J. Paleont., 77(6), 2003, pp. 1053-1068 Copyright ? 2003, The Paleontological Society 0022-3360/03/0077-1053$03.00

A GUADALUPIAN-LOPINGIAN (MIDDLE TO LATE PERMIAN) BRACHIOPOD FAUNA FROM THE JURIPU FORMATION IN THE YARLUNG-ZANGBO

SUTURE ZONE, SOUTHERN TIBET, CHINA G. R. SHI,1 SHUZHONG SHEN,2 AND LIPEI ZHAN3

'School of Ecology and Environment, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood, Victoria 3125, Australia, <grshi@deakin.edu.au>, 2Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 Eastern Beijing Road, Nanjing,

Jiangsu Province 210008, P. R. China, <szshen@nigpas.ac.cn>, and 31nstitute of Geology, Chinese Academy of Geological Sciences, Baiwanzhuang Road, Beijing 100037, P R. China

ABSTRACT-A brachiopod fauna of 16 species belonging to 11 genera and three genera and species indeterminate from the middle and upper parts of the Juripu Formation in the Yarlung-Zangbo (Indus-Tsangbo) Suture zone (=Yarlung-Zangbo River zone), southern Tibet, is described and figured for the first time. A new species, Taeniothaerus zhongbaensis, is described. The fauna is comparable with that in the Kalabagh Member of the Wargal Formation of the Salt Range, Pakistan, and is considered to be most likely Capitanian (late Guadalupian/Middle Permian) to Wuchiapingian (early Lopingian/early Late Permian) in age, as indicated by the majority of the brachiopod species and by being constrained by an underlying fusulinacean fauna (Parafusulina Zone) and an overlying ammonite fauna (Cyclolobus fauna).

INTRODUCTION

P ERMIAN SEDIMENTARY rocks are widely distributed within the Yarlung-Zangbo (Indus-Tsangbo) suture zone (roughly cor-

responding to the Yarlung-Zangbo River zone) in southern Tibet. These rocks are generally divided into two major types in view of their lithology, internal stratigraphy and contact relationships with surrounding rocks. The first type is characterized by abundant clas- tic sediments and some bioclastic limestone, with intact internal stratigraphy and a normal, or paraconformable, stratigraphical re- lationship with overlying Triassic rocks and a disconformable re- lationship, with the underlying Carboniferous deposits. The second type is distinguished mainly by gray or purple pure bioclastic lime- stone with paleobiogeographically mixed Gondwanan/Paleoequa- torial faunas (Shen et al., 2003, 2003a, 2003b, 2003c). These car- bonate successions usually crop out as isolated hills or blocks over- lying, or embedded within, a ml1ange of Mesozoic (mainly Trias- sic) flysch deposits (Wang et al., 1999; Yin, 1997) in southern Tibet. The nature of the contact between the isolated Permian lime- stone blocks and adjacent Mesozoic rock bodies is unknown either due to poor exposure or marked by a fault. These isolated Permian limestones, otherwise also known as "exotic blocks" in the liter- ature (e.g., Wang et al., 1999; Yin, 1997) are very distinct within the Yarlung-Zangbo suture zone and commonly form prominent hill tops; however, their origins remain poorly understood (see re- view and discussion in Guo et al., 1991; Yin, 1997).

The above-mentioned two Permian rock types of the Yarlung- Zangbo suture zone may be correlated with similar facies from the broad region of southern Tibet and the Himalaya. Diener (1903) first recognized and broadly defined two main types of Permian deposits in the Central Himalayan region based on both faunal and lithologic characteristics. These were informally named "Tibetan facies" and "Himalayan facies," with the former embracing all the Permian sedimentary deposits south of the Yar- lung-Zangbo suture zone and the latter being defined for the "ex- otic" limestone blocks exposed within and along the Yarlung- Zangbo suture zone. In a recent overview of the Permian, espe- cially the Lopingian (Upper Permian), stratigraphy of southern Tibet, Shen et al. (2003) have further refined the definition of Diener's "Himalayan-type" Permian facies, in that they subdivid- ed the "Himalayan facies" into two further types in view of the differences in vertical successions and depositional environments: the "Qubu-type" and the "Selong-type." The "Qubu-type" Permian facies, widely distributed throughout the Himalaya, is considered to represent a succession of marine coastal and barrier- lagoon depositional environments accumulated in the proximal

part of northern Gondwanaland. The "Selong-type" Permian de- posits, on the other hand, are confined to the central part of south- ern Tibet (principally exposed in the Jilong and Selong areas, see Shen et al., 2003 for more details) and are considered to have accumulated in a more distal offshore marine environment. In this framework, the mixed clastic and carbonate succession of the Yar- lung-Zangbo suture zone mentioned before may belong to the "Qubu-type," while the pure bioclastic limestone facies (i.e., the "exotic" limestone blocks mentioned earlier) is undoubtedly an integral part of Diener's "Tibetan-type" Permian facies [=the "Chitichun-type" Permian deposits of Shen et al. (2003)].

Permian brachiopods are common in both the "Qubu-type" and "Tibetan-type" rocks of the Yarlung-Zangbo suture zone. Diener (1897) was first to describe a large Permian brachiopod fauna from one of the Permian "exotic" limestone blocks, the Chitichun Limestone, in the southern part of the Nagri District, southwestern Tibet. More recently, Shen et al. (2003a, 2003b) documented two other brachiopod faunas from two separate lime- stone blocks in the Zhongbei mdlange in the Lhaze County of southern Tibet. In contrast, brachiopod faunas from the "Qubu- type" Permian rocks of the Yarlung-Zanbo suture zone have not been documented although they are well known from similar Permian sequences in the Himalayan region (see Shen et al., 2003 for a review of these works).

This paper is the first systematical study of the Permian bra- chiopods from the "Qubu-type" Permian facies of the Yarlung- Zangbo suture zone. This brachiopod fauna was collected by the officers of the Geological Survey of Tibet in 1979 from a section located about 25 km northwest of the Zhongba County town (Fig. 1.1). The collections were initially sent to one of us (L.-P. Zhan) in 1980 for brief identification and age determination, but have never been described. As will be seen later, the quality and quan- tity of the material for this study is limited due to the reconnais- sance nature of the field work conducted by the field geologists in this remote area, but we believe that the bulk of the collections is adequate enough to highlight the taxonomic, biostratigraphical, and potential paleobiogeographical significance of the fauna.

STRATIGRAPHY

The Permian stratigraphy of the study area is complicated by a series of faults and folds (Fig. 1.2). In general, however, the Permian rocks form a normal sequence in that they show prom- inent internal bedding structures; they can be traced laterally over approximately 100 km along strike from northwest Zhongba

1053

1054 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

-... ] 360 Legend -m./,km

,Quaternary

SHINA Tertiary (Dazhuka Fm. - L i sandstone and volcanics)

Nagri District Tertiary (Liuqu Group -

Naqu Index Map chert beds)

Ds c Lower Tertiary (Cuojiang- Changdu District EDding Group: limestone)

SLHASA Granite and granitoids

rung-zanbo River *(Tertiary)

D c XIGAZE Linzhi District Ophiolite zones (with SXigazeDistric Shannan District ultramafics)

1 " ,Upper Cretaceous (Zengzuo

, km 225

• Fm. - shale and sandstone)

00' N Late Cretaceous (Xigaze ....

::::.:. . A Gp. - sandstone and shale)

K.%A:

e-e::::::::::: .e :::-:

::.-?-mll

Lower Cretaceous (Jiashila V . .... .Fm. - shale and limestone)

, . ....u.:

. Upper Jurassic (Weimei . . :: .. ...::..:..::...••

•'.-t.-.-... .-.... Fm. - shale and sandstone)

.-.---. .. Upper Triassic (Xiukang ,m ,,-,"--?r--

-- Gp. - sandstone and slate)

......... ....... . - and sandstone) : :a:a:nama mmBam."es. :.r"Exotic" limestone blocks

a..(Juria'"-shan limestone

o......::::Upper:CaOther symbols

... City.//own.or.vi

age ..........I... ..

-....-.-... - ......[fl. LExoation

o estuied

s ion

s KM 20 River

. . . . . . . ". . a

nd-"- .)m i

. . . . . . . . . . .. .

masses? ...j r sreaed..... ...

29?00• ? ~?- ~ "xti"lmetn bok

FIGURE I1-Sketch map showing the study area and fossil locality.

County to Taji of northwest Sage and in the study area exhibit depositional, rather than structural or tectonic, contact relation- ships with adjacent Carboniferous and Triassic rocks (Geological Survey of Tibet, 1983) (Fig. 1.2). In the study area, the Permian is represented by the Juripu Formation (Geological Survey of Ti- bet, 1983), which occurs on both limbs of an anticline, overlying

disconformably on the Carboniferous slate and siltstone and par- aconformably overlain by the Triassic Qiongguo Group (Figs. 1, 2). According to the Geological Survey Team of Tibet (1983), who surveyed the area and measured the studied section, the Juripu Formation is divisible into 12 units. These units are de- scribed below in descending order (see also Fig. 2):

SHI ET AL.-PERMIAN BRA CHIOPODS FROM TIBET 1055

Lower Trias.

12 -a Cyclolobus fauna

S (Collections (8AH334, AH338, 0 * O * 0 O.*

.. ?O. ___ AH343, AH344, eC/) Q es CeO U

_

.__ •.

e

AH345) 4

_o__ • •

-- C

'0

o---

r0 a)

Limestone lens

=9 }

Cherty limestone

? .= I• Dolomitic limestone

f,..

(Collections AH332, Siltstone

8" AH333, AH368)

--• •t"IQ'•'• 15'

•

Mudstone and shale

"• •• •.

Thin-bedded limestone

10 __0

"• "•"• , L~ lThick-bedded

____ U)

L4 imestone lens

-Para

Cherty limestone

SDisconforDolomitic limestone

3 g 2Parafuslina •F

un owNature

of contact - fauna ukon

F(Collections AH332, Silts interval dominated

AH333, AH368) ~ @ M

by ammstonite and shale

3t ? % , k T'n-bedded limestone

S10 m Fossil interva dominatedded

by bParachiopodsrmity

IIZ Disconformity

2 Parafuslina Nature of contact

?. faunaunow

Fossil interval dominated Upoby ammonites

100 M Fo ssil interva dominated Fbybrachiopods

Uppe ::...::Fossil interval dominated UpperIby fusulinaceans Carb.L 0

FIGURE 2-Stratigraphic column of the Permian Juripu Formation at the studied section, also showing the stratigraphic occurrences of brachio- pods, fusulinaceans, and ammonoids.

7. Grey thin-bedded bioclastic limestone, with abundant bra- chiopods, gastropods, and bryozoans. The brachiopods are described here (collections AH332, AH333, and AH368). 20 m thick.

6. Reddish thin-bedded dolomitic limetone, locally interbed- ded with greenish-black calcareous phyllite, with brachio- pods and crinoids. 15 m thick.

5. Grey to white thick-bedded dolomitic coarse-grained cross- bedded quartz sandstone. 10 m thick.

4. Whitish medium to thick-bedded limestone intercalated with irregular chert nodules, with sparse brachiopods and corals. 159 m thick.

3. Dark grey to grey thin-bedded limestone, with fusulinacean fossils: Parafusulina sp., P. laudoni Skinner and Wilde, and P. cf. alaskensis Dunbar. 82.9 m thick.

2. Grey to greenish slate and silty slate, locally intercalated with thin-bedded sandstone. Visible thickness is about 20 m.

1. Reddish thick-bedded limestone, with thin-bedded lime- stone at the base, containing some crinoid stems. About 150 m thick.

In summary, the Juripu Formation is characterized by carbonate rocks in the lower part and fine-grained clastic sediments in the upper part, 1,100 m in total thickness (Fig. 2). Shallow marine invertebrate fossils occur throughout the formation, but are mainly concentrated in four broad intervals (Fig. 2). The lower fossilif- erous interval (unit 3) is characterized by the occurrence of fu- sulinaceans dominated by species of Parafusulina. The middle fossiliferous interval (unit 7) is dominated by brachiopods, asso- ciated with minor amount of other fossils such as gastropods, bivalves, and solitary corals. The third main fossiliferous interval occurs at unit 11 and is dominated by brachiopods. The topmost fossiliferous interval occurs at unit 12, and is distinguished by an ammonite fauna containing Medlicottia sp., Propinacoceras cf. beyrichi Gemmellarro, Cyclolobus kraffti Diener, and C. zhong- baensis Sheng (Sheng, 1989).

The brachiopods described in this study were collected from two units of the Juripu Formation. The collection from unit 7 came from a 20 m thick thin-bedded bioclastic limestone (collections AH332, AH333, and AH368) (Fig. 2), and the other from a 141 m thick unit (unit 11) of silty and micaceous slate towards the top of the section (collections AH334, AH338, AH343, AH344, and AH345) (Fig. 2). Unfortunately, in both collections no detailed lay- er by layer sampling was performed by the field geologists because of time constraint and the reconnaissance nature of their field study at the time. Therefore, it is highly likely that the sampling was incomplete, selective and biased towards more fossiliferous beds and larger specimens. Also connected with the low intensity of sampling is the fact that only limited brachiopod specimens were collected and brought back to laboratories for further study. This subsequently has resulted in most species being only represented by limited material, some of which are very fragmentary.

Nevertheless, in spite of the pitfall in the sampling procedure, some observations on the depositional environments may be made based on the preservation of the collected material. The brachio- pods from the limestone unit (unit 7) are mostly preserved as shells, most with valves conjoined. Some of the shells have been recrystallized. The abundance of conjoined shells in this unit in- dicates that the fossils suffered insignificant postmortem trans- portation before burial. By contrast, brachiopods from unit 11 are mostly preserved as internal or external molds, many of which are still with both valves conjoined and exhibiting well-preserved imprints of external ornamentations (such as the fine ribs in spi- riferids). This mode of preservation suggests that postmortem transportation of the brachiopod assemblage was insignificant (if any), and that the original shell material of the brachiopods was leached away after the burial of the fossils.

12. Grey silty slate and calcareous slate, with ironstone and calcareous nodules. Nodules contain abundant ammonites and some brachiopods, bivalves, and gastropods. The am- monite fauna has been described by Sheng (1989) (see be- low for more details). 34.64 m thick.

11. Greenish silty, micaceous slate, with abundant brachiopods, bivalves, gastropods, and crinoid stems. The unit is capped by a 10-15 cm bed of bioclastic limestone. The brachiopods from this unit are described in this study (collections AH334, AH343, AH344, and AH345). 141.42 m thick.

10. Grey-greenish calcareous slate and silty slate, intercalated with grey-brown bioclastic limestone and muddy limestone lenses in the middle part. Both the slates and limestones con- tain fragments of brachiopods and corals. 212.13 m thick.

9. Grey to black slate and phyllitic slate. 247.49 m thick. 8. Grey slate interbedded with grey thin-bedded bioclastic

limestone, with brachiopods. 30 m thick.

1056 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

DISCUSSION OF AGE

As listed in Figure 2, the brachiopods from units 7 and 11 comprise 16 species belonging to 11 genera and three genera and species indeterminate. Although the two units are separated strat- igraphically by some 490 m strata (Fig. 2), they nevertheless share four key species: Costiferina indica (Waagen, 1884), Neospirifer (Neospirifer) kubeiensis Ting (=Ding in present spelling), 1962, Retimarginifera sp., and Stenoscisma purdoni (Davidson, 1862). Given the differences in preservation mode and associated lithol- ogy and in species composition, and a significant interval of strati- graphic separation, it is highly likely that the two units may prove to contain two distinctive brachiopod assemblages of somewhat different ages. The two brachiopod assemblages as a whole, how- ever, appear to suggest a broadly comparable age, spanning Guad- alupian (Middle Permian) to Lopingian (Late Permian) age, al- though a narrower range, from Capitanian to Wuchiapingian, is considered most likely as discussed below. Therefore, considering the limited material available for our study and the joint occur- rence of four species, we treat the two brachiopod assemblages below as a single fauna for the purpose of discussing their ages, but we do not rule out the possibility that two potential brachio- pod assemblage zones may be involved in this fauna.

Of the 16 species examined here, eight have been identifiable to species level, including one new species. Costiferina indica and Sten- oscisma purdoni both have been recorded from the Kalabagh Mem- ber in the topmost part of the Wargal Formation of the Salt Range, Pakistan. According to Wardlaw and Pogue (1995) and Wardlaw and Mei (1999), the Lopingian strata started at approximately one-third up in the Wargal Formation in the Salt Range as indicated by the occurrence of advanced Clarkina and Iranognathus erwini Mei and Wardlaw. Clarkina longicuspidata Meil, Jin and Wardlaw occurs in the lower half of the Chhidru Formation, probably marking the Wuchiapingian/Changhsingian boundary. Spiriferella qubuensis Chang (=Zhang in present spelling) (in Zhang and Jin, 1976), Neos- pirifer (Neospirifer) kubeiensis, and Neospirifer (Quadrospira) tibe- tensis Ting (=Ding in present spelling), 1962 are the most common and characteristic species of the Qubuerga Formation in the Mt. Qomolangma region and the Selong Group of the Selong Xishan section in southern Tibet. The Qubuerga Formation and the Selong Group have been both dated as largely Wuchiapingian by brachio- pods (Shen et al., 2000, 2001b). Outside Tibet, these three species are also present in the Senja Formation in northwest Nepal, which is considered to be of mostly Changhsingian in age (Waterhouse, 1978; Shen et al., 2001b).

Quinquenella glabra Waterhouse, 1975 was first recorded from the Nambo Member of the Senja Formation in northwest Nepal, considered to be Changhsingian by Waterhouse (1978). Transen- natia gratiosa (Waagen, 1884) is one of the most common species in the Wuchiapingian in the Salt Range, Pakistan (Waagen, 1884), South China (Zhan in Hou et al., 1979; Liao, 1980) and Trans- caucasia (Likharev, 1937).

Other species from the two Juripu brachiopod assemblages also support a general Late Permian and most likely Wuchiapingian age. Retimarginifera sp. can be compared closely with Retimar- ginifera xizangensis Shen, Archbold, Shi, and Chen, 2000 from the Selong Group at the Selong Xishan section (Shen et al., 2000) and the Qubuerga Formation at the Qubu section (Shen et al., 2003b). Both the Selong Group and the Qubuerga Formation have been considered to be Lopingian in age in view of the constraints from the conodonts, ammonoids, and fusulinaceans found either in direct association with or immediately above the brachiopod zone in the peri-Gondwanan region.

Although Compressoproductus cannot be identified to species level due to insufficient material, this genus is very common in the Lopingian of South China (Zhan in Hou et al., 1979) and

Transcaucasia (Ruzhentsev and Sarytcheva, 1965). It is also abun- dant in the Wuchiapingian Kalabagh Member of the Wargal For- mation in the Salt Range, Pakistan (Waagen, 1883; Reed, 1944).

Our proposition of a general late Middle Permian to early Late Permian age for the Juripu Formation brachiopods is consistent with the age indications inferred from the fusulinaceans and am- monoids found in the same section. As mentioned above, a fusu- linacean fauna dominated by Parafusulina species occurs in the lower part (unit 3) of the Juripu Formation, stratigraphically below the brachiopod assemblages (Fig. 2). According to Wang et al. (1981) and Nie and Song (in Yang and Nie, 1990), similar Para- fusulina faunas are also known from the Nagri District of south- western Tibet, where they are found in association with the Mon- odiexodina Zone. In the Salt Range of Pakistan, the Monodiexodina fauna of the Amb Formation occurs with Wordian (Middle Guad- alupian) conodonts (Wardlaw and Pogue, 1995). A comparable Monodiexodina and Parafusulina association has recently also been reported from the Zhesi Formation of Inner Mongolia in northeast China by Leven et al. (2001), who assigned it to the Kubergandian (Roadian). This age determination is consistent with the general age range of Parafusulina, from the Kubergandian (Roadian) to

early Midian (Wordian) (Leven, 1993). In view of the evidence from the fusulinaceans, it therefore appears that the lower age limit of at least the lower brachiopod assemblage from unit 7 may extend as low as the Wordian (or even Roadian).

The upper age limit of the Juripu brachiopod assemblages is constrained by the distinct Cyclolobus ammonoid-bearing unit that occurs immediately and conformably above the brachiopod fauna (Fig. 2) (Sheng, 1989). Globally, Cyclolobus has rather re- stricted spatial and temporal distributions and generally indicates a late Wuchiapingian to early Changhsingian age as constrained by the associated conodont zones in the Salt Range, Pakistan (Wardlaw and Mei, 1999). Brachiopods are present in the am- monoid-bearing unit according to the Geological Survey of Tibet (1983), but they were not extensively sampled and no brachiopods from this unit were available for this study.

The two brachiopod assemblages as a whole are closest to those in the Kalabagh Member of the Wargal Formation in the Salt Range, Pakistan in overall faunal composition. Transennatia gratiosa, Cos- tiferina indica, and Stenoscisma purdoni are also recorded in the Kalabagh Member, and eight out of the 11 genera recorded here are also present in the Kalabagh Member in the Salt Range.

In summary, therefore, taking into account of the constraints set by both the Parafusulina fauna and the Cyclolobus fauna, it is possible that the two brachiopod assemblages, respectively from units 7 and 11 of the Juripu Formation, span Capitanian (late Guadalupian) to Wuchiapingian (early Lopingian) in age, although an extension to as low as Roadian cannot be ruled out at present. This broad age range for the two brachiopod assem- blages also highlights the possibility that they may represent two brachiopod zones of slightly different ages.

SYSTEMATIC PALEONTOLOGY

The systematic classification schemes adopted here follows Carter et al. (1994) for Spiriferida, Brunton et al. (2000) for Prod- uctida, Racheboeuf (2000) for Chonetidina, Williams and Harper (2000) for Orthida, and Savage et al. (2002) for Rhynchonellida. Taxonomic classifications above family level are not listed. All

specimens figured in this paper are registered and housed in Mu- seum Victoria, Melbourne, Australia, prefixed with NMV P reg- istration numbers.

Family ENTELETIDAE Waagen, 1884 Genus ENTELETES Fischer de Waldheim, 1825

ENTELETES sp. Figure 3.1

Discussion.--A conjoined shell (NMV P308043) from collec- tion AH368 of unit 7 indicates the presence of Enteletes. The

SHI ET AL.-PERMIAN BRA CHIOPODS FROM TIBET 1057

shell possesses a sulcus on the ventral valve and a corresponding fold on the dorsal valve and several weak plications on flanks. The surface of both valves is finely costellate. Details of internal structures not observed. The present shell is somewhat similar to E. tschernyscheffi Diener (1897, p. 67, pl. 5, figs. 7-10) from the Chitichun Limestone in southwestern Tibet in shell size and pro- file, but differs from the latter by its plications that commence much later and are only prominent near the anterior margin.

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

Genus QUINQUENELLA Waterhouse, 1975 QUINQUENELLA GLABRA Waterhouse, 1975

Figure 3.2-3.6

Quinquenella glabra WATERHOUSE, 1975, p. 3, pl. 1, figs. 1-3; WATER- HOUSE, 1978, p. 107, pl. 21, figs. 5-8.

Description.-Shell relatively large for genus, subquadrate in outline, widest at hinge; cardinal extremities acute, with an angle of about 70 degrees. Ventral valve moderately convex, maximum convexity at midvalve; moderately inclined laterally; ears gently convex, not well differentiated from visceral disk; no traces of hinge spines observed from the internal molds. Ventral interior with prominent median septum; median septum thick and high posteri- orly, extending anteriorly to about half of shell length, then becom- ing a median ridge. Dorsal interior with inconspicuous inner socket ridges; lateral septa thick and short; accessory septa well developed, very long, beginning from between lateral septa to near anterior margin; inner surface of dorsal valve radially papillose.

Material examined.-Six internal molds of ventral valves (NMV P308044-308049) and an internal mold of a dorsal valve (NMV P308050), all from collection AH334 of unit 11.

Discussion.-The present specimens are slightly larger than the types figured by Waterhouse (1975), but otherwise are compara- ble. Specimens from the Kuwa Member of the Senja Formation in northwest Nepal described as Glabrichonetina kuwaensis by Waterhouse (1978) also belong to Quinquenella (Shen et al., 200 1b), but differ from the present species in its smaller size and blunt cardinal extremities. Quinquenella semiglobosa Shen et al. (2001b, p. 277, fig. 3.1-3.9) from the Qubuerga Formation of the Mt. Qomolangma region differs from the present species in its more convex ventral valve, relatively small size and subquadrate cardinal extremities.

?QUINQUENELLA sp. Figure 3.7

Discussion.-An internal mold of a ventral valve (NMV P308051) from collection AH334 of unit 11 probably indicates another species of Quinquenella. The ventral valve is 6.4 mm long and 11.7 mm wide and has a much more transverse outline than the species described above. The ventral sulcus is absent, like in the above species, but the middle portion of the ventral valve is not highly raised. The ventral interior has a short median septum which extends anteriorly as a median ridge, like in Q. glabra.

Family AULOSTEGIDAE Muir-Wood and Cooper, 1960 Subfamily AULOSTEGINAE Muir-Wood and Cooper, 1960

Genus TAENIOTHAERUS Whitehouse, 1928

Type species'-Productus

subquadratus Morris, 1845, p. 284. Discussion.-The spine pattern of Taeniothaerus is of partic-

ular note. As diagnosed by Brunton et al. (2000, p. 590), the genus has two series of spines: relatively fine spines densely scattered on both valves, and clusters of coarser spines on posterior flanks. The closest ally is perhaps Reedoconcha Kotlyar, 1964, with its thick shell, dense spines, and low interarea, but the latter is clearly

distinguishable by its elongated spine ridges that may mimic rib- bing on ventral surface. The new species described below bears two types of spines: slender ones on umbo, but comparatively coarser spines appear on lateral flanks. These spine bases, al- though swollen and gently elongated in some cases, do not form elongate ridges as in Reedoconcha.

TAENIOTHAERUS ZHONGBAENSIS new species Figure 4.1-4.8

Diagnosis.-Large, subquadrate, strongly convex Taeniothae- rus with comparatively coarse spines on lateral flanks about 1-2 mm in diameter.

Description.-Shell huge, subquadrate in outline, greatest shell width at anterior one-third of shell length; hinge straight; ears small, sharply inclined; poorly differentiated from visceral region. Ventral valve strongly convex in lateral and anterior profiles; beak thick, pointed, extending slightly beyond hinge line; interarea short and narrow (relative to hinge line); sulcus commencing from umbo, shallow on visceral region, remaining inconspicuous on trail; umbonal and lateral slopes sharply inclined. Dorsal valve moderately to deeply concave; fold gently narrow and elevated. Ventral surface with dense spine bases; spine bases slightly elon- gated and swollen anteriorly, but not forming elongated ridges, 4-5 in 5 mm, about 2.5 mm long and 1 mm in diameter on umbo, becoming 2-3 in 5 mm, about 4 mm long and 1-2 mm in diameter on lateral flanks, quincunx in arrangement; dorsal valve with nu- merous distinct dimples; spines not observed. Shell greatly thick- ened posteriorly. Details of internal structures not exposed.

Etymology.-Zhongba, a county name of southern Tibet, refers to the type locality of the species.

Types.-Holotype, a ventral valve (NMV P308052, Fig. 4.7, 4.8); paratypes, a ventral valve (NMV P308053), an incomplete ventral valve (NMV P308054), and an external mold of a dorsal valve (NMV P308055). All specimens are from collection AH333 of unit 7.

Discussion.--Compared with the new species, Taeniothaerus cf.

subquadratus (Morris) of Zhang and Jin (1976) from the Qubuerga Formation at the Qubu section in the Qomolangma region, southern Tibet, is smaller, and its spine bases on the ventral valve are also much finer. A specimen figured as Buxtonia sp., also from the Qub- uerga Formation at the Qubu section, by Ding (1962, pl. 4, figs. 1, 2) is probably a Taeniothaerus in view of its adductor muscle scars in the ventral valve, but its spine pattern is unknown. Taeniothaerus subquadratus (Morris, 1845, p. 284; Parfrey, 1983, figs. 3, 4) from the Artinskian of Tasmania is comparable to the present species in its coarse spines (0.75-1 mm in diameter) on the ventral valve, but is smaller and less convex compared to our material.

Family MONTICULIFERIDAE Muir-Wood and Cooper, 1960 Subfamily COMPRESSOPRODUCTINAE Jin and Hu, 1978

Genus COMPRESSOPRODUCTUS Sarytcheva in Sarytcheva, Likharev, and Sokolskaja, 1960

COMPRESSOPRODUCTUS sp.

Figure 3.8

Discussion.-An incomplete ventral valve (NMV P308056) from collection AH368 of unit 7 can be assigned to Compresso- productus in view of its triangular outline, distinctive rugae and ribs numbering 16 in 5 mm at the anterior margin. The present specimen is closely similar to those described by Reed (1944, pl. 11, fig. 6) as C. compressus var. subcircularis from the Kalabagh Member of the Wargal Formation in the Salt Range, Pakistan in shell outline, size and rugae, but the Zhongba specimen has more and finer ribs. C. mongolicus Diener (1897, pl. 4, figs. 8-10) from the Chitichun Limestone also bears some resemblance to the pre- sent species in its outline and size; but the ribs of the Chitichun species are very fine with rather wide interspaces, hence much

1058 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

6 7 S8 9

.10 "11

zi.....

11613

15

A pk19l22

SHI ET AL. -PERMIAN BRACHIOPODS FROM TIBET 1059

like a pectinid bivalve in external appearance (Waterhouse, 1978). Compressoproductus compressus Waagen (1884, pl. 81, figs. 1- 2) differs from the present species in its much more elongated outline and coarser ribs.

Family PRODUCTELLIDAE Schuchert, 1929 Subfamily MARGINIFERIDAE Stehli, 1954

Tribe PAUCISPINIFERINI Muir-Wood and Cooper, 1960 Genus COSTIFERINA Muir-Wood and Cooper, 1960

COSTIFERINA INDICA (Waagen, 1884) Figure 3.9-3.14

Productus costatus Sowerby. DAVIDSON, 1862, p. 31, pl. 1, figs. 20, 21. Productus indicus WAAGEN, 1884, p. 687, pl. 70, figs. 1-6; pl. 71, fig.

1. DIENER, 1915, p. 66, pl. 6, fig. 15a-d. Productus spiralis Waagen. BROILI, 1916, p. 11, pl. 117, figs. 1-5. Costiferina indica (Waagen). MUIR-WOOD AND COOPER, 1960, p. 277,

pl. 95, figs. 6, 7; pl. 103, figs. 1-12; ZHANG AND JIN, 1976, p. 178, pl. 5, figs. 11, 12; pl. 6, figs. 11-13; SHIMIZU, 1981, p. 75, pl. 7, figs. 17-19; JIANG AND YAN, 1992, pl. 1, figs. 7, 8.

Dictyoclostus cf. indica (Waagen). DING, 1962, p. 456, p1. 3, fig. la, b. ?Dictyoclostus sp. indet. DING, 1962, p. 456, pl. 3, fig. 3a-d. Costiferina spiralis (Waagen). ZHANG AND JIN, 1976, p. 179, pl. 5, figs.

4, 7-10; pl. 7, fig. 2.

Description.-Shell medium in size, subquadrate in ventral out- line, hinge nearly equal to greatest shell width; ears small; car- dinal extremities nearly at right angle. Ventral valve strongly con- vex, moderately geniculated; beak broad and thick; umbonal slopes sharply inclined; sulcus broad and shallow, commencing from umbo, becoming distinct on trail. Lateral slopes strongly inclined. Dorsal valve deeply concave; strongly geniculated; dor- sal disk and ears gently concave; fold distinct, narrow on visceral region, slightly widening anteriorly on trail. Ventral visceral disk with distinct reticulation; rugae prominent on visceral region and ears; ribs originate from umbo, coarser on trail than in visceral disk, three ribs in 1 cm; nodular spine bases scattered on visceral disk, arising from reticulations of rugae and ribs, each measured about 1 mm in diameter on average.

Material examined.-Two ventral valves (NMV P308057, 308058, both from collection AH333 of unit 7) and two external molds of dorsal valves (NMV P308059, 308060, respectively from collections AH344 and AH334 in unit 11).

Discussion.-Costiferina alata Waterhouse (1966, p. 26, pl. 5, figs. 1, 4, 5; pl. 6, figs. 1-3; pl. 7, figs. 3, 5; pl. 8, fig. 4) agrees well with the present species in details of ornaments, but differs in its large ears. The specimens from the Qubuerga Formation in the Qomolangma region figured as Dictyoclostus cf. indica and Dictyoclostus sp. indet. by Ding (1962) were later assigned to Costiferina alata by Waterhouse (1966, 1978) despite the fact that they clearly do not possess the alate ears of the latter species. Costiferina spiralis (Waagen, 1884, pl. 67, fig. 6; pl. 68, fig. 3; pl. 69, figs. 1-3) from the Amb Formation of the Salt Range is closely similar to C. indica, but differs by its huge and more

transverse shell, more delicately reticulated visceral disk and more prominent ribs on trail. A specimen from the Late Permian Basleo Bed of West Timor figured as Productus spiralis by Broili (1916, pl. 117, fig. 2, non 1, 3-5) may be also a C. indica in view of its size, outline and very coarse ribs on trail. Likewise, specimens described as C. spiralis by Zhang and Jin (1976, p. 179, pl. 5, figs. 4, 7-10; pl. 7, fig. 2) are also compatible with C. indica (Zhang and Jin, 1976, p. 178, pl. 5, figs. 11, 12; pl. 6, figs. 11- 13) in all known features except for a slightly larger size.

Genus TRANSENNATIA Waterhouse, 1975

Transennatia WATERHOUSE, 1975, p. 10. Gratiosina GRANT, 1976, p. 131. Asioproductus Zhan in YANG, NIE, CHANG, AND ZHAO, 1977, p. 350

(nomen nodum). Asioproductus Zhan in Hou, ZHAN, AND CHEN, 1979, p. 85.

Type species.-Productus gratiosus Waagen, 1884, p. 691, pl. 72, figs. 3-7.

Discussion.-Gratiosina Grant (1976) was proposed using Prod- uctus gratiosus Waagen (1884) as the type species, and is therefore deemed to be an objective synonym of Transennatia Waterhouse (1975) (see also Grant, 1976, p. 131; Brunton et al., 2000, p. 447).

Asioproductus first appeared in the publication of Yang et al. (1977, p. 350), who attributed the genus to Li-pei Zhan, for ac- commodating one of the most common productid species in the Lopingian of South China, Productus gratiosus Waagen, but with- out an unequivocal designation of a type species. Subsequently, Zhan (in Hou et al., 1979) formally proposed Asioproductus with A. bellus Chan (=Zhan), 1979 as its type species. In establishing this genus, Zhan argued that A. bellus differed from Productus gratiosus Waagen in possessing a more strongly geniculated dorsal valve and, furthermore, stated that the dorsal internal details of the latter species were unknown. However, as figured by Zhan (in Hou et al., 1979), Liao (1980) and Wang et al. (1982), the degree and nature of geniculation of the dorsal valve in the South Chinese species is variable, from a well-defined geniculation to a broadly rounded curvature. As for the dorsal internal features of Productus gratiosus Waagen, these had been described in detail and clearly figured by Grant (1976, p. 132, pl. 33, fig. 23). A comparison of the dorsal features between Asioproductus bellus as figured by Zhan (in Hou et al., 1979, pl. 6, fig. 9) and Transennatia gratiosa (Waagen) from the Salt Range as figured by Grant (1976, pl. 33, fig. 23) shows little difference between the two species. We there- fore follow Brunton et al. (2000, p. 447) in considering Asioprod- uctus as a junior subjective synonym of Transennatia.

TRANSENNATIA GRATIOSA (Waagen, 1884) Figure 3.15-3.19

Productus sinuatus? Koninck. Kayser, 1883, p. 181, pl. 25, fig. 8. Productus gratiosus WAAGEN, 1884, p. 691, pl. 72, figs. 3-7; DIENER,

1897, p. 23, pl. 3, figs. 3-7; FRECH, 1911, p. 127, pl. 19, fig. 4a-b;

FIGURE 3-1, Enteletes sp.; dorsal view, NMV P308043. 2-6, Quinquenella glabra Waterhouse; 2, internal mold of a ventral valve, NMV P308049, X3; 3, internal mold of a ventral valve, NMV P308046, X3; 4, internal mold of a ventral valve, NMV P308047, X2; 5, internal mold of a ventral valve, NMV P308045, X3; 6, latex of an internal mold of a dorsal valve, NMV P308050, X2.5. 7, Quinquenella sp.; internal mold of a ventral valve, NMV P308051, X3. 8, Compressoproductus sp.; an incomplete ventral valve, NMV P308056. 9-14, Costiferina indica (Waagen); 9, external mold of a dorsal valve, NMV P308060; 10, a crushed specimen with incomplete ventral valve and external mold of dorsal valve, NMV P308058; 11-13, anterior, psotero-ventral and lateral views of a ventral valve, NMV P308057; 14, external mold of a dorsal valve, NMV P308059. 15-19, Transennatia gratiosa (Waagen); 15, anterior view of an internal mold of a ventral valve, NMV P308063, X 1.5; 16, latex of an external mold of a ventral valve, NMV P308065, X2; 17, latex of an external mold of a ventral valve, NMV P308064, X2; 18, anterior view of an internal mold of a ventral valve, NMV P308062, X 1.7; 19, internal mold of a ventral valve, NMV P308061, x 1.5. 20-23, Retimarginifera sp.; 20, 21, posterior and anterior views of an external mold of a dorsal valve, NMV P308067, X 2; 22, 23, ventral and anterior views of an internal mold of a ventral valve, NMV P308066, X 1.5. (All figures are in natural size unless otherwise illustrated).

1060 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

1V

14

2

5

? 6

3

• ~78

FIGURE 4-1-8, Taeniothaerus zhongbaensis n. sp.; 1-3, paratype, ventral, antero-ventral and lateral views of a ventral valve, NMV P308053; 4-6, paratype, lateral, ventral and dorsal views of an incomplete ventral valve, NMV P308054; 7, 8, holotype, anterior and ventral views of a ventral valve, NMV P308052. (All figures are in natural size).

CHAO, 1927, p. 44, pl. 4, figs. 6-10; GRABAU, 1934, p. 34, pl. 10, figs. 4-6.

Productus (Dictyoclostus) aff. gratiosus Waagen. HUANG, 1932, p. 32, pl. 2, fig. 3.

Productus (Dictyoclostus) cf. gratiosus Waagen. HUANG, 1932, p. 33, pl. 2, figs. 4-5.

Productus (Marginifera) gratiosus var. timorensis Hamlet. LIKHAREV, 1937, p. 118, pl. 4, figs. 11-19; pl. 9, fig. 37.

SHI ET AL.-PERMIAN BRACHIOPODS FROM TIBET 1061

Dictyoclostus gratiosus (Waagen). ZHANG AND JIN, 1961, p. 411, pl. 4, figs. 12-18; WANG, JIN, AND FANG, 1964, p. 291, pl. 45, figs. 14-19.

Asioproductus gratiosus Zhan in YANG, NI, CHEN, AND ZHAO, 1977, p. 350, pl. 140, fig. 5; FENG AND JIANG, 1978, p. 254, pl. 90, figs. 1, 2; TONG, 1978, p. 228, pl. 80, fig. 7.

Gratiosina gratiosa (Waagen). LIKHAREV AND KOTLYAR, 1978, pl. 20, fig. la, b.

Asioproductus bellus Zhan in Hou, ZHAN, AND CHEN, 1979, p. 85, pl. 6, figs. 7-13; pl. 9, figs. 8-10.

Transennatia gratiosa (Waagen). WANG, LIU, JIN, Hu, LIANG, AND LIAO, 1982, p. 214, pl. 92, figs. 6-8; pl. 103, figs. 4-9.

Description.-Shell average size for genus, subquadrate in out- line, widest at hinge, ears small and flat, well demarcated from visceral disk by grooves; cardinal extremities bluntly acute, with an angle about 70 degrees. Ventral valve strongly incurved in lat- eral profile, strongly geniculated with long trail; beak strongly in- curved, extending slightly beyond hinge; umbonal slopes sharply inclined; sulcus beginning from umbo, broad and shallow, remain- ing distinct on trail. Ventral surface with delicately reticulated or- namentation; rugae well developed on visceral disk, five rugae in 5 mm; six ribs in 5 mm on trail, ribs also present in sulcus; a pair of halteroid spines well developed on venter at geniculation, pro- jecting anteriorly; other ventral spines not observed. Ventral interior with moderately to highly raised median adductor pad.

Material examined.-Three internal molds of ventral valves (NMV P308061-308063, all from collection AH344) and two in- complete external molds of ventral valves (NMV P308064 from collection AH343, NMV P308065 from collection AH344). All specimens from unit 11.

Discussion.-Until Zhan (in Hou et al., 1979) proposed Asio- productus bellus Zhan to accommodate a relatively small, poste- riorly consipicuously reticulate marginiferid species commonly present in the Lopingian of South China, these specimens had been generally identified with Productus gratiosus Waagen from the Salt Range of Pakistan. However, as indicated above and also revealed by more recent studies by Liao (1980) and Wang et al. (1982), given the intraspecific variation of geniculation of the dorsal valve, the South Chinese species is best considered con- specific with Transennatia gratiosa (Waagen).

The specimens from the Upper Permian of North Caucasus described as Productus (Marginifera) gratiosus var. timorensis by Likharev (1937) are clearly different from those of West Timor (Hamlet, 1928) in their larger size, finer ribs and a less deep sulcus, but may be conspecific with the present species.

Genus RETIMARGINIFERA Waterhouse, 1970

Type species.-Retimarginifera perforata Waterhouse, 1970, p. 125, figs. 2.1-11, 3.

Discussion.-The close similarity between Retimarginifera and Lamnimargus Waterhouse (1975) has been noted by Shi and Shen (1997) and Shen et al. (2000). Although Lamnimargus was de- fined to possess multiple trails arising from the dorsal marginal ridge (see also Angiolini, 2001, p. 136), many known species of this genus from the Himalaya (Shi and Shen, 1997; Shen et al., 2000) and Karakorum (Angiolini in Garzanti et al., 1996) do not display this feature. Nevertheless, Retimarginifera appears to re- main distinguishable from Lamnimargus by its fewer and weaker reticulation and stronger rugae (Angiolini, 2001, p. 136). Inter- nally, as noted by Archbold (in Shen et al., 2000, p. 741), the ventral muscle scars of Lamnimargus himalayensis (Diener), type species of the genus, are deeply impressed and their adductor scar is centrally raised and posteriorly striated but essentially smooth on its lateral portions.

RETIMARGINIFERA sp. Figure 3.20-3.23

Description.-Shell small for the tribe, 14-17 mm long and about 40 mm wide, extremely transversely semi-elliptical in out- line; ears large, acute and alate, well demarcated from visceral region by grooves; widest at hinge. Ventral valve strongly in- curved in lateral profile, maximum convexity at umbo; lateral slopes moderately inclined; sulcus deep, beginning from umbo, remaining distinct on trail. Dorsal valve strongly geniculated; vis- ceral region nearly flat; fold distinct on trail. Visceral disk of both valves ornamented with delicate reticulation; concentric rugae regularly spaced, 2-3 in 2 mm on visceral disk, absent on trail; ribs interrupted by concentric rugae, absent on ears, coarser on trail, numbering two in 2 mm on trail; a pair of halteroid spines seen on venter. Ventral interior with slightly raised diductor scars and large striated adductor scars.

Material examined.-An internal mold of ventral valve (NMV P308066) from collection AH334 of unit 11, an external mold of dorsal valve (NMV P308067) from collection AH334 of unit 11, and a fragment of a ventral valve from collection AH332 in unit 7.

Discussion.-Lamnimargus himalayensis (Diener, 1899) is closely similar to the present species in ornament and size, but appears to have a longer venter and somewhat subquadrate outline. Specimens described as Retimarginifera xizangensis (Shen et al., 2000; Shen et al., 2003a) from the Selong Group at the Selong Xishan section and the Qubu section also resemble the present species, but have less distinct reticulation and a less transverse outline.

Family STENOSCISMATIDAE Oehlert, 1887 Genus STENOSCISMA Conrad, 1839

STENOSCISMA PURDONI (Davidson, 1862) Figure 5.1-5.5

Camarophoria purdoni DAVIDSON, 1862, p. 30, pl. 2, fig. 4; WAAGEN, 1883, p. 437, pl. 32, figs. 1-4; DIENER, 1897, p. 71, pl. 12, figs. 6, 8, 9; FRECH, 1911, p. 171; BROILI, 1916, p. 55, pl. 125, figs. 7-23.

Stenoscisma purdoni (Davidson). WANG, JIN, AND FANG, 1964, p. 421, pl. 70, figs. 49, 50; GRANT, 1965, p. 149, pl. 20, figs. 1-4.

Description.-Shell 25-27 mm long, about 25 mm wide and 17 mm thick, somewhat pentagonal in outline, moderately bicon- vex in lateral profile, widest at slightly anterior to shell midlength; anterior commissure moderately uniplicate. Ventral beak acute, incurved over dorsal beak; umbonal slopes sharply inclined; sul- cus beginning from anterior to umbo, shallow and widening an- teriorly, occupying about half of shell width at anterior margin. Dorsal fold raised from midvalve. Plications in sulcus beginning from umbo, but those on flanks commencing from midvalve, numbering four in sulcus and four to five pairs on flanks; dorsal fold with 5 plications; marginal stolidium not observed. Ventral interior with distinct spondylium supported by median septum. Dorsal interior not observed.

Material examined.-Two conjoined shells (NMV P308068 from collection AH334 of unit 11, NMV P308069 from collection AH332 of unit 7) and a ventral valve (NMV P308070 from col- lection AH332 of unit 7).

Discussion.-The present specimens are almost identical to those from the Salt Range as described and figured by Davidson (1862) and Waagen (1883). The specimens under the same name as the present species from West Timor (Broili, 1916) appear to have plications beginning slightly later, but this may be an intraspecific variation. The present species differs from Stenoscisma gigantea (Diener, 1897) by its relatively small size and more plications. Sten- oscisma timorensis Hayasaka and Gan (1940) and Stenoscisma se- longensis Shen and Jin (1999) differ from the present species by their smaller size and fewer but stronger plications.

1062 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

STENOSCISMA sp. Figure 5.6-5.9

Material examined.-Five internal molds of ventral valves (NMV P308071-308075), all from collection AH334 of unit 11.

Discussion.-These five internal molds have a similar size and outline to the above species, but with finer and more numerous plications. The ventral sulcus of these molds usually possesses five to seven plications and each flank has seven to twelve plications. Some plications bifurcate once anteriorly. The ventral interior possesses a characteristic spondylium supported by a me- dian septum. As such, these specimens may represent a new spe- cies; however, the material in hand is poorly preserved and in- sufficient to allow detailed comparisons with known species of the genus.

Family RHYNCHOPORIDAE Muir-Wood, 1955 Genus RHYNCHOPORA King, 1865

?RHYNCHOPORA sp. Figure 7.11

Discussion.-An incomplete internal mold of a ventral valve (NMV P308099) from collection AH334 of unit 11 seems to in- dicate the presence of Rhynchopora in the present fauna. The ventral valve has an elongately ovate outline, acute beak and about ten costae. No meaningful comparisons can be made for this specimen because of insufficient material.

Family SPIRIFERELLIDAE Waterhouse, 1968 Subfamily SIPRIFERELLINAE Waterhouse, 1968

Genus SPIRIFERELLA Tschernyschev, 1902 SPIRIFERELLA QUBUENSIS Chang (Zhang in present spelling) in

Zhang and Jin, 1976 Figure 5.10-5.11, 5.15-5.17

Spiriferella salteri Tschernyschev. DING, 1962, p. 455, pl. 3, fig. 2a-c. Spiriferella rajah (Salter). WATERHOUSE, 1966, p. 48, pl. 1, fig. 5; pl. 7,

figs. 1, 2, 7; pl. 11, fig. 2; WATERHOUSE, 1978, pl. 14, fig. 5; Yang, Hu, and Xiong in YANG AND NIE, 1990, pl. 25, fig. 2.

Spiriferella tibetana (Diener). WATERHOUSE, 1966, p. 52, pl. 12, fig. 4; pl. 13, fig. 1 (non pl. 2, fig. 2; pl. 9, fig. 3; pl. 13, figs. 2, 4, 5).

Spiriferella qubuensis Chang (Zhang in present spelling) in ZHANG AND JIN, 1976, p. 212, pl. 18, figs. 1-5.

Spiriferella nepalensis LEGRAND-BLAIN, 1977, p. 242, pl. 1, figs. 7, 11. Spiriferella oblata WATERHOUSE, 1978, p. 89, pl. 14, figs. 14-18; p. 135,

pl. 6, figs. 7-10; WATERHOUSE, 1983, p. 135, pl. 6, figs. 7-10.

Description.-Ventral valve 25-35 mm long and 23-31 mm wide; elongate in outline, moderately to strongly convex in pro- file; hinge slightly narrower than shell greatest width at midvalve; beak pointed, strongly incurved; interarea high, strongly concave; sulcus originating from beak; well defined by coarser boundary costae, with a slender median costa along midline, remaining nar- row and shallow until anterior margin; lateral slopes sharply in- clined. Sulcal bounding costae bifurcating once anteriorly; costae on lateral slopes simple, coarse, usually not bifurcating. Shell sur- face too abraded or weathered to reveal any micro-ornament. Shell strongly thickened posteriorly, ventral dental plates deeply embedded in secondary layer of shell in apical callosity, but ev- idently separated from valve wall anteriorly, ventral myophragm and stegidial plates not observed.

Material examined.-Four ventral valves (NMV P308076-

308079), all from collection AH333 of unit 7. Discussion.-As pointed out by Shen et al. (2000) and also

shown by the synonymy list above, specimens of this species from the Himalayan region have previously been referred to several different species, including S. qubuensis by Zhang in Zhang and Jin (1976), S. nepalensis by Legrand-Blain (1977), and S. oblata by Waterhouse (1978). However, a close examination of all these

species based on illustrations and descriptions reveals little mor- phological difference. Therefore, we regard them to be synony- mous, with S. qubuensis taking the precedence.

Spiriferella qubuensis differs from S. rajah (Salter) in its small- er size, much more elongate outline and the greatest shell width being anterior to hinge line. Angiolini (in Garzanti et al., 1996) proposed a new genus, Tintoriella, with S. rajah as the type spe- cies. However, as discussed by Shi and Shen (1997) and Shen et al. (2001a), the perceived differences between Spiriferella and Tintoriella are very much dependent on shell size and quality of preservation, and therefore cannot be considered significant for separating genera. A specimen figured as S. salteri Tschernyschew by Ding (1962) from the Qubuerga Formation at the Qubu section in the Qomolangma region is clearly comparable with S. qubuen- sis. Spiriferella salteri Tschernyschew (1902) from the Sakmarian in the Urals of Russia differs from the present species in having a much deeper sulcus and a higher interarea. Some of the speci- mens from the Senja Formation in northwest Nepal described by Waterhouse (1966, 1978) as S. tibetiana and S. rajah are also referable to the present species in terms of their simple costation, elongate outline and greatest shell width occurring at or near the midvalve.

SPIRIFERELLA sp. Figure 5.12-5.14

Discussion.-Four small poorly preserved internal molds of ventral valves (NMV P308080-308083) from collection AH334 of unit 11 indicate another species of Spiriferella judging by their simple costation, deeply impressed muscle scars and distinct ven- tral sulcus. This species can be readily distinguished from S. qub- uensis described above by its much smaller size. Small Elivina tibetana (Diener, 1897) is somewhat similar to the present spec- imens in size, but the other characters of our specimens are too poorly preserved to allow any meaningful comparison.

Family TRIGONOTRETIDAE Schuchert, 1893 Subfamily NEOSPIRIFERINAE Waterhouse, 1968

Genus NEOSPIRIFER Fredericks, 1924 Subgenus NEOSPIRIFER (NEOSPIRIFER) Fredericks, 1924

NEOSPIRIFER (NEOSPIRIFER) KUBEIENSIS Ting (Ding in present spelling), 1962

Figures 6.1-6.4, 7.1-7.2

Neospirifer moosakhailensis (Davidson). MUIR-WOOD AND OAKLEY, 1941, p. 30, pl. 2, figs. 12, 13; DING, 1962, p. 452, pl. 1, figs. 1-6.

Neospirifer kubeiensis Ting (Ding in present spelling), 1962, p. 453, pl. 2, figs. 1-3; Zhang in ZHANG AND JIN, 1976, p. 203, pl. 14, figs. 1-4, 8, 9; pl. 16, fig. 8; pl. 19, fig. 2, text-figs. 10, 11 (non pl. 15, figs. 1, 2); WATERHOUSE, 1978, p. 125, pl. 24, figs. 4-7; ZHAN AND WU, 1982, pl. 5, figs. 14, 20 (non 13); Yang, Hu, and Xiong in YANG AND NIE, 1990, pl. 25, fig. 9a, b; SHI AND SHEN, 1997, p. 51, fig. 7B-D.

Neospirifer ravana (Diener). WATERHOUSE, 1966, p. 40, pl. 9, fig. 2; pl. 10, fig. 4.

Neospirifer sp. ZHAN AND WU, 1982, pl. 5, fig. 15.

Description--Most specimens are too distorted to provide use- ful measurements, but one comparatively well preserved internal mold (NMV P308084) is measured 86.1 mm long, 118.4 mm wide, and 55 mm thick; shell somewhat transversely triangular; widest at or near hinge; cardinal extremities subangular to slightly rounded; lateral sides nearly straight. Ventral interarea broadly triangular; sulcus beginning from beak; broad and moderately deep; sulcal floor U-shaped; dorsal valve with highly elevated fold; anterior commissure strongly uniplicate. Surface plicate and costellate; three to four pairs of plications on flanks (e.g., Fig. 6.4); costae numbering about 6 in 5 mm near anterior margin, increasing anteriorly by bifurcation. Ventral interior with thick

SHI ET AL.-PERMIAN BRACHIOPODS FROM TIBET 1063

1 3 4 5

8

10 12 9 10 12

1416 17

FIGURE 5-1-5, Stenoscisma purdoni (Davidson); 1, 2, ventral and posterior views of a conjoined shell, NMV P308068; 3, 4, dorsal and ventral views of a conjoined shell, NMV P308069; 5, an incomplete ventral valve, NMV P308070. 6-9, Stenoscisma sp.; 6, internal mold of a ventral valve, showing the spondylium, NMV P308072; 7, internal mold of a ventral valve, NMV P308074; 8, internal mold of a ventral valve, NMV P308073; 9, internal mold of a ventral valve, NMV P308071. 10, 11, 15-17, Spiriferella qubuensis Zhang; 10, 11, lateral and ventral views of a ventral valve, NMV P308076; 15, ventral view of a ventral valve, NMV P308079; 16, ventral view of a ventral valve, NMV P308078; 17, ventral view of a ventral valve, NMV P308077. 12-14, Spiriferella sp.; 12, internal mold of a ventral valve, NMV P308081, X2.5; 13, internal mold of a ventral valve, NMV P308080; 14, internal mold of a ventral valve, NMV P308082, X2. (All figures X 1.5 unless otherwise illustrated).

and short dental plates; adminicula well developed; adductor scars elongate, deeply impressed.

Material examined.-Five internal molds of conjoined shells (NMV P308084-308088), all from collection AH334 of unit 11.

Discussion.-The species of Neospirifer (Neospirifer) recorded from the Himalayan region have recently been discussed in detail by Shen et al. (2001a). The present species differs from the type specimens of N. (N.) moosakhailensis (Davidson, 1862, p. 28, pl. 2, fig. 2a-c) by its large size, narrower and shallower sulcus, and finer costae and plications. Specimens from North Sikkim as- signed to N. (N.) moosakhailensis (Davidson) by Muir-Wood and

Oakley (1941) are closely comparable with the present species in all known features and are therefore considered conspecific with N. (N.) kubeiensis. Neospirifer (Quadrospira) tibetensis from the Qubuerga Formation in the Qomolangma region (Ding, 1962, p. 454) differs from the present species in its subquadrate outline and narrow hinge (narrower than greatest shell width) with atten- uated ears. Neospirifer ravana (Diener) of Waterhouse (1966) from northwest Nepal is herein considered conspecific with the present species as it shares all observable features with the latter, but differs from the type material of N. ravana in having fewer plications.

1064 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

5 1

~6

2

3 4

SHI ET AL.-PERMIAN BRACHIOPODS FROM TIBET 1065

Q1

3-?~

FIGURE 7-1, 2, Neospirifer (Neospirifer) kubeiensis Ding; 1, anterior view of an internal mold of a conjoined shell, NMV P308084; 2, internal mold of a ventral valve, NMV P308086. 3, 4, Neospirifer (Quadrospira) tibetensis Ding; ventral and posterior views of a nearly complete ventral valve, NMV P308089. 5, 6, Syringothyridinae gen. et sp. indet.; ventral and postero-dorsal views of an internal mold of a distorted ventral valve, NMV P308098. 7, Elythidae gen. et sp. indet.; ventral valve, NMV P308097; x2.5. 8-10, Martiniidae gen. et sp. indet.; 8, internal mold of a ventral valve, NMV P308091, x 1.5; 9, internal mold of a distorted ventral valve, NMV P308092, X 1.5; 10, internal mold of a ventral valve, NMV P308093, xi.5. 11, ?Rhynchopora sp.; internal mold of an incomplete ventral valve, NMV P308099, x3. (All figures are in natural size unless otherwise illustrated).

FIGURE 6-1-4, Neospirifer (Neospirifer) kubeiensis Ding; 1, 2, postero-dorsal and ventral views of an internal mold of a conjoined shell (dorsal valve below), NMV P308084; 3, 4, postero-ventral and antero-dorsal views of a crushed internal mold of a conjoined shell, NMV P308085. 5-7, Neospirifer (Quadrospira) sp.; antero-ventral, ventral and anterior views of an internal mold of a conjoined shell, NMV P308090. (All figures are in natural size).

1066 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

Subgenus NEOSPIRIFER (QUADROSPIRA) Archbold, 1997 NEOSPIRIFER (QUADROSPIRA) TIBETENSIS Ting (Ding in present

spelling), 1962 Figure 7.3, 7.4

Neospirifer tibetensis Ting (Ding in present spelling), 1962, p. 454, pl. 2, figs. 4-5; Zhang in ZHANG AND JIN, 1976, p. 203, pl. 15, figs. 1, 2; YANG AND ZHANG, 1982, p. 312, pl. 3, figs. 2-4; FANG AND FAN, 1994, p. 86, pl. 31, figs. 10-12; pl. 32, figs. 1, 2.

Discussion.-A nearly complete ventral valve (NMV P308089) from collection AH333 of unit 7 reliably indicates the presence of this species. The ventral valve is 59.8 mm long and 73.1 mm wide and possesses small attenuated cardinal extremities and an extremely extended anterior tongue. The hinge line is slightly narrower than the shell greatest width, a feature diagnostic of N. (Q.) tibetensis (Shen et al., 2001a).

Neospirifer (Qudrospira) plicatus Archbold and Thomas (1986, p. 133, figs. 5A-K, 6A-I) from the late Artinskian Madeline For- mation of Western Australia is closely similar to the present spe- cies in terms of shell outline, attenuated cardinal extremities and plication pattern, but is smaller and has a more highly elevated fastigum on the dorsal valve.

NEOSPIRIFER (QUADROSPIRA) sp. Figure 6.5-6.7

Discussion.-An nearly complete internal mold of a conjoined shell (NMV P308090) from collection AH334 of unit 11 pos- sesses distinct sulcus with an elongated anterior tongue and highly elevated fold. The hinge line is straight and slightly narrower than the shell greatest width. Flanks have weak plications and costae. This specimen most likely represents another species of Neospi- rifer (Quadrospira). It differs from N. (Q.) tibetensis Ting de- scribed above in its elongated anterior tongue and the highly el- evated dorsal fold.

Family MARTINIIDAE Waagen, 1883 Subfamily MARTINIINAE Waagen, 1883

MARTINIINAE genus and species indeterminate Figure 7.8-7.10

Discussion.-Six crushed internal molds of ventral valves (NMV P308091-308096) from collection AH338 of unit 11 sug- gest a species of Martinia McCoy, 1844 or Spinomartinia Water- house, 1968. The molds indicate that there are no plates within the ventral valve although distinct bifurcated vascular markings are present. The ventral sulcus varies from shallow to deep. Mar- tinia elegans (Diener, 1897, p. 54, pl. 8, figs. 1, 2; pl. 9, figs. 1, 2) from the Chitichun Limestone in southern Tibet is similar to the present species in shell size and outline, but further compar- ison is hampered due to the poor preservation of the present spec- imens. Spinomartinia queenslandica Waterhouse (1987, p. 40, pl. 11, figs. 10, 12-18; pl. 12, fig. 1) from the Middle Permian Flat Top Formation of southeast Bowen Basin, eastern Australia is also comparable particularly in terms of the strong, bifurcated to week- ly anastomosing vascular impressions, but the Australian species appears larger and more rounded in outline.

Family ELYTHIDAE Fredericks, 1924 ELYTHIDAE genus and species indeterminate

Figure 7.7

Discussion.--A small complete ventral valve (NMV P308097) from collection AH333 of unit 7 possesses a subovate outline, an incurved beak and delicate concentric lamellae. These features suggest a possible affinity with the family ELYTHIDAE, but a more detailed comparison with known genera of this family is not war- ranted because of insufficient material.

Family SYRINGOTHYRIDIDAE Fredericks, 1926 SYRINGOTHYRIDIDAE genus and species indeterminate

Figure 7.5, 7.6

Discussion.-An internal mold of a ventral valve (NMV P308098) from collection AH334 of unit 11 exhibits a transverse outline with angular cardinal extremities, two distinct divergent dental plates and simple prominent costae on flanks. The sulcus of the ventral valve begins from beak and rapidly deepens ante- riorly, smooth. The interarea is about 16 mm high. This ventral valve may be a species of Punctocyrtella Plodowski (1968) or an allied genus. Specimens recorded as? Punctocyrtella plodowskii from the Nambdo Member of the Senja Formation in northwest Nepal by Waterhouse (1978, p. 122, pl. 23, figs. 16, 17) bears some resemblance to the present specimen, but the Nepalese spec- imens are too poorly preserved to be compared.

ACKNOWLEDGMENTS

This study is supported by an Australian Research Council grant (to G. R. Shi). S. Shen acknowledges supports from the CAS Hundred Talents Program, the Major Basic Research Project of MST (G2000077700) of People's Republic of China and a grant from the National Science Foundation of China. The paper has been improved by reviewers E Alvarez, L. Angiolini, and technical editor J. Day.

REFERENCES

ANGIOLINI, L. 2001. Permian brachiopods from Karakorum (Pakistan). Rivista Italiana di Paleontologia e Stratigrafia, 107:307-344.

ARCHBOLD, N. W 1981. Quinquenella (Chonetidina, Brachiopoda) from the Permian of Western Australia. Journal of Paleontology, 55:204-210.

ARCHBOLD, N. W. 1997. Studies on Western Australian Permian brachio- pods 14. The fauna of the Artinskian High Cliff Sandstone, Perth Basin. Proceedings of the Royal Society of Victoria, 109:199-231.

ARCHBOLD, N. W., AND G. A. THOMAS. 1986. Neospirifer and Trigon- otreta (Spiriferida, Brachiopoda) from the Permian of Western Austra- lia. Alcheringa, 10(2):125-161.

BROILI, F 1916. Die Permischen brachiopoden von Timor. In J. Wanner (ed.), Palaeontologie von Timor, 7, Stuttgart, 104 p.

BRUNTON, C. H. C., S. S. LAZAREV, R. E. GRANT, AND JIN YUGAN. 2000. Productida, p. 424-609. In R. Kaesler (ed.), Treatise on Inver- tebrate Paleontology, Pt. H, Brachiopoda Revised, Volume 3. Geolog- ical Society of America and University of Kansas Press, New York and Lawrence.

CARTER, J. L., J. G. JOHNSON, R. GOURVENNEC, AND Hou HONGFEI. 1994. A revised classification of the spiriferid brachiopods. Annals of Carnegie Museum, 68:327-374.

CHAO, Y. T. 1927. Productidae of China, Pt. 1, Producti. Palaeontologica Sinica, series B, volume 5, Fascicle 2:1-133.

CONRAD, T. A. 1839. Second annual report on the palaeontological de- partment of the survey. New York Geological Survey, Annual Report, 3:57-66.

DAVIDSON, T. 1862. On some Carboniferous Brachiopoda collected in India by A. Fleming, M.D. and W. Purdon, Esq., EG.S. Quarterly Jour- nal of the Geological Society of London, 18:25-35.

DIENER, C. 1897. Himalayan fossils. The Permocarboniferous fauna of Chitichun, No.1. Geological Survey of India, Memoirs, Palaeontologia Indica, Series 15, 1(3):1-105.

DIENER, C. 1899. Himalayan fossils. Anthracolithic fossils of Kashmir and Spiti. Memoirs of the Geological Survey of India, Palaeontologia Indica, Series 15, 1(2):1-95.

DIENER, C. 1903. Permian fossils of the Central Himalayas. Memoirs of the Geological Survey of India, Palaeontologia Indica, Series 15, 1(5): 1-204.

DIENER, C. 1915. The Anthracolithic faunae of Kashmir, Kanaur and Spiti. Memoirs of the Geological Survey of India, Palaeontologia Indica, New Series 5, 2:1-135.

DING (TING) PEICHEN. 1962. Some Upper Permian brachiopods from Tibet. Acta Palaeontologica Sinica, 10(4):451-471. (In Chinese)

SHI ET AL.--PERMIAN BRACHIOPODS FROM TIBET 1067

FANG RENSEN, AND FAN JIANCAI. 1994. Middle to Upper Carboniferous- Early Permian Gondwana facies and palaeontology in Western Yunana. Yunnan Science and Technology Press, Kunming, 121 p.

FENG RULING, AND JIANG ZONGRONG. 1978. Brachiopoda, p. 231-305. In Guizhou Working Group of Stratigraphy and Paleontology (ed.), Paleontological Atlas of Southwest China. Guizhou Volume. Pt. 2 (Car- boniferous to Quaternary). Geological Publishing House, Beijing. (In Chinese)

FISCHER DE WALDHEIM, G. 1825. Notice sur la Choristite, genre de Co- quilles bivalves fossiles du gouvernement de Moscou. Programme d'invitation a la Societe Imperiale des Naturalistes de Moscou, 11 p.

FRECH, E 1911. Das Obercarbon Chinas, Die Dyas, p. 1-289. In E von Richthofen (ed.), China, Volume 5.Dietrich Reimer, Berlin.

FREDERICKS, G. N. 1924. Paleontologicheskie etiudy. 2. O verkhne-ka- mennougol'nykh spiriferidakh Urala [Paleontological studies. 2. On some Upper Carboniferous spiriferids of the Urals]. Izvestiia Geolo- gicheskogo Komiteta, 38(3):295-324. (In Russian)

GARZANTI, E., L. ANGIOLINI, AND D. SCIUNNACH. 1996. The Permian Kuling Group (Spiti, Lahaul and Zanskar; NW Himalaya): sedimentary evolution during rift/drift transition and initial opening of Neo-Tethys. Rivista Italiana di Paleontologia e Stratigrafa, 102(2):175-200.

GEOLOGICAL SURVEY OF TIBET. 1983. 1:1000000 geological map of the Xigaze (H-45) and Yadong (G-45) Districts (Xizang) and explanatory notes. Bureau of Geology and Mineral Resources of the Xizang Au- tonomous Region, Lhasa, 568 p.

GRABAU, A. W. 1934. Early Permian fossils of China, I. Early Permian brachiopods, pelecypods and gastropods of Kueichow. Palaeontologica Sinica, Series B, 8(3):1-168.

GRANT, R. E. 1965. The brachiopod Superfamily Stenoscismatacea. Smithsonian Miscellaneous Collections, 148(2):1-192.

GRANT, R. E. 1976. Permian brachiopods from Southern Thailand. The Paleontological Society Memoir 9 [(supplement to Journal of Paleon- tology, 50(3)], 269 p.

Guo TIEYING, LIANG DINGYI, ZHANG YIZHI, AND ZHAO CHONGHE. 1991. Geology of Ngari, Tibet (Xizang). China University of Geosciences Press, Wuhan, 464 p. (In Chinese)

HAMLET, B. 1928. Permische Brachiopoden, Lamellibranchiaten und Gastropoden von Timor. Jaarboek van het Mijnwezen in Nederlandsch Oost-Indie, Gravenhage, 56(2):1-109.

HAYASAKA, I., AND S. GAN. 1940. A note on Camarophoria 'purdoni' from the Permian of Timor. Journal of the Geological Society of Japan, 47(558):127-132.

Hou HONGFEI, ZHAN LIPEI, AND CHEN BINWEI. 1979. The Coal-bearing Strata and Fossils of the Late Permian from Guangdong. Geological Publishing House, Beijing, 116 p. (In Chinese)

HUANG, T. K. 1932. Late Permian brachiopod fossils of southwestern China. Palaeontologica Sinica, Series B, 9(1):1-138.

JIANG JIANJUN, AND YAN HAIJUN. 1992. On the characteristics, members, geological age and geographical distribution of Stereochia and Costi- ferina (Brachiopoda). Journal of China University of Geosciences, 6(2): 174-182. (In Chinese)

JIN YUGAN, AND HU SHIZHONG. 1978. Brachiopods of the Kuhfeng For- mation in South Anhui and Nanking Hills. Acta Palaeontologica Sinica, 17(2):101-127.

KING, W 1865. Remarks on the histology of two specimens of Rhyncho- pora geinitziana de Verneuil, from near the River Oukhla, Province of Archangel, and St. Petersburg. Annals and Magazine of Natural His- tory, series 3, 16:124-128.

KOTLYAR, G. V. 1964. Novye predstaviteli roda Productus iz verkhnei permi iuzhnogo Primor'ia [New representative of the genus Productus from the Upper Permian of South Primorye]. Vsesoiuznyi Nauchno- Issledovatel'skii Geologicheskii Institut (VSEGEI), Trudy, 93:123-134.

LEGRAND-BLAIN, M. 1977 (1976). Repartition du groupe de Spiriferella rajah (Salter, 1865) et description de Spiriferella nepalensis, nov. sp., appartenant a ce groupe. Centre National de la Recherche Scientifique, Colloques Internationaux, France, 268(2):237-250.

LEVEN, E. YA. 1993. Main events in Permian history of the Tethys and fusulinids. Stratigraphy and Geological Correlation, 1(1):51-65.

LEVEN, E. YA., T. S. GRUNT, LIN JING-DANG, AND LI LIANG-FANG. 2001.

Upper Permian stratigraphy of the Zhesi Honguer area (North China). Stratigraphy and Geological Correlation, 9(5):441-453.

LIAO ZHUOTING. 1980. Upper Permian brachiopods from western Guizh- ou, p. 241-277. In Nanjing Institute of Geology and Paleontology (ed.),

Stratigraphy and Paleontology of the Upper Permian Coal-bearing For- mation in Western Guizhou and Eastern Yunnan. Science Press, Bei- jing. (In Chinese)

LIKHAREV (=LICHAREW), B. K. 1937 (1936). Permskie brachiopody Sev- ernogo Kavkaza. Semeustva Chonetidae Hall et Clarke i Productidae Gray [Brachiopoda of the Permian System of the USSR. Fascicule 1. Permian Brachiopoda of North Caucasus: Families Chonetidae Hall & Clarke and Productidae Gray] Monografii po Palaeontologii SSSR, 39: 1-152. (In Russian)

LIKHAREV (=LICHAREW), B. K., AND G. V. KOTLYAR. 1978. Permskie brakhiopody iuzhnogo Primor'ia [Permian brachiopods of South Pri- morye], p. 63-75, 96-99. In L. I. Popeko (ed.), Verkhnii Paleozoi Sev- ero-Vostochnoi Azii [Upper Paleozoic of Northeast Asia]. Akademiya Nauk SSSR Dal'nevostochnyie Nauchnyie Tsentr Institut Tektoniki i Geofiziki, Vladivostok.

McCoY, E 1844. A Synopsis of the Characters of the Carboniferous Limestone Fossils of Ireland. Williams & Norgate, London, 207 p.

MORRIS, J. 1845. Account of the fossil Mollusca, p. 270-291. In P. E. de Strzelecki (ed.), Physical Description of New South Wales and Van Diemen's Land. Longman, Brown,Green and Longmans, London.

MUIR-WOOD, H. M. 1955. A history of the classification of the phylum Brachiopoda. British Museum (Natural History), Monograph, viii + 124 p.

MUIR-WOOD, H. M. 1962. On the morphology and classification of the brachiopod Suborder Chonetoidea. British Museum (Natural History), Monograph, viii + 132 p.

MUIR-WOOD, H. M., AND G. A. COOPER. 1960. Morphology, classifica- tion and life habits of the Productoidea (Brachiopoda). Memoir of the Geological Society of America, 81:1-447.

MUIR-WOOD, H. M., AND K. P. OAKLEY. 1941. Upper Palaeozoic faunas of North Sikkim. Memoirs of Geological Survey of India, Palaeonto- logia Indica, 31(1):1-91.

OEHLERT, D. P. 1887. Appendice sur les brachiopodes, p. 1189-1334. In P. Fischer (ed.), Manuel de conchyliologie et de paleontologie conchy- liologue, ou histoire naturelle des mollusques vivants et fossiles, vol. 4, part 11. ESavy, Paris.

PARFREY, S. M. 1983. Redescription of the Permian aulostegid brachio- pod Taeniothaerus subquadratus (Morris, 1845), the type species of Taeniothaerus Whitehouse. Memoir of the Association of Australasian Palaeontologists, 1:289-296.

PLODOWSKI, G. 1968. Neue spiriferen aus Afghanistan. Senckenbergiana Lethaea, 49(2-3):251-258.

RACHEBOEUF, P. R. 2000. Chonetidina, p. 362-423. In R. Kaesler (ed.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda Revised, Volume 2. Geological Society of America and University of Kansas Press, New York and Lawrence.

REED, E R. C. 1944. Brachiopoda and Mollusca from the Productus Limestones of the Salt Range. Memoirs of the Geological Survey of India, Palaeontologia Indica, 23(2):1-678.

RUZHENTSEV, V. E., AND T. G. SARYTCHEVA. 1965. Razivitie i smena morskikh organizmov na rubezhe Paleozoia i Mezozoia [The devel- opment and change of marine organisms at the Paleozoic-Mesozoic boundary]. Akademiia Nauk SSSR, Paleontologicheskii Institut, Trudy, 108:1-370. (In Russian)

SARYTCHEVA, T. G., B. K. LIKHAREV, AND A. N. SOKOLSKAJA. 1960. Otriad Productida, p. 221-238. In Y. A. Orlov (ed.), Osnovy Paleon- tologii; Mshanki Brakhiopody. Izdatel'stvo Akademii Nauk SSSR, Moscow. (In Russian)

SAVAGE, N. M., M. O. MANCENIDO, AND E. F OWEN. 2002. Order Rhyn- chonellida, p. 1027-1376. In R. Kaesler (ed.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda Revised, Volume 4. Geological Society of America and University of Kansas Press, New York and Lawrence.

SCHUCHERT, C. 1893. A Classification of the Brachiopoda. American Geologist, 11(3):141-167.

SCHUCHERT, C. 1929. Classification of brachiopod genera, fossil and re- cent, p. 10-25. In C. Schuchert and C. M. LeVene, Animalia pars 42 [In J.E Pompeckj (ed.), Fossilium Catalogus,volume 1] Junk, Berlin.

SHEN SHUZHONG, AND JIN YUGAN. 1999. Brachiopods from the Permian- Triassic boundary beds at the Selong Xishan section, Xizang (Tibet), China. Journal of Asian Earth Sciences, 17(4):547-559.

SHEN SHUZHONG, N. W. ARCHBOLD, G. R. SHI, AND Z. Q. CHEN. 2000. Permian brachiopods from the Selong Xishan section, Xizang (Tibet),

1068 JOURNAL OF PALEONTOLOGY, V. 77, NO. 6, 2003

China, Pt. 1, Stratigraphy, Strophomenida, Productida and Rhyncho- nellida. Geobios, 33:725-752.

SHEN SHUZHONG, N. W. ARCHBOLD, G. R. SHI, AND Z. Q. CHEN. 2001 a. Permian brachiopods from the Selong Xishan section, Xizang (Tibet), China, Pt. 2, Palaeobiogeographical and palaeoecological implications, Spiriferida, Athyridida and Terebratulida. Geobios, 34:157-182.

SHEN SHUZHONG, N. W. ARCHBOLD, AND G. R. SHI. 2001b. A Lopingian (Late Permian) brachiopod fauna from the Qubuerga Formation at Shengmi in the Mount Qomolangma region of Southern Xizang (Tibet), China. Journal of Paleontology, 75:274-283.

SHEN SHUZHONG, CAO CHANGQUN, G. R. SHI, WANG XIANGDONG, MEI SHILONG, AND JIN YUGAN. 2003. Lopingian (Late Permian) stratigra- phy and deposition in southern Tibet and its correlation over the peri- Gondwana region. Newsletter of Stratigraphy, 39:157-179.

SHEN SHUZHONG, G. R. SHI, AND N. W. ARCHBOLD. 2003a. A Wuchia- pingian (Late Permian) brachiopod fauna from an exotic block in the Indus-Tsangpo Suture zone, southern Tibet and its palaeobiogeograph- ical and tectonic implications. Palaeontology, 46:225-256.

SHEN SHUZHONG, G. R. SHI, AND N. W. ARCHBOLD. 2003b. Lopingian (Late Permian) brachiopods from the Qubuerga Formation at the Qubu section in the Mt. Qomolongma region, southern Tibet (Xizang), China. Palaeontographica Abt. A, 268:49-101.

SHEN SHUZHONG, SUN DONGLI, AND G. R. SHI. 2003c. A Capitanian (late Middle Permian) brachiopod fauna from an exotic limestone block at Xiukang in Lhaze County of Tibet. Journal of Asian Earth Sciences, 21:1125-1137.

SHENG HUAIBIN. 1989. Early Permian ammonoids from the Juripu For- mation, Zhonga, Xizang (Tibet). Geological Review, 35:324-332. (In Chinese)

SHI, G. R., AND SHEN SHUZHONG. 1997. A Late Permian brachiopod fauna from Selong, Southern Xizang (Tibet), China. Proceedings of the Royal Society of Victoria, 109(1):37-56.

SHIMIZU, D. 1981. Upper Permian brachiopod fossils from Guryul Ravine and its spur three kilometers of Barus. Palaeontologica Indica, New Series, 46:67-85.

STEHLI, E G. 1954. Lower Leonardian Brachiopoda of the Sierra Diablo. Bulletin of the American Museum of Natural History, 105(3):257-358.

TONG ZHENGXIANG. 1978. Brachiopoda, p. 210-267. In Geological In- stitute of Southwest China (ed.), Paleontological Atlas of Southwest China, Sichuan Province, Volume 2 (Carboniferous and Permian). Geo- logical Publishing House, Beijing. (In Chinese)

TSCHERNYSCHEV, T. N. 1902. Verkhnekamennougol'nye brakhiopody Ur- ala i Timana [Upper Carboniferous brachiopods from the Urals and Timan]. Trudy Geologicheskogo Komiteta, 16(2):1-749. (In Russian)

WAAGEN, W. 1883-1884. Salt Range fossils I. Productus-Limestone fos- sils. Geological Survey of India, Memoirs, Palaeontologia Indica, Se- ries 13, facicules 2- 4:391-728 , pls. 29-81.

WANG CHENGSHAN, XIA DAIXIANG, ZHOU XIANG, AND CHEN JIANPING. 1999. Geology between the Indus-Yarlung Zangbo Suture Zone and the Himalaya Mountains, Xizang (Tibet), China. Geological Publishing House, Beijing, 160 p. (In Chinese)

WANG GUOPING, Liu QINGZHAO, JIN YUGAN, HU SHIZHONG, LIANG WENPING, AND LIAO ZHUOTING. 1982. Brachiopoda, p. 186-256. In Institute of Geology and Mineral Resources (ed.), Paleontological Atlas of East China. 2. Late Paleozoic. Geolological Publishing House, Bei- jing. (In Chinese)

WANG Yu, JIN YUGAN (CHING YU-KAN), AND FANG DAWEI. 1964. Bra- chiopod Fossils of China. Science Press, Beijing, Volumes 1 and 2, 777 p. (In Chinese)

WANG YUJING, SHENG JINZHANG, AND ZHANG LINxIN. 1981. Fusulinids from Xizang of China, p. 1-80. In Chinese Academy of Sciences In- tegrated Scientific Expedition Team to the Qinghai-Xizang (Tibet) Pla- teau (ed.), The Series of the Scientific Expedition to the Qinghai-Xiz- ang Plateau. Palaeontology of Xizang, Book III. Science Press, Beijing. (In Chinese)

WARDLAW, B. R., AND K. R. POGUE. 1995. The Permian of Pakistan, p. 215-224. In P. A. Scholle, T. M. Peryt, and D. S. Ulmer-Scholle (eds.), The Permian of Northern Pangea 2, Sedimentary Basins and Economic Resources.Springer-Verlag, Berlin.

WARDLAw, B. R., AND MEI SHI-LONG.1999. Refined conodont biostratig- raphy of the Permian and lowest Triassic of the Salt and Khizor Ranges, Pakistan, p. 154-156. In Yin Hongfu and Tong linnan (eds.), Proceedings of the International Conference on Pangea and the Paleozoic-Mesozoic Transition. China University of Geosciences Press, Wuhan, China.

WATERHOUSE, J. B. 1966. Lower Carboniferous and Upper Permian bra-

chiopods from Nepal. Geologische Bundesanstalt, Jahrbuch, 12:5-99. WATERHOUSE, J. B. 1968. The classification and descriptions of Permian

Spiriferida (Brachiopoda) from New Zealand. Palaeontographica, Abt. A, 129(1-3):1-94.

WATERHOUSE, J. B. 1970. Permian brachiopod Retimarginifera gen. nov. from the Byro Group of Carnarvon Basin, Western Australia. Proceed-

ings of the Royal Society of Western Australia, 53(4):120-128. WATERHOUSE, J. B. 1975. New Permian and Triassic brachiopod taxa.

Papers of Department of Geology, University of Queensland, 7(1):1-23. WATERHOUSE, J. B. 1978. Permian Brachiopoda and Mollusca from

North-West Nepal. Palaeontographica, Abt. A, 160(1-6):1-175. WATERHOUSE, J. B. 1983. Permian brachiopods from Pija Member, Senja

Formation in Manang district of Nepal, with new brachiopod genera and species from other regions. Indian Geologists' Association Bulle- tin, 16(2):111-151.

WATERHOUSE, J. B. 1987. Late Palaeozoic Brachiopoda (Athyrida, Spi- riferida and Terebratulida) from the southeast Bowen Basin, east Aus- tralia. Palaeontographica Abt., 196:1-56.

WHITEHOUSE, F W. 1928. Notes on Upper Palaeozoic marine horizons in eastern and western Australia. Australasian Association for the Ad- vancement of Science, Report, 18:281-283.

WILLIAMS, A., AND D. A. HARPER. 2000. Orthida, p. 714-844. In R. Kaesler (ed.), Treatise on Invertebrate Paleontology, Pt. H, Brachiopoda Revised, Volume 3. Geological Society of America and University of Kansas Press, New York and Lawrence.

YANG DELI, NI SHIZHAO, CHANG MEILI, AND ZHAO RUXUAN. 1977.

Brachiopoda, p. 303-470. In Hubei Geological Institute, and Geolog- ical Bureaus of Henan, Hubei, Hunan, Guangdong, and Guangxi Prov- inces (eds.), Paleontological Atlas of Central South China, Volume 2 (Late Paleozoic). Geological Publishing House, Beijing. (In Chinese)

YANG ZUNYI, AND NIE ZETONG. 1990. Palaeontology of Nagri, Tibet (Xiz- ang). China University of Geosciences Press, Wuhan, China, 380 p.

YANG ZUNYI, AND ZHANG KANGFU. 1982. Fossils in the upper part of the Lower Permian from Surishan, Dingri, Xizang, p. 310-315. In

Monograph on Mount Xixia Bangma Scientific Expedition 1964, Sci- ence Press, Beijing. (In Chinese)

YIN JIXIANG. 1997. Stratigraphic Geology of Gondwana Facies of Qing- hai-Xizang (Tibet) Plateau and Adjacent Areas. Geological Publishing House, Beijing, 205 p. (In Chinese)

ZHAN LIPEI, AND WU RANGYONG. 1982. Early Permian brachiopods from Xainza district, Xizang (Tibet), p. 86-109. In Contribution to the Ge-

ology of the Qinghai-Xizang (Tibet) Plateau, 7. Geological Publishing House, Beijing.

ZHANG (CHANG) SHOUXIN, AND JIN (CHING) YUGAN. 1976. Late Paleo-

zoic brachiopods from the Mount Jolmo Lungma region, p. 159-271. In Xizang Scientific expedition team of Chinese Academy of Sciences

(ed.), A Report of Scientific Expedition in the Mount Jolmo Lungma Region (1966-1968), 2, Science Press, Beijing. (In Chinese)

ZHANG (CHANG) YAN, AND JIN (CHING) YUGAN. 1961. Late Permian

brachiopod fossils from Jin Xian, Anhui province. Acata Palaeontolo-

gica Sinica, 9(4):401-414.

ACCEPTED 15 FEBRUARY 2003