Gondwana Research, %! 6, No. 4, pp. 777-790.

Gondwana
0 2003 International Association for Gondwana Research, Japan.
ISSN: 1342-937X GR
GRGiZ , Research

Global Review of Permian TyZopZecta Muir-Wood and Cooper,

1960 (Brachiopoda): Morphology, Palaeobiogeographical and
Palaeogeographical Implications
G.R. Shil* and Z.Q. Chen2
School of Ecology and Environment, Deakin University, Melbourne Campus, 221 Burwood Highway, Burwood, Victoria
3125, Australia, E-mail: grshi@deakin.edu.au.au
Institute of Geology and Paleontology, Tohoku University, Aoba, Ararnaki, Sendai 980-8578, Japan

* Corresponding author

(Manuscript received July 22,2002; accepted December 21, 2002)

Abstract
A global review of the stratigraphical and geographical distribution of TyZoplecta reveals that the genus ranges in age
from Kungurian to Changhsingian (Middle to Late Permian). Tyloplecta first evolved in South China in the Kungurian
(late Early Permian). The genus went through its first diversification in the Guadalupian, suffered a major extinction at
the end of the Guadalupian, and re-diversified in the Wuchiapingian. ?: yangtzeensis persisted into the Changhsingian
as the only survivor of the genus involved in the end-Permian mass extinction. Palaeogeographically, South China is not
only the centre of origin for the genus but also an area of diversification and evolution. In addition to South China,
Tyloplecta has also been recorded from the Far East Russia, Japan, central Thailand, Laos, Cambodia, Qiangtang Terrane
of Tibet, Salt Range, Iran, Armenia, Hungary, Yugoslavia, and Slovenia. This geographic spread suggests that Tyloplecta
was primarily restricted to the Palaeotethys and is indicative of warm-water palaeoequatorial conditions. Its presence
in some of the northeast Asian terranes (e.g., parts of Japan and Far East Russia) and in the Salt Range (Pakistan) and
central and north Iran (part of the Cimmerian microcontinents) demonstrate that the genus invaded the middle
palaeolatitudinal regions in both hemispheres during the late Middle Permian in response to increased shallow marine
biotic communications between Cathaysia in the eastern Palaeotethys and southern Angaraland, and between Cathaysia
and Peri-Gondwanaland. The invasion of Tyloplecta (and some other taxa) into the southern shore waters of Angaraland
may be explained by assuming ocean surface current connections and close palaeogeographical proximities between
the South China, Sino-Korea and Bureya blocks. In comparison, the invasion of Tyloplecta into the Peri-Gondwanaland
region is more likely a result of reduced palaeogeographical distance between South China and Peri-Gondwanaland
and the appearance of the Cimmerian microcontinents as migratory stepping stones.
Key words: Brachiopoda, palaeobiogeography, Palaeotethys, Permian, Tyloplecta.

Introduction updated review on the global stratigraphical and

The Permian brachiopod productid genus Tyloplecta
Muir-Wood and Cooper, 1960 is among the most
characteristic genera that distinguish the Permian
Palaeoequatorial Realm, the Cathaysian Province in
particular, from the Gondwanan Realm to the south and
Boreal Realm to the north (Nakamura et al., 1985; Tazawa,
1991; Shi e t al., 1995). Biostratigraphically and
biogeographically the genus is of particular interest because
of its relatively short stratigraphic range, distinctive
morphological characteristics, and its common association
with typical warm-water, shallow marine faunas.
In this paper, we attempt to provide a detailed and
geographical distribution of Tyloplecta based on a refined
understanding of the taxon’s morphology and
classification. The palaeobiogeographical and
palaeogeographical implications of the genus’ global
distribution will be discussed in the broad context of
Permian marine biogeography of the Palaeotethys as a
means to elucidating the relationship between biotic
distribution and disposition of microcontinents (terranes)
in the Permian Palaeotethys. Material used for this review
includes our personal collections from South China, Japan,
and Far East Russia, as well as type collections housed at
the Nanjing Institute of Geology and Palaeontology,
Chinese Academy of Sciences, Nanjing, China.
778 G.R. SHI AND Z.Q. CHEN
Morphology and Classification of TyZopZecta
Although Tyloplecta species have been described
since 1911 (the genus was proposed by Muir-Wood and
Cooper in 1960), and the genus is morphologically
distinctive, its systematic position is still a matter of debate.
Muir-Wood and Cooper (1960) referred it to the
Dictyoclostidae Stehli when they erected the genus. Later,
Termier and Termier (1970) established a new family,
Tyloplectidae, for Tyloplecta in recognition of the presence
of numerous distinctive nodes on the ventral umbo.
However, Termier and Termier’s proposition was not
followed by subsequent workers, who continued to refer
Tyloplecta to the Dictyoclostidae (e.g., Zhang and Ching,
1976; Jin and Hu, 1978; Jin and Sun, 1981; Nakamura
et al., 1981; Sremac, 1986, Shen et al., 1992). On the
other hand, in a major effort to revise the classification
of productoids, Lazarev (1990) emphasised t h e
significance of weak ribbing confined to the areas anterior
to the umbos in Tyloplecta and accordingly placed
this genus in the Leioproductidae Muir-Wood and
Cooper, 1960. This scheme was later closely followed by
Brunton et al. (1995, 2000), who (Brunton et al., 2000)
assigned Tyloplecta to the Tyloplectini tribe within the
Leioproductinae subfamily.
Brunton et al. (2000, p. 485) diagnosed Tyloplecta as
having elongate spine bases posteriorly that become ribs
at the midlength of the corpus. However, our inspection
of the lectotype of the type species housed in the Nanjing
Institute of Geology a n d Palaeontology, Nanjing,
contradicts this observation. The lectotype shows that the
costae commence anterior to the beak (also see Brunton
et al., 2000, fig. 329b) and extends anteriorly to the trail;
on the ventral valve the costae are interrupted by
concentric rugae giving conspicuous nodes (Fig. 1A-B).
Spines are scattered on the costae but are usually
considerably narrower in diameter than costae (see
Brunton et al. 2000, fig. 329b), and the strength of costae
is consistent from the start of the protrail to anterior
margins. Other diagnostic features of the genus include
semi-costate shell (Fig. 1A-B), strongly pustulose and finely
capillate dorsal valve, spines confined to the ventral valve
with usually much coarser ones along the hinge margin
and in clusters on ears. Internally, Tyloplecta possesses a
Dictyoclostus-type cardinal process that is bilobed internally
and quadrilobed externally and supported by a broad shaft
(Fig. 1E-F); its strongly dendritic ventral muscle scars and
strong median septum, highly elevated and dendritic
adductor scars and highly ridged, hook-like brachial ridges
of the dorsal valve are also notable (Fig. 1E-F). Within
the ventral valve, the adductor scars sit on a raised
dendritic platform and diductor scars deeply depressed
below the valve floor and radially grooved (Fig. 1C-D).
Brunton et al. (2000) placed Araxilevis Sarycheva, 1965
and Pseudoantiquatonia Zhan and Wu, 1982 in the same
tribe, Tyloplectini, with Tyloplecta. But this is questionable.
Productus intermedius Abich, 1878, the type species of
Araxilevis, is strongly lamellous and lacks costae, thus very
different from t h e strongly ribbed Tyloplecta.
Pseudoantiquatonia mutabilis Zhan and Wu (1982, p. 98,
pl. 3, figs. 1-4, 8-19), also figured recently by Shi and
Shen (2000, pl. 1,figs. 7-17; pl. 2, figs. 1-9), on the other
hand, is rather finely costate on both valves with regularly
and well-defined reticulation on visceral disks, and lacks
capillae on dorsal costae and nodes on the umbonal region
of the ventral valve, both of which are otherwise typical
of Tyloplecta.
Chen and Shi (2000) compared genera of the
Liraplectini with Tyloplecta, and concluded that Tyloplecta
can be distinguished from Liraplecta Jin and Sun, 1981
and Tarimoplecta Chen and Shi, 2000 by its conspicuous
nodes on the umbonal region of the ventral valve. Kepingia
Wang and Yang, 1998, is also dorsally capillaed like
Tyloplecta, but is distinguishable from the latter by its
narrower ribs, pronounced concentric growth lines over
the whole shell, and strong reticulation on the disks.
Global Distribution of TyZopZecta
At present, twenty species of Tyloplecta are known from
around the world (Fig. 2, Table 1).Below we provide a
detailed review on the stratigraphical and geographical
distributions of these species in light of the refined
definition for the genus given above.
East Asia
In the Far East of Russia, ?: yangtzeensis (Chao) was
first reported as such by Licharew and Kotlyar (1978) from
the Chandalaz Formation in South Primorye, where it is
associated with Parafusulina stricta and Neomisellina
Zepida-Lepidolina kumaensis Zones of Capitanian age
(Kotlyar et al., 1999). In addition, Kotlyar et al. (1992)
have also reported T. yangtzeensis from a reef complex at
Nakhodka of the same region, where it is found in
association with Edriosteges poyangensis (Kayser),
Strophalosiina tibetica (Diener), Leptodus nobilis
(Waagen), Permophricodothyris grandis (Chao),
Chenxianoproductus nachodkensis Kotlyar, Anidanthus
sinosus Huang, Stenoscisma margaritovi (Huang), Peltichia
nachodkensis Kotlyar, Spinomarginifera alpha Huang,
and Caucasoproductus primoricus Kotlyar. The age of the
reef complex, as indicated by the reef-building and other
organisms (brachiopods, ammonoids, corals, and
conodonts), is Wuchiapingian according to Zakharov et
al. (1997), although a possible extension to as low as
Capitanian cannot be ruled out (see Kotlyar et al., 1992).
Gondzuann Research, V. 6, No. 4, 2003
 PERMIAN TnOPLECTA MUIR-WOOD AND COOPER, 1960 (BRACHIOPODA) 779

In Japan, Nakamura (1960) first reported Tyloplecta
(as Dictyoclostus) from the lower Kanokura Series of
the South Kitakami Belt, northeastern Japan, where it
coexists with a large mixed BorealKathaysian brachiopod
fauna (Tazawa, 1991, 1998, 2000; Tazawa and Ibaraki,
2001) and is, at some localities, directly associated with
the Monodiexodina matsubaishi Zone. Tazawa and Ibaraki
(2001) suggest a Kubergandian-Murgabian (?Roadian-
Wordian) age for the brachiopods, while Shi (in Shi and
Tazawa, 2001) preferred a general Wordian age for
this and other similarly mixed Middle Permian brachiopod
faunas in Japan, northeast China and Far East Russia.
This age assignment appears consistent with the age
indication from ammonoids from the same formation
(Ehiro, 1998).
South China appears to be the centre of origin and
diversification for Tyloplecta, with 13 species having been
described from the region. Here, the oldest species of the
genus, ?: nankingensis, was originally reported from the
basal Chihsia Formation (Kungurian) near Nanjing, South
China (Chao, 1927), but has since been found widely from
stratigraphic horizons equivalent to the lower Chihsia
Formation in many areas across South China (see Table 1
and references therein provided). The associated

Fig. 1. Morphology of Tyloplecta illustrated in both reconstructed drawings and real specimens as comparison. A-B, ventral exterior showing
the distinctive nodes on the umbonal region formed by the intersection of costae and rugae; A, a drawing based on Z yangtzeensis (Chao),
x 1.4; B, a ventral valve (NMV P309578) in ventro-anterior view from the Wuchiapingian Lungtan Formation in the Daijiagou section,
Beipei of Chongqing City, South China (for section and location details see Zeng et al., 1995), showing conspicuous nodes, X 0.85 C-D,
ventral interior of Z nankingensis (Frech) showing the details of the muscle field; C, reconstructed drawing based on specimen NMV
P309579; D, an internal mould of a ventral valve in ventro-posterior view, lower Chihsia Formation in Daijiagou section of Beipei, Chongqing
City, South China (for section and location details, see Liu et al., 1982), x 1.2 E-F, a reconstructed drawing and a dorsal interior of 'I:
nankingensis (Frech) (after Muir-Wood and Cooper, 1960, pl. 102, fig. 10; USNM124015), showing the details of the dorsal internal
structures, both X 1.2 AD - adductor scars; BR - brachial ridges; CP - cardinal process; LR - lateral ridges; MS - median septum.

Gondwana Reseauch, V. 6, No. 4, 2003

780 G.R. SHI AND Z.Q. CHEN
brachiopod genera usually include Liraplecta, Orthotichia,
Ogbinia, and Chaoina.
The upper Chihsia Formation (Roadian) and the
overlying Maokou Formation (Wordian) in South China
are characterised by the abundant appearance of T.
nankingensis, 7: grandicostata, T crassiplicata, T. magna,
T. minisulcata, and T.yangtzeensis. Of these the first two
continued from the Kungurian, while T. yangtzeensis
extends into the overlying Capitanian-Changhsingian
(Fig. 3). The remaining three species are very rich in the
Wordian and also persist into the Capitanian (Fig. 3). These
Roadian-Wordian Tyloplecta are typically associated with
many other brachiopod genera including Neoplicatifera,
Monticulifera, Cryptospirifer, Titanothyris, and Uncisteges.
The Capitanian of South China is represented by the
Lengwuan (Capitanian) reef complex, situated at the
Lengwu area of Zhejiang, eastern part of South China (Liang,
1990). So far, only T.yangtzeensis has been confirmed to
be present in this stage of South China (Table 2).
The Wuchiapingian brachiopod fauna of South China
is most diverse and abundant among its Late Permian
(Lopingian) marine faunas. There are three Wuchiapingian
Tyloplecta species, all found from the Lungtan (Longtan)
coal series (= Lungtan Formation): T. yangtzeensis, an
index species of the Lungtan Formation, ‘Ibulongensis,
and ‘Iyichunensis (Table 2). These Tyloplecta species are
usually associated with other brachiopods such as
Orthotetina frechi (Huang), Tethyochonetes soochowensis
(Ch ao) , ‘I w o ngia n a (Chao) , Edrios teges p oya nge ns is
(Kayser), Haydenella wenganensis (Huang),
Spinomurginijera lopingensis (Kayser), Leptodus nobilis
(Waagen), Gubleria huangi Wang, Oldhamina grandis
Huang, and Permophricodothyris grandis (Chao).
Although the Changhsingian of South China contains
a rich brachiopod fauna (Shen et al., 2000)) Tyloplecta
is relatively rare with only one species, T. yangtzeensis,
representing this stage (Liao, 1987; Shen et al., 1992).
Associated with this brachiopod are other diagnostic
Changhsingian brachiopods including Peltichia zigzag
Huang, P. sinensis (Huang), Perigeyerella costellata
Wang, Spinomarginifera kueichowensis Huang, S. alpha
Huang, Huatangia sulcatifera Liao and Meng, and
Permophricodothyris squamularioides (Huang) .
From the Qianghai-Tibetan Plateau of western
China, Ting (1965) reported both T. yangtzeensis and
T.nankingensis from the Bayinhe Group of the Yangkang
Valley, Tienchung district, Qinghai Province. The associated
brachiopods include Streptorhynchus lenticularis Waagen,
S. cyrtoides Ting , Meekella substriatocostata Yang and
Chang, Tenuichonetes minor (Ting), Spinomarginifera
pseudosintanensis Huang, Monticulifera plicatiformis Ting
, and Chilianshania chilianshanensis Ting. All of these are
also present in the Maokouan assemblages of South China
(Jin and Hu, 1978; Wang et al., 1982), and thus indicate
a Kuhfengian (Wordian) age.
Jin and Sun (1981, p. 137, pl. 3, Figs. 1-6, 9-11)
described Tyloplecta crassica Jin and Sun from the Upper
Permian Yinba Formation in the Jiage area of Mangkong
County and the Yinbao area of Changdu County, eastern
Tibet. This eastern Tibetan species closely resembles T.
pseudorossica Archbold, 1979 [ = T. rossica (Licharew,
1937))see below] from the Upper Permian of the northern
Caucasus in many respects, but the Caucasian species is
more transverse and possesses a nearly quadrate outline
and more pronounced reticulations on the umbonal
regions of both valves. The associated brachiopods from
the Yinbao Formation include Tethyochonetes convexa
(Yang and Fan), Peltichia sinensis Huang, Oldhamina
decipiens (Koninck), Leptodus t e n u i s (Waagen),
Spinomarginifera? lopingensis (Chao), Haydenella
kiangsiensis (Kayser), Cathaysia chonetoides (Chao),
“Productus” caucasica Licharew, Permophricodothyris
supefba (Jin and Sun), and Cartorhium mahaensis
(Huang). The fauna is overall comparable with those from
Fig. 2. Geographical distribution of
Tyloplecta Muir-Wood and Cooper.
1-South Primorye, Far East of
Russia; 2-South Kitakami
Mountains, Japan; 3-South China;
4-Qinghai-Tibet Plateau; 5-central
Thailand; 6-Laos; 7-western
Cambodia; 8-western Thailand;
9-Salt Range, Pakistan; 10-Abedeh
region of central Iran; 11-northern
Iran; 12-southern Armenia;
13-Hungary; 14-Yugoslovia;
15-Slovenia.
Gondwana Research, K 6, No. 4, 2003
 PERMIAN TYLOPLECTA MUIR-WOOD AND COOPER, 1960 (BRACHIOPODA) 78 1

Tablc 1. Dctailcd stratinrauhical and rreorrrauhical distributions of all Tvloalecta soccics.

Spccics Rcfcrcnccs Formation LOC.in
Fin. 2
7: nankingensis (Frcch) Frcch, 1911; Chao, 1927; Chihsia Fm. Kungurian 3
Wang ct al., 1982; Liu et al., 1982;
Fong and Jiang, 1978
T. nankingensis Yang, 1984 Xintan Fm. Wordian 3
T nankingensis Ding and Qi, 1983 Shuixiakou Fm Kungurian 3
T nankingensis Jin and Hu, 1978; Wang et al., 1982 Kuhfen Fm. Wordian 3
7: nankingensis Huang, 1932, 1933; Jin and Liao, 1974; Maokou Fin. Wordian 3
Tong, 1978; Zcng ct al., 1995
7: nankingensis Ting, 1965 Bayinhe Group Wordian 4
T. nankingensis Mansuy, 1914; Ishii et al., 1969; Sisophon Lst. Kung.-Wordian 7
Tcrmicr and Tcrmicr, 1970
‘r: yangtzeensis (Chao) Licharew and Kotlyar, 1978; Chandalaz Fm. Wordian-Wuchiapingian 1
Kotlyar et al., 1992
7: yangtzeensis Nakamura, 1960; Tazawa, 1976; Kanokura Series Guadalupian 2
Nakamura et al., 1981
T.yangtzeensis Kayser, 1882; Frcch, 1911; Chao, 1927; Lungtan Fm. Wuchiapingian 3
Huang, 1932, Wang, 1955;
Wang ct al., 1964; Yang ct al., 1977;
Fong and Jiang, 1978; Tong, 1978;
Zhan, 1979; 1989; Liao, 1980;
Liu ct al., 1982; Yang, 1984; Xu, 1987;
Zcng ct al., 1995
T yangtzeensis Wang ct al., 1982 Loping Fm. Wuchiapingian
T. yangtzeensis Liao, 1987 Heshan Fm. Wuchiapingian
I: yangtzeensis Ting, 1965 Uppcr Pcrmian Wuchiapingian
7: yangtzeensis Ding and Qi, 1983 Xikou Fm. Wuchiapingian
T yangtzmisis Liang, 1990 Lcngwu Fin. Capitanian
T.yangtzeensis Yang, 1984; Liao and Mcng, 1986;
Shcn ct al., 1992; Shen and He, 1994 Changhhsing Fm. Changhsingian 3
Z yangtzeensis Liao, 1980 Wuanwei Fm. Changhsingian 3
T.yangtzeensis Liao, 1987 Talung Fm. Changhsingian 3
7: yangtzeensis Kotlyar ct al., 1999 Nikitin Formation Changhsingian 11
T yangtzeensis Ishii c t al., 1969 Sisophon Lst. Wuchiapingian 5
7: yangtzeensis Yanagida, 1964; Nakamura et al., 1981 Phctchabun Bed Wuchiapingian 9
T. yangtzeensis Reed, 1944; Watcrhousc and Gupta, 1983 Wargal Fm. Capitanian 7
T cf yangtzeensis Fantini Sestini, 1965 Rutch Fm. Guadalupian 11
T.yangtzeensis Fantini Sestini and Glaus, 1966 Upper Nesen Fm. Wuchiapingian 11
T.yangtzeensis Nakamura ct al., 1981 Unit 4 Guadalupian 11
T yangtzeensis Nakamuria ct al., 1981 Unit 6 Wuchiapingian 11
7: yangtzeensis Miklukho-Maklay, 1954; Sarychcva, 1965 Kutan Fm. Wuchiapingian 112
..
I . ya ngt ze ens is Schrtter, 1963; Pesic et al., 1986 Nagyvisnyo Lst. Wuchiapingian 13
T.yangtzeensis Simic, 1933; Stojanovic-Kuzcnko, 1965; Jadar Development Wuchiapingian 14
Pesic and Sremac, 1986
T.yangtzeensis Ramovs, 1958; Pesic ct al., 1986; Zazar Fm. Wuchiapingian 15
Buscr ct al., 1986
T calloci-enea (Hcritsch) Ramovs, 1958; Pesic et al., 1986; Zazar Fm. Wuchiapingian 15
Buser et al., 1986
T. cailocrenea SchrCtcr, 1963; Pcsic ct al., 1986 Nagyvisnyo Lst. Wuchiapingian 13
T callocrmiea Simic, 1933; Stojanovic-Kuzcnko, 1965; Jadar Development Wuchiapingian 14
Pcsic and Sremac, 1986
T. alpina (Simic) Simic, 1933; Stojanovic-Kuzcnko, 1965; Zadar Fm and Jadar Wuchiapingian 14, 15
Pcsic and Sremac, 1986; Ramovs, 1958;
Pcsic ct al., 1986; Buser et al., 1986
Z Lurazi Nakamura ct al. Nakamura et al., 1981 Units 4 and 6 Capitanian-Wuchiapingian 11
T. calnbodgiensis (Mansuy) Mansuy, 1913 Basal Permian Roadian 6
7: rcdacta (liccd) Rccd, 1944; Waterhousc and Gupta, 1983 Chhidru Fm. Wuchiapingian 9

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782 G.R. SHI AND Z.Q. CHEN

Species References Formation LOC. in

Fin. 2

T. crassica Jin and Sun Jin and Sun, 1981 Tuoba Fm. Wuchiapingian 4
T. bulongensis Liao Liao, 1980; Zeng et al., 1995 Lungtan Fm. Wuchiapingian 3
T. persicn Fantini Sestini Fantini Sestini and Upper Nesen Fm. Wuchiapingian 11
and Glaus Glaus, 1966
T. adurnbrata (Reed) Reed, 1944; Waterhouse and Gupta, 1983 Chhidru Fm. Wuchiapingian 9
T. tuiiiefacta (Reed) Reed, 1944; Watcrhouse and Gupta, 1983 Chhidru Fm. Wuchiapingian 9
T. pseudorossica Licharew, 1937; Jin et al., 1979; Dzhulfa Fm. and
Archbold, 1979 Koltyar et al., 1989 Upper Permian Wuchiapingian 12, 4
T. crussiplicata Fong Fong and Jiang, 1978; Zeng et al., 1995 Maokou Fm. Wordian 3
T. liziyuensis Zeng Zeng et al., 1995 Chihsia Fm. Kungurian 3
T . rninisulcata Zeng Zeng et al., 1995 Maokou Fm. Wordian 3
T. rnagna Tong Tong, 1978 Maokou Fm. Wordian 3
T. grandicostata (Chao) Chao, 1927; Fong and Jiang, 1978; Chihsia and
Liu et al., 1982; Wang et al., 1982; Maokou Fms Kung.-Wordian 3
Yang, 1984; Zeng et al., 1995
T. yichunerisis Jin and Hu Wang et al., 1982 Loping Fm. Wuchiapingian 3
Sloplecta sp. Personal field observation Ratburi Limestone G u a d a 1up ia n 8

the P,a bed of northern Caucasia (Licharew, 1937) and
the lower Wuchiapingian faunas of South China (Jin and
Sun, 1981) (Table 2).
Southeast Asia (central Thailand, Cambodia, Laos,
Indonesia, Timorf
Yanagida ( 1 9 6 4 ) illustrated several Tyloplecta
specimens, later assigned to T.yangtzeensis by Nakamura
et al. (1981), from the Phetchabun Formation in central
Thailand (Indo-China block). At that time, Yanagida referred
the Phetchabun fauna to the Wordian, but later Grant
(1976) reassigned the fauna to the Lopingian (Table 2).
In West Cambodia, Mansuy (1914, pl. 18, fig. 6)
illustrated a shell fragment as Productus cf. sumatrensis
Koemer from the Sisophon Limestone. Grant (1976)
thought that this species could belong to Stereochia, but
the same specimen, as later also illustrated by Chi-Thuan
(1961, pl. 5, Fig. 7), shows that it bears conspicuous nodes
on the umbonal region, therefore suggestive of Tyloplecta.
Ishii et al. (1969) subdivided the Sisophon Limestone into
A-D beds, and listed T.nankingensis from Beds A-B and T.
cf. yangtzeensis from Bed C. Associated with T. nankingensis
are fusulinaceans characteristic of the Pseudofusulina
pseudolepida-Pseudofusulinaambigua pursatensis Zone and
the Surnatrina longissima-Yabeina asiatica Zone, of
Roadian to Wordian age (Sone et al., 2001). The Bed C,
bearing T. cf. yangtzeensis, contains a highly diverse
brachiopod fauna, with 36 species in 32 genera. This
brachiopod fauna, subject to future revision, appears
closest to the Wuchiapingian brachiopod faunas of South
China. However, the associated fusulinacean fauna,
assignable to the Yabeina Zone, suggests a slightly older
Capitanian age (Sone et al., 2001).
Termier and Termier (1970) also reported a diverse
fauna from the Sisophon Limestone of the same area,
including fusulinaceans: Yabeina cf. khmeriana, Verbeekina
pseudoverbeeki; b r a c h i o p o d s : T. nankingensis,
Tsch e r nys c h ew ia typ ica S t oy a n ov, Ec h i n au r is op u n t ia
(Waagen), Marginifera typica (Waagen), Costiferina spiralis
(Waagen), and rugose corals: Wentzelella sp. The fauna
has been correlated with the lower Dzhulfian of the
Transcaucasia, the lower Wuchiapingian of South China,
and the topmost Wargal Formation to Chhidru Formation
of the Salt Range, Pakistan.
In central Laos, Mansuy (1913) described Productus
sumatrensis mut. cambodgiensis Mansuy from the “Level
1” of the Permian, possibly equivalent to the Roadian-
Wordian in view of faunal composition. This Laotian form
was recently upgraded to species status and converted to
Tyloplecta by Shi and Archbold (1993).
The presence of Tyloplecta in the Indonesia-Timor region
requires clarification. Muir-Wood and Cooper (1960)
assigned Productus sumatrensis Roemer (1880; see also
Fliegel, 1901; Lang, 1925) from the Verbeekina verbeeki
Zone of Padang, west-central Sumatra and eastern
Peninsular Malaysia (Meyer, 1922) to Tyloplecta. But
Archbold (1985) placed this species in Costiferina Muir-
Wood and Cooper, 1960. More recently, Shi and Archbold
(1993) transferred the species to Stereochia Grant, 1976
on account of its well-defined reticulation, instead of
nodes, on the umbonal region of the ventral valve.
Comparable material from the Bitauni bed of Timor (Broili,
1916; Hamlet, 1928; Shimizu, 1966) and Irian Jaya
(Archbold, 1981) fits well within Costiferina or Stereochia.
Consequently, no true Tyloplecta occurs in the Indonesia-
Timor region.

Goiidzunna Research, V. 6, No. 4, 2003

 PERMIAN TYLOPLECTA MUIR-WOOD AND COOPER, 1960 (BRACHIOPODA)
6 Stage
1
I
'riassic
Zhangh .p
-singian
Wuchia-
pingian
Capi-
tanian
-
Wordian '5
Roadian
gurian
Species number
Pakistan and Iran
Reed (1944) identified two ventral valves respectively
as Productus (Dictyoclostus) yangtzeensis (Chao) var.
adumbrata Reed (1944, p. 47, pl. 9, Fig. 2) and P. (D.)
yangtzeensis tumefacta Reed (1944, p. 48, pl. 10, Fig. 5),
both from the Chhidru Formation of the Salt Range,
Pakistan. These two subspecies were later assigned to
Tyloplecta by Waterhouse and Gupta (1983, p. 232). The
lower and middle parts of the Chhidru Formation is usually
assigned to the Wuchiapingian in light of its conodont
and foraminifera1 zones (Wardlaw and Pogue, 1995;
Mertmann and Sarfraz, 2000) (Table 2).
Waterhouse and Gupta (1983) also listed T. yangtzeensis
in their revision lists of the faunas described by Reed
(1944) from the Wargal Formation. The Wargal
brachiopod fauna is similar to that of the overlying Chhidru
Formation, with many genera and species shared,
including such common genera as Waagenites, Sulcataria,
Chonetella, Waagenoconcha, Costiferina, Spiriferella,
Whitspakia, Compressoproductus, and Richthofenia. The
Wargal Formation has been dated as Capitanian by
fusulinaceans (Nakazawa et al., 1975) and conodonts
(Wardlaw and Pogue, 1995).
Tyloplecta is known to appear in the Tyloplecta Zone of
the upper Nesen and Ruteh formations of the Alborz
(Elburz) Mountains, northern Iran (Fantini Sestini, 1965;
Fantini Sestini and Glaus, 1966), and in Unit 4 (Abadeh
Formation) and Unit 6 (Hambast Formation) of the
Gondwana Reseauch, V. 6, No. 4, 2003
783
IEnd-Permian
Fig. 3. Stratigraphical distribution
and diversity patterns of
Tyloplecta Muir-Wood and
Cooper. Dashed lines
indicate possible occurrences
of species in these intervals
but are yet to be confirmed.
Abadeh region in central Iran (Taraz, 1973; Nakamura et
al., 1981) (Table 2). In the former (upper Nesen
Formation), T. yangtzeensis and T. persica are associated
with Acosarina dorashamensis Sokolskaya, Enteletes cf.
dzhagrensis Sokolskaya, Orthotetina peregrina (Abich),
Spinomarginifera ciliata (Arthaber), S. helica (Abich), S .
spinoscostata (Abich), Permophricodothyris iranica Sestini
and Glaus, I? ovata Pavlova, Armathyris araxensis Grunt,
A. f e l i n a (Arthaber), and A. protea (Abich). This
brachiopod fauna, referred to the Tyloplecta Zone by
Fantini Sestini and Glaus (1966), can be closely correlated
with the brachiopod fauna from the lower Dzhulfian of
southern Armenia (Licharew, 1937; Sarycheva, 1965). In
central Iran, the Abadeh Formation contains a diverse
brachiopod fauna as well as corals, fusulinaceans and
conodonts, all indicating a Capitanian age (Baghbani,
1997; Taraz, 1999). The overlying Hambast Formation
also yields a diverse fauna with especially abundant
ammonoids and moderately rich brachiopods including
Tyloplecta tarazi and i? yangtzensis (Iranian-Japanese
Research Group, 1981; Nakamura et al., 1981). This fauna
is closely comparable with the Wuchiapingian faunas of
South China or the Dzhulfian faunas of Armenia
(Nakamura et al., 1981; Baghbani, 1997; Taraz, 1999;
see also Table 2).
Caucasus Mountains (Armenia)
In this region, Tyloplecta pseudorossica Archbold,
784 G.R. SHI AND Z.Q. CHEN
~
originally described by Licharew (1937, p. 41-43, pl. 2,
Figs. 1-7; pl. 3, Figs. 1-4) as Productus yangtzeensis var.
rossica Licharew, occurs in the P,” bed of the Dzhulfa
Formation. The age of the P,” bed is Early Dzhulfian (Early
Wuchiapingian) (Waterhouse, 1976; Kotlyar et al., 1989).
In addition, Sarycheva (1965) reported I: yangtzeensis
from the Vedioceras-Haydenella beds of the upper Araksian
(Early Dzhulfian) in the Dorashamian stratotype section
in the Ogbin area, Dorasham, Transcaucasus.
In the northern Caucasus, T. yangtzeensis was originally
reported from the Nikitin Formation by Miklukho-Maklay
(1954). Although Nakamura et al. (1981) assigned the
Nikitin fauna to the early Wuchiapingian, other authors
(e.g., Kotlyar et al., 1989; Pronina-Nestell and Nestell,
2001) dated the fauna as late Changhsingian based on
small foraminifers and fusulinaceans.
Eastern Europe
In the Bukk Mountains of Hungary, Schrkter (1963)
reported T. yangtzeensis and T. callocrenea (Heritsch) from
the Horizon 2 of the Nagyvisnyo Limestone (Pesic et al.,
1986). Overall, the age of the Nagyvisnyo Limestone has
been considered early Dzhulfian (Wuchiapingian) by its
combined coral, brachiopod and conodont faunas
(Schrkter, 1963; Pesic et al., 1986; Kozur, 1988).
Three species of Tyloplecta are known from the Lower
Dzhulfian of western Serbia (Simic, 1933; Stojanovic-
Kuzenko, 1965; Pesic and Sremac, 1986), including
Tyloplecta yangtzeensis, ?: callocrenea (Heritsch), and
Table 2. World correlation of Permian Qloplecta-bearing strata.
Ei:aii::F
Far
J~~~~ 1 1 I
South Qinghai Indo- Salt North Central South Hungary Yugo- Slovenii
China -Tibet ChinalRmge Iran Iran Amenia
Fm.
? Tyloplecta-bearinghorizons
T. alpina (Simic) (Table 1). From Croatia, Sremac (1986)
described a ventral valve as Tyloplecta sp. from the Middle
Permian of the Velebit Mountains, but the true identity of
this valve requires further study as it is relatively small
(34 mm long and 31.9 mm wide), has a rather narrow
umbonal region (about a half of its valve width), small
ears, and is sculptured only with three broad costae in the
shallow median sulcus. Several large spine bases are present,
but umbonal nodes are absent, unlike typical Tyloplecta.
Three species, I: yangtzeensis, T. callocrenea, and T.
alpina, are known to occur in the forth horizon of the
Upper Permian Zazar Formation of Slovenia (Ramovs,
1958; Pesic et al., 1986; Buser et al., 1986). These
Tyloplecta species occur together with early Dzhulfian
brachiopods of the Iran-Armenia region and the
Wuchiapingian brachiopod faunas of South China (Buser
et al., 1986).
Diversity and Evolutionary Trends of
Tyloplecta
As outlined above and summarised in table 1and figure
3, three species, 7: nankingensis, T. grandicostata and T.
liziyaensis, occur widely within the geographic range of
this genus. The oldest species of the genus, T. nunkingensis,
began in the basal Chihsia Formation (earliest Kungurian)
in South China. Both T. nankingensis and T grandicostata
persist into the Guadalupian, which also saw the first
appearance of six new Tyloplecta species: T. cambodgiensis,
1 I 1
slavia
Gondwana Research, V. 6, No. 4, 2003
 PERMIAN TYLOPLECTA MUIR-WOOD AND COOPER, 1960 (RRACHIOPODA)
?: magna, T. minsulcata, T. yarrgtzeensis, T. crassiplicata,
and T. tarazi (Fig. 3 ) . Only two Middle Permian Tyloplecta
species, T. yangtzeensis and T. tarazi, extend into the
Lopingian. This means that 6 (out of 8, or 75%)
Guadalupian Tyloplecta species became extinct by the
beginning of the Wuchiapingian (Fig. 3). Across the
Guadalupian-Lopingian boundary, Tyloplecta became
strongly diversified in the Wuchiapingian, with eleven new
species (Fig. 3). It was also during this period that T.
yangtzeensis reached its acme and became the most
widespread species of the genus. After the Wuchiapingian
diversification, the diversity of Tyloplecta dropped rapidly
with only one species, 7: yangtzeensis, surviving into the
Changhsingian (Fig. 3).
Palaeobiogeographical Patterns
No species of Tyloplecta are known from pre-Kungurian
strata any where around the world. Three species of
Tyloplecta occurred in the Kungurian of South China, all
within the Cathaysian Province of the warm-water
Palaeoequatorial Realm (Fig. 4A), suggesting that South
China could be the centre of origin for the genus.
In the Guadalupian (Fig. 4B), Tyloplecta diversified in
South China and spread to other regions. Six of the eight
known Guadalupian species from around the world occur
in South China, four of which are endemic. TyZoplecta
became a characteristic genus for the Cathaysian Province
during this time. The first diversification of Tyloplecta not
only increased the species richness of the genus in South
China, but also resulted in the first phase of dispersal of
this genus to areas outside South China particularly into
the shallow seas flanking the eastern margins of the Sino-
Korea and Bureya blocks in East Asia (Japan and Far East
Russia) and Peri-Gondwanaland (central Tibet, central
Iran, Salt Range of Pakistan) (Fig. 4B). The southward
migration appears to have been assisted or enhanced with
a number of migratory stepping stones scattered in the
southern Palaeotethys, namely the Qiangtang Terrane,
Indo-China block, and the Shan-Thai Terrane (Fig. 4C).
Most of these ‘stepping stones’ constitute what is known
as the southern transitional zone between the Cathaysian
Province of t h e Palaeoequatorial Realm a n d the
Gondwanan Realm (Shi et al., 1995) (Fig. 4B). The Middle
Permian invasion of Tyloplecta into the South Kitakami
Belt of northeast Japan and South Primorye of Far East
Russia is also of note as it appears to coincide with the
onset of a mixed Cathaysian/Boreal shallow marine biota
in East Asia (Tazawa, 1991; Shi et al. 1995).
The Wuchiapingian saw further diversification and
geographical expansion of Tyloplecta. In addition to the
areas already invaded and occupied by Tyloplecta during
the Guadalupian, Tyloplecta penetrated further west into
Gondwana Research, 1/: 6, No.4, 2003
785
western Palaeotethys (North Iran, southern Armenia,
southwestern Russia (Caucasus Mountains), Hungary,
Slovenia, and Yugoslavia) (Fig. 4C). In the northeast of
the Palaeotethys, Tyloplecta maintained its presence in the
Bureya block of Far East Russia, but with much reduced
species richness.
Tyloplecta suffered a major extinction at the end of
Wuchiapingian. As a result, the geographical range and
diversity of Tyloplecta dramatically reduced during the
Changhsingian, with no new species and only one remnant
species from the Guadalupian, T.yangtzeensis, surviving
in both the Cathaysian and Western Tethyan provinces
(Fig. 4D).
Palaeogeographical and Plate Tectonic
Implications
The timing and pathways of migration of Tyloplecta
through the Permian and out of South China, as outlined
above and depicted in figure 4, is important in
understanding Permian palaeogeography and dynamics
of terrane movement across the Palaeotethys.
In the Sino-Mongolian-Japanese Province (Shi and
Tazawa, 2001)) a transitional biogeographical unit
characterised by an admixture of both Cathaysian and
Boreal taxa (Tazawa, 1991; Shi et al., 1995; Shi and
Tazawa, 2001), Tyloplecta species are found along with
typical Boreal and bitemperate brachiopod genera:
Waagenites, Anidanthus, Megousia, Paucispinifera,
Yakovlevia, Waagenoconcha, Neospirifer, and Spiriferella,
as well as characteristic South Chinese (or Cathaysian)
elements such as Kiangsiella, Haydenella, Spinomargini,fera,
Leptodus, Transennatia, and Compressoproductus. This
mixed brachiopod fauna is associated with fusulinaceans
of the Monodiexodina sutschanica (or M. matsubaishi) and
Parafusulina stricta Zones (Kotlyar et al., 1999). The origin
of this mixed Cathaysian/Boreal brachiopod fauna in East
Asia has been subject to much recent study, and a general
consensus is emerging (Tazawa, 1991; Shi et al., 1995;
Tazawa, 1998, 2000; Shi and Tazawa, 2001). In analogy
to the geographical distribution of Recent mixed polar/
subpolar and warm-water shallow marine faunas (eg., Sea
of Japan and the Mediterranean Sea), the mixed
Guadalupian (mainly Wordian) Cathaysian/Boreal
brachiopod fauna would suggest a mesothermal or
temperate climatic setting for the Sino-Mongolian-
Japanese Province during this time. This in turn would
imply that this province was likely located at a middle
latitudinal zone ( 2 5 O -4OO latitude) in the Northern
Hemisphere and was influenced by both warm water
(from the Panthalassa and Eastern Palaeotethys) and cold
water (from the Arctic) currents during the Middle
Permian (Fig. 4B).
786 G.R. SHI AND Z.Q. CHEN

Fig. 4. Historical patterns of palaeogeographical distribution

and migration of Typloplectu Muir-Wood and Cooper
through the Permian. (Base map is after Ziegler et
al. (1997). The subdivisions of biogeographical units
are from Shi et al. (1995) and Shi and Grunt (2000)).
Explanation of tectonic blocks o r terranes
abbreviated: Br - Bureya block, Ic - Indo-China block,
Ir - Iranian block, Qt - Qiangtang Terrane, Sc - South
China block, Sk - Sino-Korea block, St. - Shan-Thai
Terrane.

Gondwana Research, V. 6, No. 4, 2003

 PERMIAN TYLOPLECTA MUIR-WOOD AND COOPER, 1960 (BRACHIOPODA)
The distribution and dispersion history of Tyloplecta in
the southern Palaeotethys may also be interpreted in the
broad context of the contemporaneous plate tectonic
configurations and associated palaeogeographical changes
of the Palaeotethys. The first appearance of the genus in
the region was in the Roadian (Fig. 4B). By early late
Lopingian (Wuchiapingian), the genus had spread further
to the west, to the Iranian block of north Iran and Armenia
(Fig. 4C). This westerly progression in the dispersion of
Tyloplecta suggests a possible westerly migration route for
the genus via such stepping stones as the Qiangtang and
Shan-Thai terranes during the early Middle Permian. At
this point, i t is also w o r t h y noting t h a t t h e
palaeogeographical distribution pattern of Tyloplecta, as
outlined above, is similar to that of Cryptospirifer,an index
Wordian brachiopod genus for the Cathaysian Province
(Shi and Shen, ZOOO), and also similar to the distribution
of the Capitanian-Wuchiapingian Shanita-Hemigordius
(Hernigordiopsis)foraminiferid fauna (Nestell and Pronina,
1997). These joint distribution patterns of both brachiopod
genera and foraminiferans among the cluster of
microcontinents o r terranes in t h e southern and
southwestern Palaeotethys are therefore likely to indicate
a much closer geographical proximity between these
terranes as well as between them and South China and
Indo-China blocks during the late Middle Permian
(Wordian-Capitanian) a n d early Late Permian
(Wuchiapingian) . This conclusion supports the view of
Shi et al. (1995) and Shen and Shi (2000) that the
biogeographical evolution of northern Iran through the
Permian is closely comparable with that of the general
‘Cimmerian continental strip’ (Sengor, 1979). This would
suggest that tectonically northern Iran may have behaved
in a similar fashion to t h a t of t h e Cimmerian
microcontinents through the Permian. However, there is
a notable difference in Permian biogeography between
north Iran and the rest of the Cimmerian microcontinents,
namely that the Iranian block was incorporated into the
Western Tethyan Province in the Changhsingian while the
rest of the Cimmerian microcontinents remained within
the Cathaysian Province (Shen et al., 2000). Shen and
Shi (2000) explained this difference as a result of the
Iranian block’s unique position within the NW-SE
orientated Cimmerian microcontinental strip- being on
the northwestern end of the continental strip- and its
continuous northward drifting through the Permian.
Conclusions
Globally there are 20 species (including species
remaining in open nomenclature) currently known from
Tyloplecta,ranging from Kungurian to Changhsingian. The
genus went through its first diversification in the Wordian,
Gondzuana Research, V: 6, No.4, 2003
78 7
suffered a major extinction in the end-Guadalupian, and
re-diversified in the Wuchiapingian. T. yangtzeensis is the
only species that persisted into the Changhsingian and
the Permian-Triassic mass extinction. A detailed global
review of the spatio-temporal distributions of this genus
suggests that South China was the centre of origin for the
genus and that the genus began dispersal to other regions
in the Wordian. Further analysis of the distribution
patterns of the genus in Eastern Europe, South and South
East Asia indicates that the genus dispersed from Eastern
Palaeotethys (South China) to Western Palaeotethys (e.g.,
North Iran, Armenia, and Eastern Europe) in a staged
fashion and via some ‘stepping stones’ (i.e., the Cimmerian
microcontinents). The palaeogeographical spread and
associated faunas of Tyloplecta suggests that the genus is
generally indicative of warm-water palaeoequatorial
conditions, but could also exist in temperate zones of
mesothermal conditions in the form of mixed cold- and
warm-water faunas.
Acknowledgments
This study is supported by a research grant from the
Australian Research Council (to GRS). We are grateful for
the critical reviews by Ian Metcalfe and Jun-ichi Tazawa
which have helped improve this paper.
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