middle permian brachiopods from central peninsular ...directory.umm.ac.id/data...

Middle Permian brachiopods from central Peninsular Malaysia Ð faunalaf®nities between Malaysia and west Cambodia

Masatoshi Sonea,1,*, Mohd Shafeea Lemanb, Guang R. Shia

aSchool of Ecology and Environment, Deakin University, Rusden Campus, Clayton, Vic. 3168, AustraliabSchool of Environmental Sciences and Natural Resources, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia

Accepted 27 April 2000

Abstract

A moderately diverse Permian brachiopod fauna is described from a new rock unit, the Bera Formation, in the Bera District, central

Pahang, Peninsular Malaysia. The fauna consists of 19 taxa, including 14 genera and 17 (both identi®ed and unidenti®ed) typically Tethyan

species. The fauna appears to be correlative on the basis of brachiopods with the Neoschwagerina-Yabeina fusulinid Zones in Indochina and

South China. In particular, it has strong linkage to Member C (Yabeina beds) of the Sisophon Limestone, west Cambodia. This is indicated by

three of the Bera species Ð Urushtenoidea chaoi (Ching), Spyridiophora gubleri Termier and Termier, and Transennatia termierorum sp.

nov., being shared with the Cambodian fauna. A possible early Capitanian (Middle Permian) age is proposed for the Bera brachiopod fauna.

q 2001 Elsevier Science Ltd. All rights reserved.

1. Introduction and regional geology

This paper describes the largest known Permian brachio-

pod fauna from the so-called Central Basin (or the Central

Belt) of Peninsular Malaysia, and reports on the southern-

most occurrence of such a fauna in that region. Most of the

fossils examined in this study were collected by Sone and

Leman in the Bera District in February 1998. Additional

material came from a collection in the palaeontology labora-

tory of the National University of Malaysia (Universiti

Kebangsaan Malaysia), collected originally by Leman. All

described specimens are registered in the same university

with pre®x UKM-F.

The regional geology of the Bera District is known from

only two brief reports (Cook and Suntharalingam, 1970;

MacDonald, 1970). The rock unit from which the current

fossil horizon is studied has been named the Bera Formation

(Leman et al., in press). The area where it occurs was

previously considered to be part of the Triassic Semantan

Formation Ð for example, on the most recent of®cial geolo-

gical map of Peninsular Malaysia (Geological Survey of

Malaysia, 1985). The Semantan Formation, as revised,

outcrops west of the Bera Formation, which in turn is

¯anked on the east by ?Jurassic to Cretaceous continental

sediments of the Bertangga Sandstone, but exposures to east

and west are poor (Leman et al., in press). The northern and

southern boundaries of the Bera Formation are also uncer-

tain, but in the north it is associated with an andesitic vol-

canic unit thought to be of a Late Permian age (Cook and

Suntharalingam, 1970). The occurrence of the fusulinid,

Parafusulina sp., from a small limestone outcrop near

Tasik Bera implies that the lower part of the Bera Formation

may extend down into the Kungurian (upper Early Permian)

(Cook and Suntharalingam, 1970).

2. Stratigraphy

The fossils of this report come from the ªSungai Bera

sectionº (grid reference WF151446; the National Map

Malaysia 1:50 000 Sheet 4157) exposed on the eastern

side of Bera Road, 17.3 km south of Felda Sebertak junction

(Fig. 1). This section is equivalent to Locality BF2 of

Leman et al. (in press). The outcrop was originally prepared

for construction of a resort. The exposed strata extend

approximately 90 m vertically and 300 m laterally, and

appear to have undergone gentle but pervasive soft-sedi-

ment deformation westwards. It is thus dif®cult to trace

the sequence from one side to the other. The general strike

is N50±708W and dip 70±908SW. The lithology consists

mainly of mudstone±siltstone interbedded with sandstone,

Journal of Asian Earth Sciences 19 (2001) 177±194

1367-9120/01/$ - see front matter q 2001 Elsevier Science Ltd. All rights reserved.

PII: S1367-9120(00)00026-2

www.elsevier.nl/locate/jseaes

* Corresponding author.

E-mail address: [email protected] (M. Sone).1 Present address: Institute for Environment and Development

(LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor,

Malaysia.

becoming dominantly tuffaceous in the upper part of the

section. The surface is bleached soft sediment. More

outcrops occur along the road; these are much smaller

than the Sungai Bera section.

Three major fossiliferous stratigraphic intervals can be

discriminated; these are referred to as Assemblages A±C

in ascending order (Fig. 2). Assemblage A is predominantly

a coquina of shell debris in grey mudstone; it often contains

fossiliferous calcareous concretions. This assemblage holds

the richest brachiopod deposit in this report. Assemblage B

has less common brachiopods in nearly white, ®ne- to

medium-grained sandstone. Assemblage C consists princi-

pally of abundant lyttoniids in purple to black tuffaceous

mudstone and siltstone.

The brachiopods are listed in Table 1. The genus Trans-

ennatia is the most abundant, although most specimens are

too poorly preserved for satisfactory illustration. The three

assemblages share few taxa, and may appear to represent

distinct faunules, but sampling was not exhaustive. The

three assemblages are considered as a single fauna in the

following discussion.

3. Correlation and age

At present, there is no globally accepted Permian chron-

ological standard. The time-scale compiled by the Interna-

tional Subcommision on Permian Stratigraphy (Jin et al.,

1997) is applied in this study, with slight modi®cation to

fusulinid biozonation for more appropriate correlation

within South-East Asia (Fig. 3). The brachiopod fauna of

the Sungai Bera section appears to correlate with the other

brachiopod faunas in the Neoschwagerina-Yabeina Zones

of the Middle Permian, most likely to a lower Capitanian,

This is based on the three brachiopod species shared with

Member C (Yabeina beds) of the Sisophon Limestone, west

Cambodia.

The Permian strata of west Cambodia are exposed mainly

in the Sisophon and Battambang regions (Fig. 4), as lime-

stone hills with intercalations of tuff, shale and mudstone

strata, collectively called the Sisophon Limestone (Ishii et

M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194178

Fig. 1. Possible extent of the Bera Formation and the location of the Sungai Bera section (modi®ed after Leman et al., in press).

Table 1

Brachiopod species from each assemblage of the Sungai Bera section

Species Assemblages

A B C

Derbyia sp. X

Urushtenoidea chaoi (Ching, 1963) X X

Urushtenoidea sp. X

Transennatia termierorum sp. nov. X

Transennatia sp. indet. X

Spinomarginifera sp. X

Echinauris sp. X

Spyridiophora gubleri Termier and Termier, 1970a X

Kozlowskia sp. X

Phricodothyris sp. X

Orthotetes aff. picta Fang, 1983 X

Strophalosiina sp. X

Linoproductus sp. X

Rhynchonellid family indet. X

Orthothetina cf. iljinae Sokol'skaya, 1965 X

Gubleria aff. ninglangensis Fang and Jiang, 1992 X

Gubleria sp. X

Eolyttonia? sp. X

Spiriferinid family indet. X

al., 1969). Brachiopods from the limestone have been

studied by several palaeontologists (e.g., Mansuy, 1913,

1914; Chi-Thuan, 1961; Termier and Termier, 1960,

1970a, 1970b). Ishii et al. (1969) listed 50 brachiopod

species from their collections gained from that unit; Naka-

mura (1979a) later revised the list. Based on lithology, Ishii

et al. (1969) divided the limestone into four members, A±D

in ascending order (Fig. 3). This division is supported by

fusulinid faunas, supplemented by coral and brachiopod

data. Because the four members are de®ned on lithologies

and because a few key fusulinid species range through more

than one member, the boundaries of each member cannot be

precisely assigned chronologically. The most likely biostra-

tigraphic ranges of the four members are indicated in Fig. 3,

based on studies of Ishii et al. (1969) and Waterhouse

(1976).

Member C consists of shale and calcareous mudstone

with calcareous nodules, sandwiched by the limestone

facies of Members B and D (Ishii et al., 1969). It contains

abundant brachiopods, and is characterised by fusulinid

species such as Yabeina asiatica, Lepidolina multiseptata,

Sumatrina longissima and Verbeekina verbeeki. Y. asiatica

is the primitive form of the genus, suggesting the lowest part

of the Yabeina Zone (Ishii, 1966). L. multiseptata and S.

longissima are typical species of the middle Midian (the

Y. globosa Zone) of the eastern Tethys (Leven, 1992,

1993). Species of Sumatrina and Verbeekina are not

known to extend up to the upper Midian (see Kobayashi,

1997). The Midian Stage is approximately equivalent to the

Capitanian Stage (Jin et al., 1997), although the lower

boundary of the former is probably slightly older than that

of the latter, as pointed out by Leven and Grant-Mackie

(1997). Therefore, an early Capitanian age is the most de®n-

able for Member C of the Sisophon Limestone.

The most important species in the Bera fauna is Urushte-

noidea chaoi (Ching) whose many reported occurrences are

con®ned to the Kuhfengian Stage (Wordian) of South China

and Member C of the Sisophon Limestone. It is one of the

index fossils for the lower Maokouan Sub-series in the

Nanjing Hills, southeastern China (He and Shi, 1996). An

equally important form is Spyridiophora gubleri Termier

and Termier. It has previously been found only in the

Yabeina beds of the Sisophon Limestone (Mansuy, 1913;

Chi-Thuan, 1961; Termier and Termier, 1970a). Nakamura

(1979a) also listed its occurrence in Member C. Transenna-

tia termierorum sp. nov. is elsewhere known only in the

Yabeina horizon of the Sisophon Limestone (Termier and

Termier, 1970a). These three species indicate strong linkage

between the Bera fauna and that in Member C (the Yabeina

Zone) of the Sisophon Limestone.

Species of Orthotetes, Gubleria, Phricodothyris and

Orthothetina in the Bera fauna closely resemble those

recorded in the Gnishik-Khachik beds of the Trans-Cauca-

sus and in the Maokou horizons of South China. In particu-

lar, the species of Orthotetes and Gubleria are alike to those

in west Yunnan, southwestern China. Additionally, the Bera

fauna displays some similarity to the Capitanian brachio-

pods of the Lengwu Formation, Zhejiang Province, South

China (see Liang, 1990). Almost all Bera genera occur in

the Lengwu fauna, including relatively rare taxa such as

M. Sone et al. / Journal of Asian Earth Sciences 19 (2001) 177±194 179

Fig. 2. Stratigraphic log of the Sungai Bera section with indications of each

Assemblage A to C (modi®ed after Leman et al., in press).

Strophalosiina and Spyridiophora; the latter are uncommon

or absent in the underlying and overlying Maokou and

Wuchiaping Formations. There are no species in common

between the Bera and Lengwu faunas. This may be due to

slightly different ages, or slightly different climatic condi-

tions, or both.

No genus of the Bera fauna is particularly suggestive of a

Late Permian age. Transennatia, Orthothetina and Gubleria

are regularly seen in the Wuchiapingian Stage of South

China and in the Djul®an Stage of the Trans-Caucasus.

However, some species of these genera occur in Middle

Permian rocks of South China, west Cambodia, Japan,

south Primorie and possibly Timor.

The presence of Urushtenoidea in both Assemblages A

and C is good evidence for precluding the possibility of a

Late Permian age. The genus has never been found in post-

Kuhfengian horizons in South China; its closest relative

Uncisteges persisted only into the lower part of the overlying

Lengwuan horizon; its youngest occurrence, on present data, is

in Member C of the Sisophon Limestone. Spyridiophora and

Kozlowskia are consistent with a pre-Wuchiapingian age.

These genera are dominant in the Early Permian, and have

never been unequivocally reported from Late Permian

rocks. Their youngest occurrence may in fact be in the

Lengwu Formation.


Fig. 3. Age of the Sungai Bera section brachiopods and possible correlation with other horizons in the Central Basin of Peninsular Malaysia, the Sisophon

Limestone of west Cambodia, and South China. Only part of the Early Permian is shown. Time-scale after Jin et al. (1997); fusulinid biozonation and the

stratigraphy of localities in the Central Basin adopted from Fontaine et al. (1994); Jengka Pass, Lst. � Limestone Member, S & S. � Sandstone-shale

Member; Aring Formation is partly adapted from Yanagida and Aw (1979); Sumalayang Limestone is from Igo et al. (1979); divisions of the Sisophon

Limestone are after Ishii et al. (1969).

Fig. 4. Possible distributions of the Indochina and East Malaya terranes, and

locations of the Bera District and the Sisophon and Battambang regions: 1,

Uttaradit-Nan Suture; 2, Raub-Bentong Line (modi®ed after Metcalfe,

1998).

There is a generally accepted view that large lyttoniids

such as Leptodus and Gubleria are typically Late Permian

(e.g., Shen and Shi, 1996) and, in the case of Oldhamina,

this seems to be true. It is only partly true for brachiopod

faunas from the Trans-Caucasus and South China, where

the Lengwu fauna is now known to contain numerous lytto-

niids, but notably without Oldhamina. Many lyttoniid forms

occur in the Middle Permian of Indochina (e.g., Mansuy,

1912, 1913, 1914; Termier and Termier, 1960; Chi-Thuan,

1961), and in the Neoschwagerina horizon of the Sosio

Limestone, Italy (e.g., Rudwick and Cowen, 1967). Lyt-

toniids are in fact more common in the Middle rather than

Late Permian of Japan, with several reports from the lower

Kanokuran Series (e.g., Tazawa, 1976, 1987; Tazawa and

Matsumoto, 1998). The presence of numerous lyttoniids in

Assemblage C is thus not necessarily indicative of a Late

Permian age; it may in fact be the Middle Permian.

Permian strata are prominent in the Central Basin of

Peninsular Malaysia. Ages with respect to the age of the

Sungai Bera section are indicated in Fig. 3. These strata

are mostly limestones dated by fusulinids, corals, algae

and/or radiolarians. In contrast, only a few Permian brachio-

pods have been described, despite many reported occur-

rences (e.g., Jones et al., 1966; Gobbett, 1973; Fontaine et

al., 1994). Igo (1964) and Nakamura (in Nakazawa, 1973)

described some poorly preserved brachiopods of little bio-

stratigraphic value. Yanagida and Aw (1979) described a

small suite of Permian brachiopods from Kelantan; they

considered the fauna to be late Djul®an.

The so-called ªLeptodus Shaleº fauna in north-central

Pahang has been known for over a half century (Muir-

Wood, 1948, p. 5, footnote; Jones et al., 1966). Leman

(1993, 1994) reported 49 species of brachiopods from the

same fauna, but they have not been described. Based on

the presence of abundant lyttoniids, the fauna was thought

to be Late Permian, but this is questionable. Thus, based

on brachiopods, it is still dif®cult to make precise correla-

tions within the Central Basin. Nevertheless, considering

available data from palaeontology and regional geology,

the Sungai Bera section is likely to more or less align

stratigraphically with the nearest limestone units at

Kampung Awah and Jengka Pass (Fig. 3). Both the lime-

stones yield Yabeina asiatica which is known elsewhere

only in the Sisophon Limestone (Ishii, 1966; Igo, 1967;

Ishii et al., 1969; Fontaine et al., 1994). North of the Bera

District, small outcrops of late Middle Permian age have

been studied in recent years (e.g., Fontaine et al., 1994;

Jasin et al., 1995). There are thus prospects for better

understanding of Permian correlations in central Pahang,

speci®cally with respect to the Bera brachiopod fauna.

4. Palaeogeographic implications

All taxa identi®ed to generic and speci®c levels accord

with this being a warm-water fauna. This is indicated by

Orthothetina, Transennatia, Gubleria, Spinomarginifera,

Urushtenoidea and Strophalosiina, all of which are typical

Tethyan elements. The close af®nities between the Bera and

Sisophon faunas were discussed above. Relationships with

the Middle Permian brachiopods of South China are less

obvious. Differences in Middle Permian brachiopods

between Indochina and South China have been pointed

out by Nakamura et al. (1985). It is, therefore, assumed

that the Central Basin with the Bera fauna was located in

the same or nearby climatic zone as west Cambodia during

the age-interval in question, but that there were slight differ-

ences in ecology or geographic position relative to tropical

South Chinese Cathaysia, hindering faunal interchange.

This conclusion is consistent with the tectonic model of

Metcalfe (1996a, 1996b, 1998), namely, that the East

Malaya terrane accommodating the Central Basin was part

of the Indochina terrane (Fig. 4).

5. Systematic palaeontology (M. Sone)

The systematic study follows the supra-ordinal classi®ca-

tion of Williams et al. (1996), and the classi®cations of

Williams and Brunton (1993) for the Strophomenida, Brun-

ton et al. (1995) and Brunton and Lazarev (1997) for the

Productida, and Carter et al. (1994) for the Spiriferida and

Spiriferinida. The suborder Strophalosiidina Schuchert,

1913 refers to Brunton (written communication, 27 January

2000).

Class Strophomenata Williams et al., 1996

Order Strophomenida OÈ pik, 1934

Suborder Orthotetidina Waagen, 1884

Superfamily Orthotetoidea Waagen, 1884

Family Orthotetidae Waagen, 1884

Subfamily Orthotetinae Waagen, 1884

Genus Orthotetes Fischer de Waldheim, 1829

Orthotetes aff. picta Fang, 1983

Fig. 5; 1±2.

cf. Orthotetes picta Fang, 1983, p. 94. pl. 1, ®gs. 4±6.

Description. Ventral shell medium for genus, 27 mm wide

and 26.5 mm long with greatest width at hinge; outline U-

shaped; pro®le fairly ¯at; ears not demarcated; interarea

very low; umbo inconspicuous; rectimarginate; plications

absent. Ornament of moderate costellae with angular crests,

increasing by intercalation, numbering approximately 18

per 10 mm at anterior margin; growth lines few, well

de®ned, with two high ones at mid-length and near the

anterior margin, with the anteriormost one extending from

the end of the hinge. Ventral interior with long median

septum, extending to nearly mid-valve, slightly thickened

anteriorly, height unknown; spondylium minute, laterally

semi-ovate; dental ridges uniting to form a spondylium;


adductor ®eld weakly concave. Details of dorsal valve

unknown.

Remarks. The ¯attened ventral shell with a small apical

spondylium and a median septum suggests Orthotetes. Its

U-shaped outline, low in¯ation and rectimarginate commis-

sure makes it close to O. picta Fang (1983, p. 94. pl. 1, ®gs.

4±6) from the Neoschwagerina horizon of the Xiaoxinzhai

section in Gengma, west Yunnan. There are de®nable, but

subtle differences in the condition of the costellation Ð the

Yunnan species displays slightly coarser (7±8 in 5 mm) and

more intercalated costellae. This, however, may be due to

differing stages of their maturity; the larger Bera valve

seems older ontogenetically with more growth lines. It is

thought to be close to O. picta, but additional material is

required for con®dent identi®cation. No other previously

described species appears close to the present form.

Family Derbyiidae Stehli, 1954

Subfamily Derbyiinae Stehli, 1954

Genus Derbyia Waagen, 1884

Derbyia sp.

Fig. 5; 3±4

Remarks. The ventral shell with a high and thick median

septum bisecting the umbo is indicative of Derbyia, but it is

small for the genus, almost equidimensional (about 23 mm

wide to 22 mm long), and the hingeline is the widest part.

The outline is U-shaped, the ears are not differentiated, and

the pro®le is convex in the median region, but contrastingly

¯at on the ¯anks. The interarea is low and narrow, and

cardinal extremities are obtuse. The umbo is prominent.

The shell is rectimarginate without plications, and is undis-

torted. The septum is relatively short. Costae are coarse, low

and blunt, increasing by intercalation, with width fairly

uneven.

The present species is characterised by the umbonal

region being prominent, raised and demarcated by a strong

concentric line. D. altestriata Waagen illustrated by Fantini

Sestini (1965, p. 38, pl. 3, ®g. 7) from the upper Ruteh

Limestone, north Iran, displays a very similar elevated

umbo with a primary lamina. It differs from the Bera form

in possessing a more transverse outline and ¯attened median

region.

Family Meekellidae Stehli, 1954

Subfamily Meekellinae Stehli, 1954

Genus Orthothetina Schellwien, 1900

Orthothetina cf. iljinae Sokol'skaya, 1965

Fig. 5; 5±6

cf. Meekella cf. baschkirica Tschernyschew; Mansuy,

1914, p. 22, pl. 2, ®g. 15.

cf. Orthothetina iljinae Sokol'skaya, 1965, p. 204, pl. 30,

®gs. 1a and b, 2.

cf. Orthothetina iljinae Sokol'skaya; Tong, 1978, p. 214,

pl. 77, ®gs. 9a±c.

Description. Shell medium sized, approximately 46 mm

wide and 47 mm long, with greatest width at mid-valve;

outline sub-pyramidal; pro®le gently convex; ¯anks

smoothly rounded; sulcus and plications absent; umbonal

region ¯attened, not recurved. Ornament of radial capillae,

moderately strong, even in width, numbering 18±20 per 10

mm at anterior margin. Concentric crenulation present near

the frontal margin, 4 mm in maximum width Ð growth

lamella otherwise not de®ned on surface. Ventral interior

with pair of dental plates, high, arising directly from valve

¯oor, extending at least one-third of valve-length, gently

divergent internally, slightly converging apically. Dorsal

characters unknown.

Remarks. The non-plicate ventral valve with strong,

parallel dental plates clearly indicates Orthothetina. The

present form is characterised by a sub-pyramidal outline

and low in¯ation. It closely resembles O. iljinae

Sokol'skaya (1965, p. 204, pl. 30, ®gs. 1a and b, 2) from

the Gnishik and Khachik Formations of the Trans-Caucasus

by lacking growth lines but instead having a de®ned

concentric elevation anteriorly. O. iljinae, as illustrated by

Tong (1978, p. 214, pl. 77, ®gs. 9a±c) from the Limestone

Member of the upper Chihsia Formation, Guizhou, South

China, has somewhat more ¯attened shells than the Trans-

Caucasian form.

O. phetchabunensis Yanagida (1964, p. 14, pl. 2, ®gs. 5

and 6) from the upper Middle Permian of central Thailand is

similar in outline and size, but its costellation is slightly

stronger than the Bera form. The Thai species displays

concentric rugae over the ventral surface, a feature lacking

in the present form. A Cambodian shell illustrated by

Mansuy (1914, p. 22, pl. 2, ®g. 15) as Meekella cf. basch-

kirica Tschernyschew from Phnom Takream comes close to

the Bera shell in having a sub-pyramidal outline and lacking

concentric lines. The specimen is, however, much smaller

than the Bera form, so precise identi®cation is dif®cult. It


Fig. 5. 1±2, Orthotetes aff. picta Fang, internal and external moulds of ventral shell showing a trace of spondylium with dental ridges, £ 1.5, UKM-F258,

Assemblage B. 3±4, Derbyia sp., rubber cast of ventral exterior, ventral internal mould, £ 2.0, UKM-F259, Assemblage A. 5±6, Orthothetina cf. iljinae

Sokol'skaya, rubber cast of ventral exterior, ventral internal mould showing a pair of dental plates, £ 1.5, UKM-F260, Assemblage C. 7±14, 16±18,

Urushtenoidea chaoi (Ching); 7±11, rubber cast of dorsal interior, dorsal external mould Ð ventral, left lateral views, and anterior view showing mould

of short trail with re¯ected dorsal hollow spines, £ 2.0, enlarged anterior view showing delicate concentric laminae, £ 4.5, UKM-F261, Assemblage A. 12±

14, broken ventral internal mould Ð ventral, anterior and lateral views, £ 2.0, UKM-F262, Assemblage A. 16±18, internal mould of ventral trail Ð ventral

view showing trace of ªmarginal fenceº and ªtransverse ridgeº, lateral views (right and left), £ 1.5, UKM-F263, Assemblage C. 15, Urushtenoidea sp., dorsal

external mould, £ 2.0, UKM-F264, Assemblage A. 19, Strophalosiina sp., rubber cast of dorsal exterior with stereozone, £ 1.5, UKM-F265, Assemblage B.

shows no plication, hence is more likely Orthothetina rather

than Meekella.

Order Productida Sarytcheva and Sokol'skaya, 1959

Suborder Strophalosiidina Schuchert, 1913

Superfamily Aulostegoidea Muir-Wood and Cooper,

1960

Family Aulostegidae Muir-Wood and Cooper, 1960

Subfamily Chonosteginae Muir-Wood and Cooper, 1960

Genus Urushtenoidea Jing (Jin) and Hu, 1978

Discussion. Urushtenoidea has a complicated spine arrange-

ment on the anterior margin of both valves. The genus has

considerable similarity to Urushtenia Likharev. The former,

however, differs from the latter most obviously in having

scaly concentric lamellae (composed of spinous structure)

on the ventral trail. The ventral geniculation is much stron-

ger in Urushtenoidea; its angle often exceeds 608. Urushte-

noidea has a ¯attened dorsal disc with a poorly de®ned trail,

whereas Urushtenia has a geniculate dorsal valve with a

moderately long trail. Species of Urushtenoidea commonly

show ®ner costae than Urushtenia.

Occurrence. Urushtenoidea has hitherto been recorded

only from the eastern Tethys of South China, Cambodia,

Laos and Japan. This report con®rms the presence of the

genus in Peninsular Malaysia. Urushtenoidea is limited to

the Xiangboan to Kuhfengian Stages (Roadian to Wordian)

of South China, Members A±C of the Sisophon Limestone,

the Parafusulina-Neoschwagerina horizons of Laos, and the

Lower Kanokuran Series of Japan, all of which are corre-

lated with the Guadalupian Series.

Urushtenoidea chaoi (Ching (Jin), 1963),

Fig. 5; 7±14, 16±18

Urushtenia chaoi Ching, 1963, p. 15, pl. 1, ®gs., 1±4, 9±

12, 16; pl. 2, ®gs. 7±8, 11±17.

cf. Urushtenia chenanensis (Chan); Tong, 1978, p. 218,

pl. 78. ®gs. 16a±c.

Urushtenia chaoi Ching; Tong, 1978, p. 218, pl. 78, ®g.

18a±d.

Urushtenoidea chaoi (Ching); Jing (Jin) and Hu, 1978, p.

116, pl. 2, ®g. 10.

Urushtenoidea chaoi (Ching); Nakamura, 1979b, p. 230,

pl. 2, ®g. 4.

Urushtenoidea chaoi (Ching); Hu, 1983, pl. 3, ®gs. 6a±c.

Urushtenia chaoi Ching; Zhang et al., 1983, p. 266, pl.

93, ®gs. 12a±e.

Urushtenoidea chaoi (Ching); Zeng, 1987, pl. 1, ®gs. 1±

5, 8±14, 16±19, 26±27, 29±30; pl. 2, ®gs. 4, 7, 9.

Description. Shell average- to large-sized for genus, 19.5±

23 mm wide and 16±18 mm long, with maximum width at

mid-length; outline cylindrical; ears not differentiated;

median fold not well de®ned. Ventral valve moderately

geniculate; trail long; umbo unknown. Dorsal valve almost

¯at on disc, trail about 2 mm. Radial costae well marked on

trail but poor on disc, numbering about 40; terminations of

dorsal costae developing into hollow spines anteriorly along

edge of re¯exed marginal ledge; spines incurved dorsally at

approximately 35±408; concentric wrinkles poorly de®ned

on disc; concentric laminar delicate over dorsal trail, dense

and regular (Fig. 5; 9±11). Dorsal interior with robust bi®d,

sessile cardinal process; lobes sub-ovate, large; alveolus

absent; posterior platform weakly in¯ated; buttress plates

absent; muscle scars large, palmate, hanging anteriorwards;

breviseptum narrow and long; brachial ridges weak and

small; ear baf¯es strong; marginal rim moderately elevated.

In specimen UKM-F263 (Fig. 5; 16±18), moulds of

dorsal marginal endospines (`marginal fence spines' of

Zeng, 1987, p. 502) individually appear as cylindrical

hollows lying on the interior of costal crests. A row of

endospines collectively forming a high `marginal fence'

(term from Jing (Jin) and Hu, 1978, p. 258) behind the

ventral trail. The spines are laterally supported by the

band of the `marginal fence transverse ridge' (term from

Zeng, 1987, p. 502) at mid-length of spines Ð appearing

as a fracture in this specimen.

Remarks. Three shells (one dorsal and two ventral) were

available. The present form is referred to Urushtenoidea

mainly because of its short dorsal trail and spinous nature

of the ventral trail. It is broadly similar to the type species,

U. chaoi (Ching). There are minor differences from the

Chinese form. The Bera form has slightly ®ner costae and

a less strongly elevated posterior platform. The present form

is very much like the Cambodian form, U. chaoi of Zeng

(1987), in having comparable hanging palmate-shaped

adductor scars. Zeng's materials are from the Neoschwager-

ina margaritae bed of the Sisophon region. Jing (Jin) and

Hu (1978), in de®ning the genus, referred to it having a large

alveolus. This feature is not developed on the present mate-

rial, and is not always apparent in other congeneric Chinese

or Cambodian species.

Occurrence. U. chaoi has been reported from Member C

of the Sisophon Limestone, Cambodia, and the Maokou

Formation, Kuhfeng Formation, and Hsiaochiang Lime-

stone of South China, all horizons correlative with the

Neoschwagerina-Yabeina Zones.

Urushtenoidea sp.

Fig. 5; 15

Urushtenoidea maceus (Ching); Nakamura, 1979b, p.

227, (not pl. 1, ®gs. 1±4), pl. 2, ®gs. 1±3.

Remarks. One dorsal valve referred to Urushtenoidea has a

short dorsal trail (1.5 mm). It is medium-sized for the genus

(21 mm wide and 12 mm long). It clearly differs from U.

chaoi in possessing a more transverse outline and slightly

coarser costae. Its cardinal process lobes are globular in

shape, unlike the semi-oval shape of those in U. chaoi,


suggesting that this is a second species of Urushtenoidea in

Assemblage A. Its raised ventral ears are well impressed on

the mould.

Three specimens illustrated by Nakamura (1979b, p. 227,

pl. 2, ®gs. 1±3) as U. maceus (Ching) each from Members

A, B and C of the Sisophon Limestone are close to the

present form in outline and costae, and are probably conspe-

ci®c. In addition, Jing (Jin) and Hu (1978) placed U. maceus

in Uncisteges, but Nakamura (1979b) has not accepted this.

Urushtenia khmeriana Termier and Termier (1970b, p. 446,

text-®gs. 2A±D, pl. 29, ®gs. 1±7) from Phnom Te Bon,

Sisophon, has a ¯attened dorsal shell and scaly laminae on

its ventral trail. It is thus doubtlessly a species of Urushte-

noidea. It has costae and raised ventral ears similar to those

of the present form, but no dorsal view is available.

Genus Strophalosiina Likharev, 1935

Strophalosiina sp.

Fig. 5; 19

Remarks. A single mould of a dorsal exterior was available.

The shell is large for the genus (26 mm wide and 18.5 mm

long), transverse, with a semi-circular anterior margin,

moderate geniculation. It has a well-de®ned dorsal trail

(3.5 mm long) with no concentric wrinkles, unlike that of

Urushtenoidea. The stereozone is smooth, seemingly very

thick, wide and reaching the hinge; it is widest (2 mm) in

front. Ornament of ®ne and sharp costae with weak and

irregular rugae are present on the disc but are absent from

the trail. No dorsal external spines are discernible.

The large transverse shell with the wide and thick stereo-

zone suggests Strophalosiina. The stereozone does not seem

to have any re¯ected hollow spines or funnels, unlike those

of other genera of Chonosteginae such as Urushtenia and

Chonosteges. The present form shares several similarities

with the type species, S. tibetica (Diener) from Tibet,

Cambodia, Timor, Iran and the Trans-Caucasus. The Malay-

sian species, however, differs most obviously in being larger

and having ®ner costae Ð at least 45 in number, whereas

typical S. tibetica has about 30. S. zhejiangensis Liang

(1990, p. 154, pl. 18, ®gs. 9±20) from the lower Lengwu

Formation, China, is less transverse than the Bera form, and

has many large nodules on the dorsal disc.

Superfamily Lyttonioidea Waagen, 1883

Family Lyttoniidae Waagen, 1883

Subfamily Lyttoniinae Waagen, 1883

Genus Gubleria Termier and Termier, 1960

Comments. Gubleria was originally proposed by Termier

and Termier (1960) with the type species, G. disjuncta

from the Sisophon Limestone. The genus is characterised

most importantly by the discontinuous nature of its ventral

median septum and corresponding dorsal median holes.

Cooper and Grant (1974, p. 411) and Grant (1976, p.

162), however, asserted that Gubleria is invalid, being

immature, or a variation of Leptodus Kayser.

In this study, on the contrary, numerous specimens of

large lyttoniids, thought to belong to two species, were

collected. Almost all specimens possess more or less devel-

oped median incisions or segments, indicative of Gubleria,

whereas no form clearly assignable to Leptodus was found.

This seems to contradict to the view of Cooper and Grant

(1974) that Gubleria has individual rather than a generic

characters.

Gubleria aff. ninglangensis Fang and Jiang, 1992

Fig. 6; 1±8

cf. Gubleria ninglangensis Fang and Jiang, 1992, p. 327,

pl. 1, ®gs. 3±5.

Description. Shell size varied, average width 19 mm and

length more than 57 mm; conical, spatulate, weakly twisted;

outline long and narrow, umbo strongly obtuse, gradually

expanding and ¯attening towards mid-length, then twisted

anteriorly; lateral pro®le ¯at to slightly convex. Ventral

valve interior with obsolete muscle ®eld; median septum

highest medianly, faintly continuous, rough incisions

de®ned anteriorly; lateral septa symmetrical, short and

thin, numerous Ð 4 to 5 in 10 mm, at least 18 for longest

but still incomplete specimen UKM-F287 (Figs. 6; 7); inter-

septal spaces much wider than septal ridges, deeply grooved

along septal margins. Dorsal internal plates ¯at and long, later-

ally grooved along margins; median trough segmented irregu-

larly. No trace of papilla observed. Other details unknown.

Remarks. Seven specimens (®ve ventral and two dorsal

internal moulds) are identi®ed as a species of Gubleria by

possessing a discontinuous or segmented median line on

both ventral and dorsal valves. This species is characterised

by its elongate shape. G. ninglangensis Fang and Jiang

(1992, p. 327, pl. 1, ®gs. 3±5) from the lower Emishan

Basalt Formation (lower Capitanian), west Yunnan, is

very close in outline and septal apparatus; it is believed to

be closely allied to the present form. Collemataria tilita

Liang (1990, p. 229, pl. 40, ®g. 2; pl. 41, ®g. 11) from the

lower Lengwu Formation also has a similar outline with an

anteriorly segmented median septum, and is possibly allied

as well. The type species, G. disjuncta Termier and Termier

(1960, p. 241, text-pl. 3, ®gs. 11±14; ®g. 1), is represented

by a couple of fragmentary internal plates. They are ®nely

papillose and show median holes that are irregular in size

and shape. Other details are unknown, preventing useful

comparison.

Gubleria sp.

Fig. 6; 9±12

Remarks. Three ventral shells are assigned to Gubleria. The

shell is medium sized (about 36 mm wide and more than 47

mm long). The outline is ovate, with an umbonal angle



Fig. 6. 1±8, Gubleria aff. ninglangensis Fang and Jiang; 1±2, ventral internal mould of umbonal coneÐventral and lateral views, £ 1.5, UKM-F282, 3,

anterior part of ventral internal mould, £ 1.5, UKM-F283, 4, ventral internal mould, £ 1.5, UKM-F284, 5, internal mould of dorsal internal plate showing

segmentation of median slit, £ 1.5, UKM-F285, 6, internal mould of dorsal internal plate, £ 1.5, UKM-F286, 7, ventral internal mould, £ 1.5, UKM-F287, 8,

ventral internal mould of two imbricating shells, £ 1.5, UKM-F288. 9±12, Gubleria sp.; 9, ventral internal mould displaying anteriorly segmented median

septum, £ 1.5, UKM-F289, 10, ventral internal mould, £ 1.5, UKM-F290, 11±12, ventral internal mould, rubber cast of ventral exterior, £ 1.5, UKM-F291.

13±14, Eolyttonia? sp., fragment of ventral internal mould, £ 1.0, £ 3.0, UKM-F292, (All specimens are from Assemblage C).

ranging 80±1058, and the pro®le is fairly ¯at. They may

belong to Gubleria aff. ninglangensis because of their simi-

lar septal apparatus. The present material, however, is much

wider and has greater umbonal angles. Specimen UKM-

F291 is problematic in that it seems to have an incurved

beak (?) pointing dorsally (Fig. 6; 11).

The present form super®cially recalls Leptodus nobilis

(Waagen). The chief differences, however, concern the

median septum. The Bera form displays a weakly raised

median septum with segmentation. Mature individuals of

L. nobilis typically have a high, thick, continuous, blade-

like median septum. This feature is well illustrated by

Noetling (1905, pl. 17, ®gs. 1 and 2; pl. 18, ®gs. 1 and 2)

for specimens of L. nobilis from the Wargal and Chhidru

Formations of the Salt Range, Pakistan.

Genus Eolyttonia Fredericks, 1924

Eolyttonia? sp.

Fig. 6; 13±14

Remarks. A fragment of a ventral internal mould exhibits

the distinctive ¯uting of its angustilobate septal apparatus,

suggesting possible location in Eolyttonia. It is small (9 mm

in width) for the genus, but its true length is uncertain. It is

longitudinally conical in shape, narrow and slightly wider

anteriorly. The ventral interior has well-formed septa with

crests broadly grooved and low. The lateral septa are trans-

verse, and the interseptal troughs are rounded. It may be a

juvenile, although the internal structure is relatively well

expressed.

Suborder Productidina Waagen, 1883

Superfamily Productoidea Gray, 1840

Family Productellidae Schuchert, 1929

Subfamily Marginiferinae Stehli, 1954

Tribe Paucispiniferini Muir-Wood and Cooper, 1960

Genus Transennatia Waterhouse, 1975

Transennatia termierorum sp. nov.

Fig. 7; 1±10, 13±15

Spyridiophora ? timorensis Termier and Termier, 1970a,

p. 58, pl. 5, ®g. 1, text-®gs. 1±4.

Etymology. After Professors Henri Termier and GenevieÁve

Termier, who greatly contributed to knowledge on the

South-East Asian Permian brachiopods, and ®rst described

this species.

Type specimen. The specimen illustrated as Spyridio-

phora ? timorensis Termier and Termier (1970a, pl. 5, ®g.

1) (not Hamlet) is designated as the holotype.

Diagnosis. Shell medium-sized, geniculation abrupt,

sulcus and fold strong anteriorly, ears proportionally large

and projecting, costae notably converging, branching, and

sturdy on the trail, coarse reticulation present on the disc.

Description. Shell size ranging from 13±23.5 mm wide,

9±14.5 mm long, more than 8.5 mm high, commonly small;

outline sub-quadrate, widest part at hinge; ears large,

convex and costate; ventral umbo pointed; geniculation

abrupt with angle about 508; dorsal disc fairly ¯attened,

semi-circular in shape, almost without a fold, notably reti-

culate; trail long. Surface coarsely ornamented by sturdy

costae, branching on the ¯anks, converging in the sulcus

and on the fold, with crests rounded, and similarly rounded,

wide and shallow interspaces; reticulation con®ned to disc,

concentric rugae faint on posterior part of the trail. Ventral

interior with wide and ¯at platform, well elevated, extend-

ing from apex; muscle scar obsolete or obscure.

Remarks. This form is characterised by its abrupt and

strong geniculation and extended ears. It is most like Spyr-

idiophora ? timorensis Termier and Termier (1970a, pl. 5,

®g. 1, text-®gs. 1±4) from Sisophon, Cambodia, but lacks

the row of spine bases along the ears seen in the Cambodian

material; this could be an artifact of preservation. Termier

and Termier (1970a) originally de®ned their timorensis as a

synonym of Productus gratiosus Waagen, as illustrated by

Broili (1916, pl. 2, ®gs. 4±5, 7±13), from the Basleo, Koeka

bei Koeafeoe and Bitauni areas of Timor. However, Grant

(1976, p. 134) asserted that only the Basleo specimens

belong to Transennatia, the others being Retimarginifera.

This view was followed by Archbold (1984, p. 115) and

Archbold and Bird (1989, p. 108). Waterhouse (1978, p.

119) considered Termiers' timorensis to be a junior syno-

nym and homonym of Productus timorensis Hamlet.

T. timorensis is close to T. termierorum in size and orna-

ment. The former can, however, be distinguished from the

latter by having much smaller ears and a strongly folded

dorsal disc. There is admittedly considerable variation

among the materials of T. timorensis as understood by Arch-

bold and Bird (1989, p. 108). For instance, the Kasliu form

of Archbold and Bird (1989, p. 107, ®gs. 3E±I) is excep-

tionally large and has ®ne ornament, whereas the Basleo-

Wesleo forms of Hamlet (1928, pl. 2, ®gs. 2±4) have strong

ornament and a deep fold. The small specimen in ®g. 7 of

Diener (1897, pl. 3) identi®ed as P. gratiosus from the Chit-

ichun Limestone, Tibet, has a deep sinus and sturdy costae;

it strongly resembles T. termierorum. This Tibetan material

is possibly conspeci®c, although it is slightly wider in

outline. T. insculpta Grant (1976, p.135, pl. 32, ®gs. 1±

37; pl. 33, ®gs. 1±16) from the Rat Buri Limestone of south-

ern Thailand is similar. Its geniculation is, however, not as

abrupt as that of the new species. T. termierorum differs

from the type species, T. gratiosa (Waagen, 1884, pl. 72,

®gs. 3±7) from the Salt Range, Pakistan, most obviously in

having larger ears. The geniculation of T. gratiosa varies

from shell to shell; smaller ones tend to be much less geni-

culate. This trend is probably common in the genus; it

occurs also in T. insculpta Grant. The small but highly

geniculate Bera shells may, therefore, be mature forms.

Occurrence. This form was originally recorded from the

Yabeina bed of a limestone hill, Phnom Takream, in the

Battambang region (Termier and Termier, 1970a, p. 58),

suggesting that its occurrence is con®ned to an upper part


of Member B to Member D, most likely Member C of the

Sisophon Limestone.

Transennatia sp. indet.

Fig. 7; 11±12

Remarks. The small conjoined shells (9 mm wide and 7.5

mm long) with no geniculation is probably a juvenile. It,

nevertheless, displays features diagnostic of the genus, such

as strong reticulation, tiny ears and branching costae. It

occurs in Assemblage C.

Tribe Marginiferini Muir-Wood and Cooper, 1960

Genus Spinomarginifera Huang, 1932

Spinomarginifera sp.

Fig. 7; 16±17

Remarks. One squashed mould of a dorsal interior was

available. The distinctive appearance of its muscle ®eld

suggests alliance with Spinomarginifera. The shell is

small for the genus; its dorsal disc is about 13 mm wide

and 13.5 mm long. It seems to have a row of endospines

around the anterior half of the disc (characteristic of the

genus) but this is poorly preserved. There is too little infor-

mation for speci®c assignment.

Subfamily Overtoniinae Muir-Wood and Cooper, 1960

Tribe Costispiniferini Muir-Wood and Cooper, 1960

Genus Echinauris Muir-Wood and Cooper, 1960

Echinauris sp.

Fig. 7; 18

Remarks. One internal mould of a dorsal disc was available.

It is medium-sized (18 mm wide and 15.5 mm long). Its pan-

like appearance with a low and short median septum indi-

cates Echinauris. As common in the genus, brachial ridges

are not de®ned. The marginal ridge is moderate. The exter-

nal features are not unknown, but rugose radiations occur

internally, probably corresponding to the external spines.

Considering the poorly developed internal structure, it

may be an immature shell, although it is not small in size.

A Cambodian shell ®gured by Termier and Termier

(1970b, p. 450, ®g. 6) as E. opuntia (Waagen) has a longer

median septum than the Bera form. Productus khmerianus

Mansuy (1914, p. 17, pl. 6, ®gs. 3a±d) from Phnom Roang,

Sisophon, possibly belongs to Echinauris, as suggested by

Waterhouse and Piyasin (1970, p. 130) and Grant (1976, p.

123). Prior to them, Ishii et al. (1969, p. 48) reported the

same species as Echinauris khmerianus from Member D of

the Sisophon Limestone. Its interior is not known, hindering

comparison with the Bera material, although the size is

close.

Family Productidae Gray, 1840

Subfamily Productinae Gray, 1840

Tribe Spyridiophorini Muir-Wood and Cooper, 1960

Genus Spyridiophora Cooper and Stehli, 1955

Spyridiophora gubleri Termier and Termier, 1970a

Fig. 7; 19

Productus gratiosus Waagen; Mansuy, 1913, p. 115, pl.

13, ®g. 1.

Productus gratiosus Waagen; Chi-Thuan, 1961 p. 276, pl.

5, ®gs. 8 and 9.

Spyridiophora gubleri Termier and Termier, 1970a, p. 60,

pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G±I.

Lectotype. A holotype was not selected by Termier and

Termier (1970a). Their specimen of conjoined valves (pl.

5, ®gs. 4±6) is here selected as the lectotype for this species.

Description. About medium size for genus, 15.5 mm wide

and about 9 mm long, widest at hinge and of similar width at

mid-length; outline sub-rectangular with ears well ¯attened

and ornamented; anterior margin broadly emarginate;

pro®le gently concave; trail short; geniculation weak,

restricted to median region between ¯anks; median fold

notably shallow, poorly delimited on visceral disc, very

broad on anterior trail. The umbonal region is missing, so

true length and height are uncertain. Ornament distinctively

reticulate; radial costae strong, coarse, rounded, of even

width, converging slightly in the sulcus, numbering about

16 at mid-length, increasing occasionally by bifurcation;

intertroughs of similar width, rounded; concentric rugae

strong, but ®ner than costae; external spines not de®ned.

Remarks. The present dorsal specimen is damaged but is

suf®ciently well preserved to retain the main characteristics

of Spyridiophora gubleri. The Bera form is almost identical

to all the Cambodian shells ®gured by Mansuy (1913, pl. 13,

®g. 1), Chi-Thuan (1961, pl. 5, ®gs. 8 and 9) and Termier

and Termier (1970a, pl. 5, ®gs. 4±6, text-®gs. 8±10, 12G±

I). In addition, Colani (1919, p. 10, pl. 1, ®g. 2) re-illustrated

the Mansuy's (1913) Spyridiophora specimen named

Productus gratiosus for comparison with her Vietnamese

gratiosus; the latter is probably a species of Transennatia.

Spyridiophora is known to bear the distinctive elevation


Fig. 7. 1±10, 13±15, Transennatia termierorum sp. nov.; 1±4, ventral internal mould Ð ventral, lateral (right), anterior, and lateral (left) views, £ 3.0, UKM-

F266, 5±8, dorsal external mould with accompanying natural cast of shell remaining anteriorly Ð posterior, ventral, anterior, and lateral views, £ 3.0, UKM-

F267, 9±10, dorsal external mould Ð posterior and anterior views, £ 3.0, UKM-F268, 13±15, dorsal external mould Ð ventral, anterior, and posterior views,

the last showing a hole corresponding to the ventral beak, £ 2.0, UKM-F269, Assemblage A. 11±12, Transennatia sp. indet., dorsal external mould, dorsal

internal mould resting on ventral external mould, £ 3.0, UKM-F270, Assemblage C. 16±17, Spinomarginifera sp., dorsal internal mould and rubber cast,

£ 3.0, UKM-F271, Assemblage A. 18, Echinauris sp., rubber cast displaying dorsal disc interior, £ 2.0, UKM-F272, Assemblage A. 19, Spyridiophora

gubleri Termier and Termier, incomplete dorsal external mould, £ 3.0, UKM-F273, Assemblage A. 20±23, Spiriferinid family indet., incomplete mould of a

ventral interior Ð ventral, lateral, posterior and dorsal views, the last showing interarea, £ 3.5, UKM-F274, Assemblage C.


Fig. 8. 1±11, Kozlowskia sp.; 1±5, ventral internal mould of semi-mature shell (Arrows indicate spine bases.) Ð ventral, posterior, lateral (right and left) and

anterior views, £ 2.5, UKM-F275, 6±11, internal mould of conjoined shells Ð ventral, posterior, lateral (right) and anterior views of ventral interior, and

dorsal view and rubber cast of dorsal interior, £ 2.0, UKM-F276, Assemblage A. 12±14, Linoproductus sp.; 12, ventral internal mould, £ 1.5, UKM-F277,

13, ventral internal mould, £ 1.5, UKM-F278, 14, ventral internal mould, £ 1.5, UKM-F279, Assemblage B. 15, Rhynchonellid family indet, incomplete

ventral internal mould, £ 2.0, UKM-F280, Assemblage B. 16±17, Phricodothyris sp., rubber cast of ventral exterior, £ 2.0, enlarged section showing

microornament, £ 6.0, UKM-F281, Assemblage A.

of the brachial adductor scars, called a `spyridium' (term

proposed by Cooper and Stehli, 1955, p. 472). It is well

developed in the type species, S. distincta Cooper and Stehli

from West Texas. By comparison, the spyridium of S.

gubleri, as illustrated by Termier and Termier (1970a, pl.

5, ®g. 6), is less salient. Grant (1976, p. 136) regarded the

Chi-Thuan's (1961) materials as Transennatia instead of

Spyridiophora, but made no mention of the Termier and

Termier (1970a) designation. Waterhouse (1978, p. 119)

compared the Mansuy's (1913) material to his Nepalese

Transennatia sp., but does not seem to have accepted the

Termiers' work. However, considering the fact that the

Termiers' material clearly displays the typical spyridium,

the present form is placed in Spyridiophora without

hesitation.

Productus craticulatus Reed (1931, p. 11, pl. 1, ®gs., 1

and 2) from the Neoschwagerina craticulifera bed of the

Bamian Limestone, Afghanistan, obviously belongs to

Spyridiophora, judged from the description and illustra-

tions. It appears to be close to S. gubleri, especially as

regards its ornament. The South-East Asian forms are

more transverse than the Afghan form, but this may be no

more than within the same species. Because of its distinctive

reticulation and transverse outline, Productus margaritatus

Mansuy (1913, p. 28, pl. 2, ®g. 6) from Kham-keut, Laos, is

possibly a species of Spyridiophora. The same species was

illustrated by Huang (1932, p. 30, pl. 1, ®gs. 22±24) from

the Late Permian of Guizhou, South China. This species has

®ner reticulation than S. gubleri. Grant (1976, p. 136) noted

that this species is unlike Transennatia. Liang (1990)

proposed two species of Spyridiophora, namely S.

huadongensis (p. 159, pl. 24, ®gs. 1±21) and S. plegma

(p. 161, pl. 24, ®gs. 22 and 23), both from the Lengwu

Formation. They both differ from S. gubleri in having

much ®ner ornament.

Occurrence. S. gubleri has previously been found only in

the Sisophon Limestone at Phnom Tup in the Sisophon

region (Mansuy, 1913; Chi-Thuan, 1961), and at Phnoms

Anseh, Totung and Takream in the Battambang region

(Termier and Termier, 1970a). Nakamura (1979a) also

recorded this species in Member C of the same formation.

Species of Spyridiophora are extremely rare in the Tethyan

Province. They have been recognised only in the Early to

Middle Permian of South China, Indochina, Afghanistan

and the Carnic Alps.

Tribe Kozlowskiini Brunton et al., 1995

Genus Kozlowskia Fredericks, 1933

Kozlowskia sp.

Fig. 8; 1±11

Remarks. This species is represented by two specimens of

mature and semi-mature shells. The shells are slightly large

for the genus, 20 mm wide, 15 mm long and the disc is 8 mm

high for the mature specimen UKM-F276. The body cavity

is of moderate size. The cardinal process is sessile, and the

breviseptum is short. The brachial ridges are located very

distally. Two small dorsal endospines are discerned.

Generic assignment to Kozlowskia is con®rmed by the

distinctive pattern of the ventral strut spines. Three large

stout spines are seen on the ventral valve of the semi-mature

specimen UKM-F275 Ð one on each lateral slope (Fig. 8;

2±4) and one on the anterior of the disc (Fig. 8; 5). This is

further supported by the appearance of the dorsal interior,

speci®cally the strong ear baf̄ es, large pair of adductors,

high breviseptum and dorsal marginal ridge forming a wide

platform. The indistinct or absent sulcus and very weak

ventral ornament also characterise the genus.

Two species of Kozlowskia, K. cornuta Grant (1976, p.

118, pl. 26, ®gs. 24±29; pl. 27, ®gs. 35±38) and K. opipara

Grant (1976, p. 121, pl. 28, ®gs. 1±27), were found in the

Rat Buri Limestone of southern Thailand. Their dorsal inter-

iors differ from the Bera form. Grant (1976, p. 118) noted

that species of Kozlowskia were restricted to the Pennsylva-

nian and Early Permian (Wolfcampian); his two Thai

species were the only Asian occurrences known at the

time of his study. Since then, only two additional Asian

occurrences have been reported, both from Early Permian

horizons in Xinjiang, northwestern China (Wang and Yang,

1993; Wang, 1995), both being reports of the type species,

K. capacii d'Orbigny.

Waterhouse (1981, p. 13) indicated that Marginifera

sensu Diener (1897, pl. 4, ®gs. 11±13; pl. 5, ®gs. 1 and 2)

from the Chitichun Limestone (Kalabaghian) of Tibet may

be Kozlowskia. However, as con®rmed by Waterhouse

(written communication, 24 July 1999), this was in error

because he previously discussed the Diener's specimens in

terms of Marginifera, as a species separate from the type

species, M. typica (see Waterhouse and Piyasin, 1970, p.

127; Waterhouse, 1978, p. 30). Marginifera and Kozlowskia

often resemble each other considerably, as pointed out by

Cooper and Grant (1975, p. 969). The two genera, however,

can be differentiated by the very different spine patterns on

the ventral valve. They are now classi®ed into the different

families by Brunton et al. (1995).

Superfamily Linoproductoidea Stehli, 1954

Family Linoproductidae Stehli, 1954

Subfamily Linoproductinae Stehli, 1954

Genus Linoproductus Chao, 1927

Linoproductus sp.

Fig. 8; 12±14

Remarks. Three impressions of the ventral interior display

characteristic costellae of Linoproductus. The shells are

medium to large for the genus, ranging 29±40 mm wide

and 30±40 mm long. They are semi-ovate to semi-elongate

in outline, and lack a median fold. Costellae are irregular

and uneven, numbering 7±8 in 5 mm medianly. Rugae are

present only on the lateral trail as broad wrinkles. Parallel

transverse lines intersect costellae in the largest specimen

UKM-F278 (Fig. 8; 13).


Class Rhynchonellata Williams et al., 1996

Order Rhynchonellida KuÈhn, 1949

Rhynchonellid family indet.

Fig. 8; 15

Remarks. A poorly preserved impression of a ventral inter-

ior is the only rhynchonellid in the Bera fauna. It is small

(approximately 20 mm wide and 16 mm long), and has

simple, weak costae Ð four in the sulcus and ®ve on each

¯ank.

Order Spiriferida Waagen, 1884

Suborder Delthyridina Ivanova, 1972

Superfamily Reticularioidea Waagen, 1884

Family Elythidae Fredericks, 1924

Subfamily Phricodothyridinae Caster, 1939

Genus Phricodothyris George, 1932

Phricodothyris sp.

Fig. 8; 16±17; 9; 1

Remarks. One incomplete ventral specimen was available.

The shell is sub-orbicular, medium-sized (30 mm wide and

about 22 mm long), with double-barrelled (biramous) spine-

bases, suggesting location in Phricodothyris. It is broadly

convex without sulcation, with its umbo moderately high

and obtuse at about 908. The spine arrangement of the

Bera shell is similar to that of P. pyriformis Pavlova

(1969, p. 93, pl. 8, ®gs. 1±3) (Fig. 9; 2), but differs slightly

in that the Bera species has a third row of spine bases (Fig. 9;

1) and slightly coarser concentric laminae. P. rotiformis

Liang (1990, p. 290, pl. 63, ®gs. 17±23) from the lower

Lengwu Formation, South China, is also close; it seems to

have a similar spine arrangement of three rows (Fig. 9; 3).

Only a mould of the micro-ornament was illustrated for the

Chinese species; the exact shape of its spines is, therefore,

uncertain.

Order Spiriferinida Ivanova, 1972

Suborder Spiriferinidina Ivanova, 1972

Spiriferinid family indet.

Fig. 7; 20±23

Remarks. A single mould of a ventral interior is the only

spiriferinid in the Bera fauna. It is small, spiriferiform, with

simple, strong plicae indicating Spiriferinidina. The shell is

about 9.5 mm wide, 7.5 mm long and 4.5 mm high in size,

and has three ribs on each ¯ank.

Acknowledgements

M. Sone's research was carried out for his B.Sc. Honours

degree at Deakin University in 1998±99, and was ®nanced

by his parents, Masao and Toshiko Sone. He is indebted to

Prof. N.W. Archbold, Dr. S. Shen and Mr. Z. Chen for their

invaluable discussions. Drs. J.B. Waterhouse, I. Metcalfe

and Prof. J.A. Talent are thanked for critically reading the

manuscript. Dr. Ahmad Jantan and Prof. Ibrahim Komoo are

acknowledged for providing ®nancial supports from their

government IPRA Projects 02±02±02±0004 and 02±02±

02±0015, respectively. This paper was completed while

M. Sone was supported by a fellowship of LESTARI.

References

Archbold, N.W., 1984. Western Australian occurrences of the Permian

brachiopod genus Retimarginifera. Alcheringa 8 (1/2), 113±122.

Archbold, N.W., Bird, P.R., 1989. Permian Brachiopoda from near Kasliu

village, West Timor. Alcheringa 13 (1/2), 103±123.

Broili, F., 1916. Die permischen Brachiopoden von Timor. In: Wanner, J.

(Ed.), PalaÈontologie von Timor 7, Stuttgart, pp. 1±104.

Brunton, C.H.C., Lazarev, S.S., Grant, R.E., 1995. A review and new

classi®cation of the brachiopod order Productida. Palaeontology 38

(4), 915±936.

Brunton, C.H.C., Lazarev, S.S., 1997. Evolution and classi®cation of the

Productellidae (Productida), Upper Paleozoic brachiopods. Journal of

Paleontology 71 (3), 381±394.

Carter, J.L., Johnson, J.G., Gourvennec, R., Hou, Hong-fei, 1994. A revised

classi®cation of the spiriferid brachiopods. Carnegie Museum, Annals

63 (4), 327±374.

Caster, K.E., 1939. A Devonian fauna from Columbia. Bulletin of Amer-

ican Paleontology 24 (83), 1±218, pls. 1±14.

Chao, Y.T., 1927. Productidae of China, Part 1: Producti. Palaeontolgia

Sinica, Series B 5 (2), 1±244, 16 pls.

Chi-Thuan, T.T., 1961. Les brachiopodes permiens du Phnom-Tup (Siso-

phon Ð Cambodge). Annales de la FaculteÂ des Sciences, UniversiteÂ de

Saigon 1961, 267±308, 8 pls. (in French).

Ching, Y., 1963. Urushtenia from the Lower Permian of China. Acta

Palaeontologica Sinica 11 (1), 1±31, 2 pls. (in Chinese with English

abstract).

Colani, M., 1919. Sur quelques fossiles Ouralo-Permiens de Hongay.

Service GeÂologique de l'Indochine, Bulletin 6 (5), 1±27, 2 pls. (in

French).

Cook, R.H., Suntharalingam, T., The geology and mineral resources of

Pahang Tenggara (Sectors ªEº and ªFº). Geological Survey Malaysia

Annual Report 1970, pp. 104±116.

Cooper, G.A., Grant, R.E., 1974. Permian brachiopods of West Texas, II.

Smithsonian Contributions to Paleobiology 15, 233±793, pls. 24±191.

Cooper, G.A., Grant, R.E., 1975. Permian brachiopods of West Texas, III

(Part 1 and 2). Smithsonian Contributions to Paleobiology 19, 794±

1920, pls. 192±502.


Fig. 9. Comparison of microornament among three Phricodothyris species

(all illustrations £ 25), 1, Phricodothyris sp. in this paper, 2, P. pyriformis

Pavlova (1969, text-®g. 58), 3, P. rotiformis Liang (1990, pl. 63, ®g. 23).

Cooper, G.A., Stehli, F.G., 1955. New genera of Permian brachiopods from

West Texas. Journal of Paleontology 29 (3), 469±474, pls. 52±54.

Diener, C., 1897. Himalayan fossils. The Permocarboniferous fauna of

Chitichun No.1. Palaeontologia Indica, Series 15 1 (13), 1±105.

Fang, R., 1973. The Early Permian Brachiopoda from Xiaoxinzhai of

Gengma, Yunnan, and its geological signi®cance. Contribution to the

Geology of the Qinghai-Xizang (Tibet) Plateau 11, 110±119, 7 pls. (in

Chinese with English abstract).

Fang, R., Jiang, N., 1992. Discovery and its geological signi®cance of

Gubleria from Permian Emeishan Basalt Formation in Ninglang

Xipiaoluo of northwestern Yunnan. Yunnan Geology 11 (4), 325±

329, 1 pl. (in Chinese with English abstract).

Fantini Sestini, N., 1965. Bryozoans, brachiopods and molluscs from Ruteh

Limestone (Permian). The geology of the Upper Djadjerud and Lar

Valleys (North Iran) II, Palaeontology. Rivista Italiana di Paleontologia

e Stratigra®a 71 (1), 13±108, pls. 2±8.

Fischer de Waldheim, G., 1829. Quelques fossiles du Gouvernement de

Moscou. SocieÂteÂ ImpeÂriale des Naturalistes de Moscou Bulletin 1 (12),

375±376.

Fontaine, H., Ibrahim, A., Khoo, H.P., Nguyen, D.T., Vachard, D., 1994.

Geological Papers Volume 4. Geological Survey of Malaysia, Ministry

of Primary Industries, pp. 1±175.

Fredericks, G.N., 1924. Ussuriiskii verkhnii paleozoi. I. Brachiopoda

(Upper Paleozoic of the Ussuriland, I: Brachiopoda.). Materialy po

Geologii i Poleznym Iskopaemym Dal'nego Vostoka 28, 1±53, 1 pl.

(in Russian).

Fredericks, G.N., 1933. Paleontologicheskie etiudy, 4: O nekotorykh

verkhne-paleozoiskikh brakhiopodakh Evrazii (palaeontological

notes, 4: on some Upper Paleozoic brachiopods of Eurasia). Materialy

Tsentral'nogo Nauchno-Issledovatel'skogo Geologo-Razvedochnogo

Instituta (TSNIGRI), Paleontolgiia i Stratigra®ia Sbornik 2, 24±33 (in

Russian).

Geological Survey of Malaysia, 1985. Geological Map of Peninsular

Malaysia, 8th ed. (1:500,000). Directorate of Geological Survey Malay-

sia, Kuala Lumpur.

George, T.N., 1932. The British Carboniferous reticulate Spiriferidae.

Geological Society of London Quarterly Journal 88, 516±575, pls.

31±35.

Gobbett, D.J., 1973. Chapter 4, Upper Paleozoic. In: Gobbett, D.J., Hutch-

ison, C.S. (Eds.), Geology of the Malay Peninsula (West Malaysia and

Singapore). Wiley/Interscience, New York, pp. 61±96.

Grant, R.E., 1976. Permian brachiopods from southern Thailand. Journal of

Paleontology (Paleontological Society Memoir 9) 50 (3), 1±269.

Gray, J.E., 1840. Synopsis of the contents of the British Museum, 42nd ed.

London, 308 pp.

Hamlet, B., 1928. Permische Brachiopoden, Lamellibranchiaten und

Gastropoden von Timor. Jaarboek van het Mijnwezen in Neder-

landsch-IndieÈ 56 (2), 1±115, 12 pls.

He, X., Shi, G.R., 1996. The sequence of Permian brachiopod assemblages

in South China. In: Cooper, P., Jin, J. (Eds.), Proceedings of the Third

International Brachiopod Congress, 2±5 September 1995, Subdury,

Ont., Canada, pp. 111±115.

Hu, S., 1983. Brachiopods from the Hsiaochiangpien Limestone, southern

Jiangxi. Acta Palaeontologica Sinica 22 (3), 338±345, pls. 1±3 (in

Chinese with English abstract).

Huang, T.K., 1932. Late Permian Brachiopoda of southwestern China.

Palaeontologia Sinica, Series B 9 (1), 1±113, 9 pls.

Igo, H., 1964. Permian fossils from northern Pahang, Malaya. Geology and

Paleontology of Southeast Asia 1, 191±207, pl. 2.

Igo, H., 1967. Some Permian fusulinids from Pahang, Malaya. Geology and

Palaeontology of Southeast Asia 3, 30±38, pls. 4±9.

Igo, H., Rajah, S.S., Kobayashi, T., 1979. Permian fusulinacean from the

Sungai Sedili area, Johore, Malaysia. Geology and Palaeontology of

Southeast Asia 20, 95±118, pls. 15±26

Ishii, K., 1966. On some fusulinids and other Foraminifera from the

Permian of Pahang, Malaya. Journal of Geosciences, Osaka City

University 9 (4), 131±136, pls. 5±6.

Ishii, K., Kato, M., Nakamura, K., 1969. Permian limestones of West

Cambodia Ð lithofacies and biofacies (contribution to the geology

and palaeontology of Cambodia, Part 3). Palaeontological Society of

Japan, Special Papers 14, pp. 41±55, pls. 9±11.

Ivanova, E.A., 1972. Osnovnyye zakonomernosti evolyutsii spiriferid

(Brachiopoda) [main features of spiriferid evolution (Brachiopoda)].

Paleontologicheskii Zhurnal 1972 (3), 28±42 (in Russian).

Jasin, B., Said, U., Rahman, R.A., 1995. Late Middle Permian radiolarian

from the Jengka area, central Pahang, Malaysia. Journal of Southeast

Asian Earth Sciences 12 (1/2), 79±83.

Jing (Jin). Y., Hu, S.Z., 1978. Brachiopods of the Kuhfeng Formation in

south Anhui and Nanking Hills. Acta Palaeontologica Sinica 17 (2),

101±127, 4 pls. (in Chinese with English abstract).

Jin, Y., Wardlaw, B.R., Glenister, B.F., Kotlyar, G.V., 1997. Permian

chronostratigraphic subdivisions. Episodes 20 (1), 10±15.

Jones, C.R., Gobbett, D.J., Kobayashi, T., 1966. Summary of fossil record

in Malaya and Singapore 1900±1965. Geology and Palaeontology of

Southeast Asia 2, 309±359.

Kobayashi, F., 1997. Middle Permian biogeography based on fusulinacean

faunas. In: Ross, C.A., Ross, J.R.P., Brenckle, P.L. (Eds.), Late Paleo-

zoic Foraminifera; their Biostratigraphy, Evolution and Paleoecology,

and the Mid-Carboniferous Boundary. Cushman Foundation for Fora-

miniferal Research Special Publication vol. 36, pp. 73±76.

KuÈhn, O., 1949. Lehrbuch der PalaÈozoologie. Schweizerbart'sche Verlags-

buchhandlung, Stuttgart, 326 pp.

Leman, M.S., 1993. Upper Permian brachiopods from northwest Pahang,

Malaysia. In: Thanasuthipitak, T. (Ed.), Proceedings of the Interna-

tional Symposium on Biostratigraphy of Mainland Southeast Asia:

Facies & Paleontology. Chang Mai, Thailand, pp. 203±218.

Leman, M.S., 1994. The signi®cance of Upper Permian brachiopods from

Merapoh area, northwest Pahang. Geological Society of Malaysia

Bulletin 35, 113±121.

Leman, M.S., Mohamed, K.R., Sone, M. On the new Permian Bera Forma-

tion from the Bera District, Pahang, Malaysia, Proceedings of the

Annual Geological Conference 2000. 8±9 September 2000, Penang.

Geological Society of Malaysia. In press

Leven, E.Ya.(Ja.), 1992. Problems of Tethyan Permian stratigraphy. Inter-

national Geology Review 34 (10), 976±985.

Leven, E.Ya.(Ja.), 1993. Phylogeny of the sumatrinids and questions on the

zonal subdivision of the Midian Stages of the Permian System. Paleon-

tological Journal 27 (3), 29±35.

Leven, E.Ja., Grant-Mackie, J.A., 1997. Permian fusulinid foraminifera

from Wherowhero Point, Orua bay, Northland, New Zealand. New

Zealand Journal of Geology and Geophysics 40, 473±486.

Liang, W., 1990. Lengwu Formation of Permian and its brachiopod fauna in

Zhejiang Province. People's Republic of China, Ministry of Geology

and Mineral Resources, Geological Memoirs, Series 2. 10. Geological

Publishing House, Beijing, pp. 1±522, 84 pls. (in Chinese with English

summary).

Likharev, B., 1935. Bemerkungen uÈber einige oberpalaÈozoische Brachio-

poden. Zentralblatt fuÈer Mineralogie, Geologie und PaÈlaeontologie,

Abt. B 9, 369±373.

MacDonald, S., 1970. Geology and mineral resources of the Lake Chini Ð

Sungei Bera Ð Sungei Jeram area of south-central Pahang. Geological

Survey of Malaysia, Map Bulletin 1, pp. 1±44.

Mansuy, H., 1912. Mission du Laos 1, GeÂologie des environs de Luang-

Prabang. Service GeÂologique de l'Indochine MeÂmoires 1 (4), 1±32, pls.

1±7 (in French).

Mansuy, H., 1913. Faunes des Calcaires aÁ Productus de l'Indochine,

PremieÁre SeÂrie. Service GeÂologique de l'Indochine MeÂmoires 2 (4),

1±133, 13 pls. (in French).

Mansuy, H., 1914. Faunes des Calcaires aÁ Productus de l'Indochine, Deux-

ieÁme SeÂrie. Service GeÂologique de l'Indochine MeÂmoires 3 (3), 1±59,

pls. 1±7 (in French).

Metcalfe, I., 1996a. Pre-cretaceous evolution of SE Asian terranes. In: Hall,

R., Blundell, D.J. (Eds.), Tectonic Evolution of Southeast Asia, Geolo-

gical Society Special Publication 106, pp. 97±122.


Metcalfe, I., 1996b. Gondwanaland dispersion, Asian accretion and evolution

of Eastern Tethys. Australian Journal of Earth Sciences 43, 605±623.

Metcalfe, I., 1998. Palaeozoic and Mesozoic geological evolution of the SE

Asian region: multidisciplinary constraints and implications for biogeo-

graphy. In: Hall, R., Holloway, J.D. (Eds.), Biogeography and Geolo-

gical Evolution of SE Asia. Backbuys, Leiden, pp. 25±41.

Muir-Wood, H.M., 1948. Malayan Lower Carboniferous Fossils and their

Bearing on the Visean Palaeogeography of Asia. British Museum

(Natural History), London, pp. 1±118.

Muir-Wood, H.M., Cooper, G.A., 1960. Morphology, classi®cation and life

habits of the Productoidea (Brachiopoda). Geological Society of Amer-

ican Memoir 81, 1±447, 135 pls.

Nakamura, K., 1979a. Kenkyu Seika Houkokusho, ªHimalaya Zenentai

oyobi Honpousan Perumuki Wanzokurui no Hikaku Kenkyuº (the

report of research results, ªcomparison between Permian brachiopods

from the Himalaya Frontal Margin and Japanº). Monbu-syo Science

Research Subsidy (General Science Category C) for the Syowa Year 53

(in Japanese), unpublished.

Nakamura, K., 1979b. Additional occurrences of Urushtenoidea (brachio-

pods) from the Permian of Asia. Journal of Faculty of Science,

Hokkaido University, Series 4. Geology and Mineralogy 19 (1/2),

221±233, 3 pls.

Nakamura, K., Shimizu, D., Liao, Z., 1985. Permian palaeobiogeography of

brachiopods based on the faunal provinces. In: Nakazawa, K., Dickins,

J.M. (Eds.), The Tethys. Tokai University Press, Tokyo, pp. 185±198.

Nakazawa, K., 1973. On the Permian fossils from Jengka Pass, Pahang,

Malay Peninsula. Tohoku University, Science Report Series 2 (Geol-

ogy), Special Volume 6 (Hatai Memorial Volume), 277±296, pls. 30±

31

Noetling, F., 1905. Untersuchungen uÈber die Familie Lyttoniidae Waag.

emend. Noetling. Palaeontographica 51, 129±153, pls. 15±18.

OÈ pik, A.A. 1934. UÈ ber Klitamboniten. Acta Commentationes, Series A, 26

(3), 1±239, 48 pls. Tartu University, Dorpat.

Pavlova, E.E., 1969. Razvitie brakhiopod semeistva Reticulariidae (devel-

opment of brachiopods of the family Reticulariidae). Akademia Nauk

SSSR, Palaeontologicheskii Institut, Trudy 120, pp. 1±128 (in Russian).

Reed, F.R.C., 1931. Upper Carboniferous fossils from Afghanistan.

Palaeontologia Indica 19, 1±39, 4 pls.

Rudwick, M.J.S., Cowen, R., 1967. The functional morphology of some

aberrant strophomenide brachiopods from the Permian of Sicily. Soci-

eta paleontologica Italiana, Bollettino 6 (2), 113±176, pls. 32±43.

Sarytcheva, T.G., Sokol'skaya, A.N., 1959. O klassi®katsin lozhnoporis-

tykh brachiopod (on the classi®cation of pseudopunctate brachiopods).

Vol. 125, pp. 181±184. Akademiia Nauk SSSR, Doklady.

Schellwien, E., 1900. BeitraÈge zur Systematik der Strophomeniden des

oberen Palaeozoicum. Neues Jahrbuch fuÈer Mineralogie, Geologie

und PalaÈeontologie 190, 1±15, 1 pl.

Schuchert, C., 1913. Class Brachiopoda. The Lower Devonian deposits of

Maryland. Maryland Geological Survey, Baltimore. 290±449.

Schuchert, C., 1929. Brachiopoda (generum et genotyporum index et

bibliographia). In: Schuchert, C. and LeVene, C.M. (Eds.), Fossilium

Catalogus. I. Animalia. W. Junk, Berlin. 140 pp.

Shen, S., Shi, G.R., 1996. Diversity and extinction patterns of Permian

Brachiopoda of South China. Historical Biology 12, 93±110.

Sokol'skaya, A.N., 1965. Brachiopoda: Strophomenida. In: Ruzhensev,

V.E., Sarytcheva, T.G. (Eds.), Razvitie i smena morskikh organismov

na rubezhe paleozoia i mezozoia (the development and change of

marine organisms at the Palaeozoic±Mesozoic boundary). Akademia

Nauk SSSR, Palaeontologicheskii Institut, Trudy 108, pp. 203±209 (in

Russian).

Stehli, F.G., 1954. Lower Leonardian Brachiopoda of the Sierra

Diablo. Bulletin of the American Museum of Natural History

105, 257±358.

Tazawa, J., 1976. The Permian of Kesennuma, Kitakami Mountains: a

preliminary report. Earth Science (Chikyu Kagaku). Journal of the

Association for the Geological Collaboration in Japan 30 (3), 175±

185, 3 pls.

Tazawa, J., 1987. Kamiyase-chiku Kaseki Chyosa Houkoku-syo (the report

of the fossil survey in the Kamiyase area). Publication of the Kesennuma

Educational Committee, Miyagi Prefecture, pp. 1±37 (in Japanese).

Tazawa, J., Matsumoto, T., 1998. Middle Permian brachiopods from the

Oguradani Formation, Ise district, Hida Gaien Belt, Central Japan.

Science Reports of Niigata University, Series E (Geology), 13, 1±19

Termier, H., Termier, G., 1960. Contribution aÁ la classi®cation des Brachio-

podes: le lophophore des Collolophides nov. ord. Appendice: les Oldha-

minideÂs du Cambodge. SocieÂteÂ GeÂologique de France Bulletin, Series, 7

1 (3), 233±244, 2 pls. (in French).

Termier, H., Termier, G., 1970a. Le genre Spyridiophora (Brachiopodes

ProductoõÈdeÂs) dans le Permien asiatique. SocieÂteÂ GeÂologique du Nord,

Annales 90 (2), 57±62 1 pl. (in French).

Termier, H., Termier, G., 1970b. Les ProductoõÈdeÂs du Djoul®en (Permien

supeÂrieur) dans la TeÂthys orientale: essai sur l'agonie d'un phylum.

SocieÂteÂ GeÂologique du Nord, Annales 90 (4), 443±461, pls. 29±31

(in French with English abstract).

Tong, Z., 1978. Brachiopoda, Carboniferous±Permian. Palaeontological

Atlas of Southwestern China, Sichuan Province, 2. Geological Publish-

ing House, Beijing, pp. 10±267, pls. 77±92 (in Chinese).

Waagen, W., 1883. Salt Range Fossils: Productus Limestone fossils, Part 4

(2), Brachiopoda Palaeontologia Indica, Series 13 (1), 391±546.

Waagen, W., 1884. Salt Range Fossils: Productus Limestone fossils, Part

4(2) Brachiopoda. Palaeontologia Indica, Series 13 (1), 547±728.

Wang, C., 1995. Brachiopod fauna from the Kangkelin Formation in

Akesu, the Xinjiang Autonomous Region. Journal of Changchun

University of Earth Sciences 25 (1), 15±23, 4 pls. (in Chinese with

English abstract).

Wang C., Yang, S., 1993. Brachiopod fauna around the Carboniferous±

Permian boundary from the Balikelike Formation in Keping, Xinjiang.

Journal of Changchun University of Earth Sciences 23 (1), 1±9, 3 pls.

(in Chinese with English abstract).

Waterhouse, J.B., 1975. New Permian and Triassic brachiopod taxa. Papers

of Department of Geology, University of Queensland 7 (1), 1±23, pls.

1±2.

Waterhouse, J.B., 1976. Permian brachiopod correlations for South-east

Asia. Geological Society of Malaysia Bulletin 6, 187±210.

Waterhouse, J.B., 1978. Permian Brachiopoda and Mollusca from north-

west Nepal. Palaeontographica, Abt. A 160 (1±6), 1±175, 26 pls.

Waterhouse, J.B., 1981. Early Permian brachiopods from Ko Yao Noi and

near Krabi, southern Thailand. In: Waterhouse, J.B., Pitakpaivan, K.,

Mantajit, N. (Eds.), The Permian Stratigraphy and Palaeontology of

Southern Thailand. Geological Survey of Thailand (Memoir vol.

4(2)), pp. 43±213.

Waterhouse J.B., Piyasin, S., 1970. Mid-Permian brachiopods from Khao

Phrik, Thailand. Palaeontographica, Abt. A 135 (3±6), 83±197, pls.

14±32

Williams, A., Brunton, C.H.C., 1993. Role of shell structure in the classi-

®cation of the Orthotetidine brachiopods. Palaeontology 36 (4), 931±

966.

Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L.E., Popov, L.,

1996. A supra-ordinal classi®cation of the Brachiopoda Philosophical

Translations of the Royal Society of London. Biological Science 351,

1171±1193.

Yanagida, J., 1964. Permian brachiopods from central Thailand. Faculty of

Sciences, Kyushu University Memoirs, Series D. Geology 15 (1), 1±22,

pls. 1±3

Yanagida, J. Aw, P.C., 1979. Upper Carboniferous, Upper Permian and

Triassic brachiopods from Kelantan, Malaysia. Geology and Palaeon-

tology of Southeast Asia 20, 119±141, pls. 27±29

Zhang, Y., Fu, L., Ding, P., 1983. Phylum Brachiopoda. In: Palaeontolo-

gical Atlas of Northwest China (Shaanxi±Gansu±Ningxia Provinces),

Upper Palaeozoic 2. Geological Publishing House, pp. 244±425, pls.

88±143 (in Chinese).

Zeng, Q., 1987. Some structures and life habits of Urushtenoidea. Acta

Palaeontologica Sinica 26 (4), 497±503, 2 pls. (in Chinese with English

abstract).


middle permian brachiopods from central peninsular ...directory.umm.ac.id/data...

Documents