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Journal of Asian Earth Sciences 19 (2001) 177±194
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Middle Permian brachiopods from central Peninsular Malaysia Ð faunal
af®nities between Malaysia and west Cambodia
Masatoshi Sone a,1,*, Mohd Shafeea Leman b, Guang R. Shi a
a

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

b

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 brachiopod fauna from the so-called Central Basin (or the Central
Belt) of Peninsular Malaysia, and reports on the southernmost 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 laboratory 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 geological map of Peninsular Malaysia (Geological Survey of
Malaysia, 1985). The Semantan Formation, as revised,

outcrops west of the Bera Formation, which in turn is
* Corresponding author.
E-mail address: masasone@looksmart.com (M. Sone).
1
Present address: Institute for Environment and Development
(LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor,
Malaysia.

¯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 uncertain, but in the north it is associated with an andesitic volcanic 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-sediment 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,

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

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

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 principally of abundant lyttoniids in purple to black tuffaceous
mudstone and siltstone.
The brachiopods are listed in Table 1. The genus Transennatia 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 chronological standard. The time-scale compiled by the International 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 limestone hills with intercalations of tuff, shale and mudstone
strata, collectively called the Sisophon Limestone (Ishii et
Table 1
Brachiopod species from each assemblage of the Sungai Bera section
Species

Assemblages
A

Derbyia sp.
Urushtenoidea chaoi (Ching, 1963)
Urushtenoidea sp.
Transennatia termierorum sp. nov.
Transennatia sp. indet.
Spinomarginifera sp.
Echinauris sp.
Spyridiophora gubleri Termier and Termier, 1970a
Kozlowskia sp.
Phricodothyris sp.
Orthotetes aff. picta Fang, 1983
Strophalosiina sp.
Linoproductus sp.
Rhynchonellid family indet.
Orthothetina cf. iljinae Sokol'skaya, 1965
Gubleria aff. ninglangensis Fang and Jiang, 1992
Gubleria sp.

Eolyttonia? sp.
Spiriferinid family indet.

B

X
X
X
X

C

X

X
X
X
X
X
X

X
X
X
X
X
X
X
X
X

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

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).

al., 1969). Brachiopods from the
studied by several palaeontologists
1914; Chi-Thuan, 1961; Termier
1970a, 1970b). Ishii et al. (1969)

limestone have been
(e.g., Mansuy, 1913,
and Termier, 1960,
listed 50 brachiopod

179

species from their collections gained from that unit; Nakamura (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 biostratigraphic 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®nable for Member C of the Sisophon Limestone.
The most important species in the Bera fauna is Urushtenoidea 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. Transennatia 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-Caucasus and in the Maokou horizons of South China. In particular, 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 brachiopods 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

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

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. ˆ Limestone Member, S & S. ˆ 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).

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 conditions, 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 postKuhfengian 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. 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).

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

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 lyttoniids, 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). Lyttoniids 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 brachiopods have been described, despite many reported occurrences (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 biostratigraphic 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 (MuirWood, 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 correlations 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 limestones 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

181

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 differences 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®cation of Williams et al. (1996), and the classi®cations of
Williams and Brunton (1993) for the Strophomenida, Brunton 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
È pik, 1934
Order Strophomenida O
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 Ushaped; 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;

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

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

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 commissure 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 undistorted. 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

183

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 TransCaucasian 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. baschkirica 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.

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

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 arrangement 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 stronger in Urushtenoidea; its angle often exceeds 608. Urushtenoidea 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 correlated 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 Neoschwagerina 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 material, 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 Limestone 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,

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

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 conspeci®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 Urushtenoidea. 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 stereozone 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 Malaysian 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.

185

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 developed 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); interseptal spaces much wider than septal ridges, deeply grooved
along septal margins. Dorsal internal plates ¯at and long, laterally grooved along margins; median trough segmented irregularly. 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

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

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).

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

ranging 80±1058, and the pro®le is fairly ¯at. They may
belong to Gubleria aff. ninglangensis because of their similar septal apparatus. The present material, however, is much
wider and has greater umbonal angles. Specimen UKMF291 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, bladelike 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 transverse, 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 Spyridiophora ? 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;

187

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 reticulate; 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, extending 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 Spyridiophora ? 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 synonym and homonym of Productus timorensis Hamlet.
T. timorensis is close to T. termierorum in size and ornament. 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 Archbold and Bird (1989, p. 108). For instance, the Kasliu form
of Archbold and Bird (1989, p. 107, ®gs. 3E±I) is exceptionally large and has ®ne ornament, whereas the BasleoWesleo 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 Chitichun 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 southern 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 geniculate. 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

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

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

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 information 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 panlike appearance with a low and short median septum indicates Echinauris. As common in the genus, brachial ridges
are not de®ned. The marginal ridge is moderate. The external 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

189

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, UKMF266, 5±8, dorsal external mould with accompanying natural cast of shell remaining anteriorly Ð posterior, ventral, anterior, and lateral views, £ 3.0, UKMF267, 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.

190

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

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.

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

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 illustrations. 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 P