glen Park Formation of Illinois and Missouri, USA. Among these, none has a distinctive sulcus and fold as in C. ambigua. Although C. cf. oblonga (Tolmachoff) of Carter (1987, p. 66, pl. 21, figs. 1–16) from the Tournaisian Banff Formation of western Alberta is comparable with C. ambigua in terms of ventral foramen and a median groove on the fold, it is elongately oval in outline. Owing to sharing similar shell outline, external ornamentation and internal structures, Gruntathyris Gretchishnikova, 1996, has been treated as a synonym of Composita by Alvarez et al. (1998). As such, all of the three species described by Gretchishnikova (1996) from the Upper Devonian to Lower Carboniferous of Transcaucasus are suitable for Composita. Of these, C. innae (Gretchishnikova, 1996, p. 53, pl. 8, figs. 1–2) is differentiated from C. ambigua by its more elongate outline, weakly developed median sulcus, and lack of a median groove on the median fold. Internally, the hinge plates of C. innae are separated posteriorly and united anteriorly, whereas C. ambigua has the united hinge plate (see Fig. 12). Composita subtau (Gretchishnikova, 1996, p. 56, pl. 8, fig. 4) bears prominent concentric lamellae and possesses a relatively deeper median sulcus on the ventral valve, and C. pseudointermedia (Gretchnishnikova, 1996, p. 54, pl. 8, fig. 3) is more

Material examined.NMV P309570–571 and two broken specimens, of these, NMV P309570 and two broken specimens are sectioned. Measurements.See Table 8. Occurrence.Cosmopolitan species; Tournaisian to Viseán of Britain, France-Belgium Basin, Urals-Russian Platform, Spitzbergen, Altai Mountains of Siberia, South China, and Qaidam Basin of northwest China. TABLE 8—Shell dimensions (mm) of Composita ambigua (Sowerby, 1822). Specimens

Width

Length

Thickness

NMV P309570 NMV P309571

16.8 22.0

15.5 21.6

8.9 11.8

FIGURE 13—Exterior of Lochengia lochengensis, showing concentric growth lines and plicae [based on lectotype specimen NMV P309570, figured by Jin (1983, pl. 1, fig. 11)]. Scale bar represents 30 mm.


859

FIGURE 14—1–10, Lochengia qinghaiensis n. sp. 1–2, 5, NMV P309572, a broken specimen in ventral, dorsal and anterior views, 31; 3, 3–4, 6–9, Holotype, NMV P309573, a specimen in posterior, ventral, anterior, lateral, and dorsal views, 31.4; enlarged shell showing external ornaments, 33.0. All from the Chengqianggou Formation of the Chuanshan.

circular and much smaller, and lacks a sulcus and fold. Subsequently, the latter two species both cannot be confused with C. ambigua. Subfamily LOCHENGIINAE Ching and Yang (in Yang et al., 1977) Genus LOCHENGIA Yoh, 1929 (5 FLEXATHYRIS Grunt, 1980) Lochengia YOH, 1929, p. 70; GRABAU, 1931a, p. 478; CHING AND YANG in YANG ET AL., 1977, p. 412; JIN, 1983, p. 227–228; ALVAREZ AND RONG, 1995, p. 604–605; ALVAREZ ET AL., 1998, p. 483. Flexathyris GRUNT, 1980, p. 61; GRUNT, 1986, p. 89–91.

Type species.Lochengia lochengensis Ching and Yang in Yang et al., 1977, p. 412. Nomenclature discussion.Alvarez and Rong (1995) have discussed the status of Lochengia Yoh, 1929 and Cryptospirifer Grabau, 1931b, recognizing both as being valid and available taxonomic names. Lochengia is now restricted to Early Carboniferous, while Cryptospirifer is confined to the Permian. Revised diagnosis.Typical Lochengiinae, ventribiconvex to plano-convex valves in profile, sulcus and fold present but usually weak; anterior commissure slightly sulciplicate. Description.Medium to large Lochengiinae; transversely

oval in outline; anterior commissure broadly sulciplicate; shell thick without punctae. Ventral foramen mesothyridid; median sulcus shallow, narrow. Dorsal umbo flattened to slightly swollen; median fold low, indistinct. Shell surface weakly plicate anteriorly in juveniles, strongly plicate to slightly costate posteriorly at maturity; lamellose growth lines prominent (Fig. 13). Teeth stout, rounded; dental plates low, long, divergent along margins of fanshaped muscle scar regions; Cardinal plate slightly rectangular to subtrapezoidal, performate apically; jugal saddle similar to that of Athyris. Included species.See Table 4. Discussion.Lochengia is often confused with several forms of the Lochengiinae, notably Cryptospirifer Grabau, 1931b; Galeatathyris Jin, 1983; and Titanothyris Jin and Hu (in Wang et al., 1982). However, Cryptospirifer is characterized by equally biconvex valves, rectimarginate anterior commissure, and absence of a median sulcus and fold and radial ornament. Despite possessing radial ornament like Lochengia, Titanothyris is usually more densely and strongly costate. Galeatathyris is distinct from Lochengia in having an oval outline, a concavo-convex profile, a more highly convex ventral umbo and weaker radial ornaments, and the presence of a low median fold, instead of a median sulcus, on the ventral valve.

860

JOURNAL OF PALEONTOLOGY, V. 77, NO. 5, 2003 when compared with L. lochengensis. Lochengia enteletiformis (Grabau), validated by Ching and Yang (in Yang et al., 1977) and redescribed by Jin (1983, p. 228–229, pl. 1, figs. 1–3, 9, 12), clearly differs from L. qinghaiensis by its smaller size, relatively elongate outline and higher, more sharply crested plicae. Two Russian species: Lochengia prokofjevi (Grunt, 1980, p. 61–62, pl. 4, figs. 17–18) and L. variabilis (Mo¨ller, 1862, p. 78, pl. 6, figs. 3–6), also described by Nalivkin (1975, p. 203, pl. 94, figs. 4–6) and by Poletaev (1975, p. 92–93, pl. 22, figs. 5–6), are considered herein congeneric with the new species in view of ventribiconvex to plano-convex profile, weak sulcus and fold, slightly uniplicate anterior commissure, and typical ornaments of variedly developed plicae and prominent lamellose growth lines. When compared with the new species, L. prokofjevi is distinct by its slightly concave dorsal valve and smaller number of plicae, while L. variabilis is differentiated by its lower and greater number of plicae, usually eight or nine plicae on each valve. ACKNOWLEDGMENTS

FIGURE 15—Serial sections of Lochengia qinghaiensis n. sp. (based on specimen NMV P309572). dp, dental plates; hp, hinge plate.

LOCHENGIA

new species Figures 14, 15 Diagnosis.Average sized (see measurements) and flattened Lochengia, with rounded and broad plicae, two on each flank. Description.Transversely oval shell; ventri-biconvex to slightly plano-convex in lateral profile; anterior commissure moderately sulciplicate. Ventral beak small, strongly incurved, overhanging hingeline; umbonal region convex; median sulcus shallow in juveniles, deep at maturity, originating anterior to umbo, defined by pair of pronounced sulcal-bounding plicae. Dorsal median fold low, originating anterior to umbo. Plicae broad, originating anterior to umbo, low, rounded in juveniles, and high with ridge-shaped crests at maturity; two plicae on each flank; lamellose growth lines fine, distinct, about 15–16 in 10 mm at midvalve. Teeth stout, rounded, slightly curved dorsally; dental plates low, thin, fairly long. Cardinal plate slightly subtrapezoidal, flattened; dorsal foramen large, round, situated posterior to cardinal plate, open, under delthyrium (Fig. 15). Etymology.Named for Qinghai Province, northwest China. Type.NMV P309573 is herein designated as holotype. From the Chuanshangou section of the Qaidam Basin. Other material examined.NMV P309572 and several broken shells. Measurements.See Table 9. Occurrence.Late Tournaisian to early Viseán of the Qaidam Basin, northwest China. Discussion.Lochengia lochengensis (Grabau) of Ching and Yang (in Yang et al., 1977, p. 412) and Jin (1983, p. 228, pl. 1, figs. 5–7, 10, 11) is probably the closest ally to the new species, but it has a lower, more rounded and greater number of plicae QINGHAIENSIS

TABLE 9—Shell dimensions (mm) of Lochengia qinghaiensis new species. Specimens

Width

Length

Thickness

NMV P309572 NMV P309573*

56.0 38.6

38.0 23.0

14.5 8.6

* holotype.

G. R Shi acknowledges the continue supporting from the Australian Research Council for his research on the Late Paleozoic brachiopod faunas and biogeography of the Asian-western Pacific region. Two research grants (to G. R. Shi) from the Australian Research Council also enabled Zhan Li-Pei and ZQC to visit Australia in 1995 and 2002, respectively. Our thanks are also rendered to Dr. John Carter of Carnegie Museum, Pittsburgh, Professor Fernando Alvarez of Universidad de Oviedo, Oviedo, Spain, and Professor Jed Day of Illinois State University, Normal for their critical reviews and constructive suggestions. Dr. Howard Brunton of the Natural History Museum, London is also greatly acknowledged for his advice and providing some information of his studying materials of Lamellosathyris lamellosa (Le´veille´). This study is also financially supported by a Japan Society for the Promotion of Science grant (P01103, to ZQC). REFERENCES

ABRAMYAN, M. S. 1957. Brakhiopody verkhnefamenskikh i etrenskikh otlozhenii iugo-zapadnoii [Brachiopods of the Upper Famennian and Etroeungt deposits of S.W. Armenia]. 142 p., 16 pls. Erevan: Izd-vo AN Armianskoi SSR. (In Russian) ALVAREZ, F. 1999. Ornamentacioń desarrollada por algunos braquio´podos del orden Athyridida. Trabajos de Geologıá, 21:33–36. (In Spanish) ALVAREZ, F., AND C. H. C. BRUNTON. 2001. Chapter 12: fundamental differences in external spine growth in brachiopods, p. 108–118. In C. H. C. Brunton, L. R. M. Cocks, and S. L. Long (eds.), Brachiopods: past and present. The Systematics Association Special Volume Series, Volume 63, London: Taylor and Francis, 442 p. ALVAREZ, F., C. H. C. BRUNTON, AND W. STRUVE. 1996. On Athyris (Brachiopoda) and its type species ‘Terebratula’ concentrica VON BUCH. Senckenbergiana lethaea, 76(1/2):65–105. ALVAREZ, F., AND J. Y. RONG. 1995. On the taxonomic status of Lochengia and Cryptospirifer (Brachiopoda, Athyridida). Journal of Paleontology, 69(3):604–605. ALVAREZ, F., J. Y. RONG, AND A. J. BOUCOT. 1998. The classification of athyridid brachiopods. Journal of Paleontology, 72(5):827–855. BOUCOT, A. J., J. G. JOHNSON, AND R. D. STATON. 1964. On some atrypoid, retzioid, and athyridoid Brachiopoda. Journal of Paleontology, 38 (5):805–822, pls. 125–127. BOUCOT, A. J., J. G. JOHNSON, AND R. D. STATON. 1965. Suborders Retziidina and Athyrididina, p. H649–H667. In R. C. Moore (ed.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda, Volume 2. Geological Society of America and University of Kansas Press, Lawrence. BROWN, T. 1849. Illustrations of the fossil conchology of Great Britain and Ireland, with descriptions and localities of all species, 273 p., 98 pls. London. BRUNTON, C. H. C. 1980. Type specimens of some Upper Palaeozoic Athyridide brachiopods. Bulletin of the British Museum, London, 34: 219–234.

CHEN ET AL.—CARBONIFEROUS ATHYRIDID BRACHIOPODS FROM CHINA BRUNTON, C. H. C. 1984. Silicified brachiopods from the Viseán of County Fermanagh, Ireland (III). Rhynchonellids, spiriferids and terebratulids. Bulletin of the British Museum, London, 38(2):27–130. BRUNTON, C. H. C., F. ALVAREZ, AND D. I. MACKINNON. 1996. Morphological terms used to describe the cardinalia of articulate brachiopods: homologies and recommendations. Historical Biology, 11:9–41. BRUNTON, C. H. C., S. S. LAZAREV, R. E. GRANT, AND J. G. JIN. 2000. Productidina, p. 424–609. In A. Williams, C. H. C. Brunton, and S. J. Carlson with 44 other authors (eds.), Treatise on Invertebrate Palaeontology, Pt. H, Brachiopoda (revised), Volume 3, Linguliformea, Craniiformea and Rhynchonelliformea (part). Geological Society of America and The University of Kansas Press, Lawrence. ¨ ber Terebrateln, mit einem Versuch, sie zu clasBUCH, L. VON. 1834. U sificiren und zu beschreiben. Abhandlungen der Koeniglichen Akademie der Wissenschaften zu Berlin, 1833:21–144, pls. 1–3. (In German) BUCKMAN, S. S. 1906. Brachiopod Nomenclature. Annals and Magazine of Natural History, Series 7, 18(107):321–327. CARTER, J. L. 1967. Mississippian brachiopods from the Chappel Limestone of central Texas. Bulletins of American Paleontology, 53 (238): 253–488, pls. 13–45. CARTER, J. L. 1987. Lower Carboniferous brachiopods from the Banff Formation of Western Alberta. Bulletin of the Geological Survey of Canada, 53(378), 99 p., 29 pls. CARTER, J. L. 1988. Early Mississippian Brachiopods from the Glen Park Formation of Illinois and Missouri. Bulletin of the Carnegie Museum, Number 27, 82 p. CARTER, J. L. 1990. Subdivision of the Lower Carboniferous in North America by means of articulate brachiopod generic ranges. Courier Forschungsinstitut Senckenberg, 130:145–155. CARTER, J. L. 1999. Tournaisian (Early Osagean) brachiopods from a bioherm in the St. Joe Formation near Kenwood, Oklahoma. Annals of Carnegie Museum, 68:91–149. CARTER, J. L., AND R. C. CARTER. 1970. Bibliography and index of North American Carboniferous brachiopods (1898–1968). The Geological Society of America, Memoir, 128, 382 p. COOPER, G. A., AND J. T. DUTRO. 1982. Devonian Brachiopods of New Mexico. Bulletins of American Paleontology, 82–83(315), 215 p., 45 pls. DAVIDSON, T. 1858–63. British Fossil Brachiopoda. Permian and Carboniferous species. Palaeontographical Society (Monograph), London, 2(4):1–51, pls. 1–4; (5):1–280, pls. 5–60. DAVIDSON, T. 1881. On genera and species of spiral-bearing Brachiopoda, from specimens developed by the Rev. Norman Glass. Geological Magazine, 8:1–13. FRECH, F. 1916. Geologie Kleinasiens im Bereiche der Bagdadbahn. Ergebnisse einiger Reisen und palaeontologische Untersuchungen. Zeitschrift der Deutschen Geologischen Gesellschaft, 68(1–3):1–325, 24 pls. (In German) GAETANI, M. 1968. Lower Carboniferous brachiopods from central Elburz, Iran. Rivista Italiana di Paleontologia e Stratigrafia, 74(3):665– 744, pls. 47–52. GALITZKAJA, Y. 1977. Ranne- i srednekamennougol’nye produktidy severnoi Kirgizii [Early and Middle Carboniferous Productida of northern Kirghizia]. Akademiia Nauk Kirgizkoi SSSR, Frunze. 298 p., 64 pls. (In Russian) GOBBETT, D. J. 1963. Carboniferous and Permian brachiopods of Svalbard. Norsk Polarinstitutt Skrifter, Number 127, 201 p., 25 pls. GRABAU, A. W. 1931a. The Permian of Mongolia. A report on the Permian fauna of the Jisu Honguer Limestone of Mongolia and its relations to the Permian of other parts of the world. Natural History of Central Asia, Volume 4, 665 p., 35 pls. American Museum of Natural History, New York. GRABAU, A. W. 1931b. The Brachiopoda, Pt. 2. In Studies for Students, Science Quarterly of the National University of Peking, 2(3):397–422. GRETCHISHNIKOVA, I. A. 1996. The new athyridids (Brachiopoda) from Upper Devonian and Lower Carboniferous of the Transcaucasus. Paleontologicheskii Zhurnal, 1996(3):51–61, 1 pl. (In Russian) GRUNT, T. A. 1965. Nadsemeistvo Athyridacea [Superfamily Athyridacea], p. 237–253. In V. E. Ruzhencev and T. G. Sarycheva (eds.), Razvitic i smena morskikh organizmov na rubezhe paleozoia i mesozoia [Evolution and change of marine organisms at the Paleozoic-Mesozoic

861

boundary]. Akademiia Nauk SSSR Trudy Paleontologicheskogo Institut [Translation by D. A. Brown, 1968, Australian National University Publication 17], 108, 430 p. GRUNT, T. A. 1968. Rod Spirigerella i ego tipovoi vid Spirigerella derbyi. Paleontologicheskii Zhurnal, 1968(3):42–48. (In Russian) GRUNT, T. A. 1976. Gladkie atirididy (Brachiopoda) V Kamennougol’nykh otlozheniiakh podmoskov’ia [‘Smooth’ athyridides in the Carboniferous deposits of the Moscow Region]. Paleontologicheskii Zhurnal, 3:71–86. (In Russian) GRUNT, T. A. 1980. Atirididy Russkoi platformy [Athyridides of Russian Platform]. Akademiia Nauk SSSR, Trudy Paleontologicheskogo Institut (Moscow), 182, 164 p. (In Russian) GRUNT, T. A. 1986. Sistema brakiopod otriada Atiridida [Classification of brachiopods of the oder Athyridida]. Akademiia Nauk SSSR, Paleontologicheskogo Institut, Trudy, 215, 200 p. (In Russian) HALL, J. 1858. Palaeontology of Iowa, p. 473–724, 29 pls. In J. Hall and J. D. Whitney (eds.), Report of the Geological Survey of the State of Iowa: Embracing the Results of Investigations Made During Portions of the Years 1855, 56 and 57, Volume 1, Pt. 2. Albany: C. van Benthuysen. HALL, J., AND J. M. CLARKE. 1893. An introduction to the study of the genera of Palaeozoic Brachiopoda. New York Geological Survey, 8(2): 1–317(1893),319–394(1895), pls. 21–84. HALL, J., AND J. M. CLARKE. 1895. The new species of Brachiopoda described in Paleontology of New York, Volume 8, Pt. 1–2, 1892–1894, New York State Geologist, Annual Report, 14:325–402, 14 pls. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of Zoological Nomenclature (4th edition.) International Trust for Zoological Nomenclature, London, 306 p. JANISCHEWSKY, M. E. 1935. Fauna serpukhovskikh sloev raiona 43-go lista 10- verstnoi karty Evropeiskoi chasti SSSR. Uchenye Zapiski Leningradskogo Gosudarstvennogo Universiteta (LGU), Seriia GeologoPochvenno- Geograficheskaia, 1:77–96, 6 pls. (In Russian) JANISCHWSKY, M. E. 1954. Fauna brakhiopod nizhnego karbona leningradskoi oblasti. Leningrad: Izd-vo LGU, 279 p., 49 pls. (In Russian) JIN, Y. G. 1983. On the Family Lochengiidae. Bulletin of the Nanjing Institute of Geology and Palaeontology, Academia Sinica, 6:225–248, 3 pls. (In Chinese) JIN, Y. G., AND R. S. FANG. 1983. Early Carboniferous brachiopods from Shidian, Yunnan. Acta Palaeontologica Sinica, 22(2):150–159, 3 pls. (In Chinese) JIN, Y. G., H. K. XU, D. L. SUN, AND S. L. YE. 1979. Brachiopoda. p. 60–225, pls. 18–57. In Palaeontological Atlas of northwestern China, Qinghai Volume, 293 p., 96 pls., Beijing: Geological Publishing House. (In Chinese) KAMMER, T. W., P. L. BRENCKLE, J. L. CARTER, AND W. I. AUSICH. 1990. Redefinition of the Osagean-Meramecian boundary in the Mississippian stratotype region. Palaios, 5:414–431. KING, W. 1850. A monograph of the Permian fossils of England. Palaeontographical Society, Monograph, 3(1), 258 p., 28 pls. KONINCK, L. G. DE. 1887. Faune du Calcaire Carbonifere de la Belgique. VI. Brachiopodes. Musee Royal d’Histoire naturelle de Belgique, Annales, 14, 154 p., 37 pls. LEGRAND-BLAIN, M., J. J. DELVOLVE, AND M. HANSOTTE. 1996. Carboniferous Brachiopods from the Arize Massif (Ariege), French Pyrenees. Geobios, 29(2):177–207. LE´VEILLE´, C. 1835. Apercu geologique de quelques localites tres riches en coquilles sur les frontieres de France et de Belgique. Societe geologique de France, Memories, Series 1 (2):29–40, 2 pls. (In French) LITVINOVITCH, N. V. 1962. Kamennougol’nye i permskie otlozheniia zapadnoi chasti Tsentral’nogo Kazakhstana [The Carboniferous and the Permian deposits of the western part of central Kazakhstan]. Materialy po Geologii Tsentral’nogo Kazakhstan, 4:1–389, 40 pls. (In Russian) LU, J. D., AND J. M. ZHAO. 1962. Tetracorals of the Lower Carboniferous of the Qilianshan Region. Memoirs of the Geology of Qilianshan, 4(3): 356–384. Geological Publishing House, Beijing. (In Chinese) MARTINEZ-CHACON, M. L., AND M. LEGRAND-BLAIN. 1992. Braquio´podos, p. 91–144, 10 pls. In S. Rodriguez, M. E. Arribas, M. J. ComasRengifo and eleven others (eds.), Analisis paleontolo´gico y sedimentolo´gico de la cuenca carbonı´fera de los Santos de Maimona (Badajoz). Coloquios de Paleontologia, Universidad Complutense de Madrid, Madrid, 44, 303 p. (In Spanish)

862


MCCOY, F. 1844. A synopsis of the characters of the Carboniferous Limestone fossils of Ireland. 207 p., 29 pls. Dublin. METCALFE, I. 1996. Gondwanaland dispersion, Asian accretion and evolution of eastern Tethys. Australian Journal of Earth Sciences, 43:605– 620. MO¨LLER, V. I. VON. 1862. Geologicheskie i paleontologicheskie zametki ob osadkakh gorno- izvestkovoi formatsii sklonov khrebta Ural’skogo [Carboniferous palaeontology and geology of the Ural Mountains]. Gornyi Zhurnal, 4(2):163–208, pls. 1–9. (In Russian) MU, E. Z., Z. Y. YANG, C. M. YU, Z. L. ZHU, AND J. X. YIN. 1963a. The geological sections between the Huaitoutala Coal mine and Keluke. Geology of Chilianshan, 2(1):197–199. Science Press, Beijing. (In Chinese) MU, E. Z., Z. Y. YANG, C. M. YU, Z. L. ZHU, AND J. X. YIN. 1963b. The outcrop sections west to Huaitoutala Coal mine of the Oulongbuluke county. Geology of Chilianshan, 2(1):200–201. Science Press, Beijing. (In Chinese) MU, E. Z., Z. Y. YANG, C. M. YU, Z. L. ZHU, AND J. X. YIN. 1963c. The Chengqianggou section of the Oulongbuluke county. Geology of Chilianshan, 2(1):201–203. Science Press, Beijing. (In Chinese) MUIR-WOOD, H. M., AND G. A. COOPER. 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Memoir of the Geological Society of America, 81, 447 p., 135 pls. NALIVKIN, D. V. 1937. Brakhiopody verkhnego i srednego devona i nizhnego karbona Severo- Vostochnogo Kazakhstana [Brachiopoda of the Upper and Middle Devonian and Lower Carboniferous of northeastern Kazakhstan]. Tsentral’nyi Nauchno-Issledovatel’skii GeologoRazvedochnyi Institut, Trudy, 99, 200 p., pls. 1–39. (In Russian) NALIVKIN, D. V. 1975. Brakhiopody, p. 154–203, pls. 62–94. In D. L. Stepanov (ed.), Paleontologicheskii Atlas Kamennougol’nykh Otlozhenii Urala [Paleontological Atlas of Carboniferous deposits of the Urals]. Vsesoiuznyi Neftianoi Nauchno-Issledovatel’skii Geologo-Razvedochnyi Institut, Trudy, 383. (In Russian) NALIVKIN, D. V. IN NALIVKIN, D. V., AND N. N. FOTIYEVA. 1973. Brakhiopody pogranichnykh otlozhenii turneiskogo i vizeiskogo iarusov zapadnogo’sklona Urala. Moscow: Nauka, 119 p. (In Russian) OZAKI, J. 1939. On some lower Carboniferous Brachiopods from Central Hunan, China. Shanghai Science Institute Journal, Series 2, 2(2):225– 282, pls. 35–43. PHILLIPS, J. 1836. The Mountain Limestone District. Illustrations of the geology of Yorkshire, 2, 253 p., 25 pls. London. POLETAEV, V. I. 1975. Rannekamennougol’nye i bashkirskie gladkie spiriferidy i atiridy Donetskogo basseina. 140 p., 27 pls., Izdatel’stvo ‘Naukova Dumka’, Kiev. (In Russian) ROTAI, A. 1931. Brakhiopody i stratigrafiia nizhnego karbona Donetskogo basseina [Brachiopods and stratigraphy of Lower Carboniferous of the Donetz Basin]. Glavnogo Geolog-Razvedochnogo Upravleniya, Trudy, 73:35–144, pls. 1–10. (In Russian) ROTAI, A. 1941. Klass Brachiopoda, p. 85–117, pls. 14–29. In L. S. Librovich (ed.), Atalas rukovodiashchikh form iskopaemykh faun SSSR, Tome 4, Nizhnii otdel kamennougol’noi sistemy, Leningrad: Gosgeolizdat. SARYCHEVA, T. G., AND A. V. N. SOKOLSKAYA. 1952. Opredelitel’ paleozoiiskikh brakhiopod podmoskovnoi kotloviny [A description of the Paleozoic Brachiopoda of the Moscow Basin]. Akademiia Nauk SSSR, Paleontologicheskii Institut, Trudy, 38, 307 p., 71 pls. (In Russian) SARYCHEVA, T. G., A. V. N. SOKOLSKAYA, G. A. BESNOSOVA, AND S. V. MAXIMOVA. 1963. Brakhiopody i paleogeografiia karbona Kuznetskoi kotloviny [Carboniferous brachiopods and paleogeography of the Kuznetsk Basin]. Akademiia Nauk SSSR, Paleontologicheskii Institut, Trudy, 95, 547 p., 64 pls. (In Russian) SCOTESE, C. R., AND W. S. MCKERROW. 1990. Revised world maps and introduction. In W. S. McKerrow and C. R. Scotese (eds.), Paleozoic Paleogeography and Biogeography. Geological Society Memoir, 12:1– 21. STAINBROOK, M. A. 1947. Brachiopoda of the Percha Shale of New Mexico and Arizona. Journal of Paleontology, 21(4):297–328, pls. 44–47. SHI, X. D. 1983. Discovery of the Tournaisian in the Mountain Amunike of the northeastern Qaidam Basin, with a discussion on the subdivision

of the Lower Carboniferous in the Mountain Oulongbuluke. Contributions to Geology of Qinghai-Xizang Plateau 2:41–68. Geological Publishing House, Beijing. (In Chinese) SOWERBY, J. 1822. The Mineral Conchology of Great Britain, 4:17–114, pls. 319–383. London. STRUVE, W. 1965. Beitraege zur Kenntnis devonischer Brachiopoden, 12: Atrythyris n. g. und ihre Arten (Athyridae). Senckenbergiana lethaea, 46(2/3):217–228. (In German) STRUVE, W. 1992. Neues zur stratigraphie und fauna des rhenotypen Mittel Devon. Senckenbergiana Lethaea, 71:503–624. (In German) WANG, G. P., Q. Z. LIU, Y. G. JIN, S. Z. HU, W. P. LIANG, AND Z. T. LIAO. 1982. Brachiopoda, p. 186–256, pls. 74–102. In Palaeontological Atlas of east China (Late Paleozoic), 2. Geological Publishing House, Beijing, 495p., 157 pls. (In Chinese) WANG, Z. J. 1981. Early Lower Carboniferous stratigraphy of northwest and north China. Geological Review, 27(6):533–538. (In Chinese) WANG, Z. J. 1987. Lower Carboniferous stratigraphy and coral fossil sequence in the Amunike Mountain of the north Qaidam Basin. Bulletin of the Institute of Geology, Chinese Academy of Geological Sciences, 16:51–99, 15 pls. (In Chinese) WANG, Z. J. 1990. Kunlun-Qaidam Region, p. 67–92. In Z. J. Wang (ed.), The Carboniferous System of China. Stratigraphy of China, 8. Geological Publishing House, Beijing, 419 p., 22 pls. (In Chinese) WELLER, S. 1914. The Mississippian Brachiopoda of the Mississippi Valley Basin. Illinois State Geological Survey, Monograph, 1:1–508, 83 pls. WILLIAMS, J. S. 1943. Stratigraphy and fauna of the Louisiana Limestone of Missouri. United States Geological Survey Professional Paper, 203, 133 p., pls. 1–9. WILLIAMS, A., AND C. H. C. BRUNTON. 1997. Morphological and anatomical terms applied to brachiopods, p. 421–440. In A. C. Williams, C. H. C. Brunton, and S. J. Carlson (eds.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda (revised) 1, Introduction. Geological Society of America and the University of Kansas Press, Lawrence, 539 p. WOLFART, R. 1965. Die Fauna des karbons vom Jebel Abd-Et-Aziz (Nordost-Syrien), 1: Anthozoa, Bryozoa, Brachiopoda. Geologisches Jahrbuch, 83:277–326, 6 pls. WU, W. S., L. X. ZHANG, X. G. ZHAO, Y. G. JIN, AND Z. T. LIAO. 1987. Carboniferous stratigraphy in China. Science Press, Beijing, 160 p. YANAGIDA, J. 1962. Carboniferous Brachiopoda from Akiyoshi, southwest Japan, Pt. 1. Memoirs of Faculty of Sciences, Kyushu University, Series D (Geology), 12(1):87–127, 8 pls. YANG, D. L., S. Z. NI, M. L. CHANG, AND R. X. ZHAO. 1977. Brachiopoda, p. 306–470, pls. 129–188. In Palaeontological Atlas of CentralSouth China, 2. Geological Publishing House, Beijing. (In Chinese) YANG, J. Z., J. Z. SHENG, W. S. WU, AND L. H. LU. 1962. The Carboniferous of China; Symposium of First All-China Stratigraphy Congress. Science Press, Beijing, 112 p. (In Chinese) YANG, S. P. 1964. The Lower and Middle Carboniferous brachiopod fauna from the northern slope of the Mountain Borohoro, Xinjiang. Science Press, Beijing, 179 p., 23 pls. (In Chinese) YANG, S. P. 1978. Lower Carboniferous brachiopods of Guizhou Province and their stratigraphic significance, p. 78–142, pl. 17–39. In Professional Papers of Stratigraphy and Palaeontology, 5. Geological Publishing House, Beijing. (In Chinese) YANG, S. P. 1990. Brachiopoda, p. 325–332. In Z. J. Wang (ed.), The Carboniferous System of China. Stratigraphy of China, 8. Geological Publishing House, Beijing, 419 p., 22 pls. (In Chinese) YOH, S. S. 1929. Geology and mineral resources of northern Kwangsi. Geological Survey of Kwangtung and Kwangsi, Annual report, 2(2): 59–108, 2 pls. YU, C. C., AND Y. D. LIN. 1961. Discussion on the correlation of Early Carboniferous stratigraphy of southern and northern slopes of the Chilianshan as based upon the study of its Anthozoa content. Acta Geologica Sinica, 41(2):154–174. (In Chinese) ZHANG, Z., F. M. ZHANG, Z. X. ZHANG, AND Z. WANG. 1983. Brachiopoda, p. 262–386, pls. 86–145. In Regional Geological Survey Team of Xinjiang, Institute of Geosciences of Xinjiang, Geological Survey Group of Petroleum Bureau of Xinjiang (eds.), Palaeontological Atlas of Northwestern China (Xinjiang Part), Paleozoic Pt. 2. Geological Publishing House, Beijing, 785 p., 226 pls. (In Chinese) ACCEPTED 13 DECEMBER 2002