Palaeobiogeography of Kazanian-Midian (Late Permian) western Pacific Brachiopod faunas

Journal ofSourheast

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Asian Earth Sciences, Vol. 12. No. 112. pp. 129-141, 1995 Copyright 0 1995 Elsevier Science Ltd Printed in Great Britain. All rights reserved 074%9547/95$9.50 + 0.00

Palaeobiogeography of Kazanian-Midian (Late Permian) Western Pacific Brachiopod Faunas G. R. Shi and N. W. Archbold School of Aquatic Science and Natural Resources Management, Deakin University, Rusden Campus, 662 Blackburn Road, Clayton, Victoria, Australia (Received 18 November 1994; accepted for publication 21 February 1995)

data matrix of the presence/absence occurrence data of 129 genera from 16 fauna1 stations (selected from 19 initially compiled fauna1 stations on the basis of sampling and study adequacies) is analysed by cluster analysis, nonmetric multidimensional scaling, principal coordinate analysis and minimum spanning tree. Three core fauna1 groups are revealed and interpreted as representing three biotic provinces. They are the Verkolyma Province embracing the Kolyma Massif, Verchoyan Mountains, east Zabaikal and northern Mongolia; the Cathaysian Province of Japan, northern and southern China and the Indo-China block; and the Austrazean Province of eastern Australia and New Zealand. The overall evolving pattern of Permian marine provincialism of the western Pacific is summarized from our previous, as well as current studies, on the basis of three Abstract-A

discrete time slices: Asselian-Tastubian, Baigendzhinian-Early Kungurian and Kazanian-Midian. An attemnt is also made to exnlain the marked change of marine provinciality during the Permian and climatic in the western Pacific in the’ context of plate tectonics, transgression/regression amelioration.

Introduction This paper continues our work on the western Pacific Permian marine stage-by-stage palaeobiogeography. The western Pacific region as used herein includes northeast Siberia, Mongolia, the Korean Peninsula, Japan, China, the Indian subcontinent, the Himalaya, Southeast Asia, Australia and New Zealand (Fig. 1). Our previous quantitative studies @hi and Archbold, 1993, 1995b) have analysed the brachiopod data of Asselian-Tastubian and Baigendzhinian-Early Kungurian time intervals, respectively, and showed that the Permian marine provinciality of the region changed markedly during the Early through to mid-Permian (Asselian to Kungurian). However, further investigations into the Late Permian are deemed necessary in order to reveal the dynamic nature of the western Pacific marine provincialism. For this purpose, a data base consisting of Kazanian to Midian brachiopod faunas from the western Pacific region was compiled and analysed by a set of multivariate statistical methods, and the computed results are interpreted herein in conjunction with our earlier studies.

Data and Methods The procedure for collecting and treating raw data follows that outlined by Shi and Archbold (1993). A raw data set of 19 fauna1 stations, or operational geographical units (OGUs), was initially compiled from all published literature (Table 1). An OGU is a geological entity, such as an intracratonic basin, a marginal (“geosyncline’‘-type) basin, or an epicontinental seaway. OGUs are as extensive as possible to embrace all correlative fossil communities from various substrates and water depths, provided that geological evidence is clear

that only one basin is involved. Therefore, an OGU may represent a fauna from one locality or a composite (synthetic) record of several assemblages from the same basin. This sampling method is designed to minimize the potential effect of local ecology on large-scale palaeobiogeographic analysis. For each OGU, a list of species, genera and families is provided and revised from the original literature wherever necessary and possible in terms of modern taxonomy. The taxonomic data base (Shi and Archbold, 1994) providing details of the fauna1 lists and revisions is available either in hard copy or disk format from the authors at a small charge to cover the cost. Three taxonomic levels were reviewed in our data base mainly for the sake of internal taxonomic consistency, but only generic data were used in quantitative analyses, following Shi and Archbold (1993, 1995b). All but three OGUs (Yanji of Jilin Province, China, Hainan Island of China and Shikoku of Japan) were included in cluster analysis, ordination and minimum spanning tree. The three “outlying” OGUs were excluded because they do not satisfy one or more of the subjective criteria suggested by Shi (1993b) and elaborated by Shi and Archbold (1993) in selecting raw data to be analysed quantitatively. These OGUs were however subjected to discriminant analysis and reallocated into the core groups defined by the cluster and ordination analyses (see below). Having removed these outlying OGUs from the raw data, a secondary data matrix was derived, consisting of the presence/absence (i.e. l/O) data of 129 genera, or operating taxonomic units (OTUs), from 16 OGUs (see Appendix 1 for a list of these genera). This data set was then analysed in both Q and R modes by cluster analysis (CA), nonmetric multidimensional scaling (NMDS) and principal coordinate analysis (PCO). Q-mode analysis aims to reveal relationships between OGUs (samples),

129

130

G. R. SHI and N. W. ARCHBOLD

h. .

0

Fig. 1. Major pre-Mesozoic tectonostratigraphical units in the western Pacific region. Br-Brook Street terrane, New Zealand; Ch-Chukotka Block; In-Indo-China Block; Iz-Inner Zone, Japan; Ju-Juggr Basin; Ka-Karamea terrane, New Zealand; Ki-Kitakami Massif; Ko-Kolyma Block; Ls-Lhasa terrane; NC-North China Block; Oz-Quter Zone, Japan; Qd-Qaidam Basin; Qt-Qiangtang terrane; Si-Siberia; Sl-Songliao Massif; St-Shan-Thai terrane; Ta-Tarim Basin; To-Torlesse terrane, New Zealand; Yz-Yangtze Block; Tectonic boundaries. (l)--Verchoyan Fold Belt; (2)---Koryak Fold Belt; (3)-Hingan Fold Belt; (4~Sikhote Alin Fold Belt; (5)--Yinshan-Tumen Fold Belt; (6)--Altay Fold Belt; (7bTianshan Fold Belt; (8)-Kunlun Fold Belt; (9)-Banggong Co-Nujiang Suture; (IO&Himalayan terranes/sutures (undifferentiated); (1 I)--Sanjiang Fold Belt; (12~Uttaradit-Nan Suture; (13)-Qinling Fold Belt; (14)-Tasman Fold Belt.

while R-mode is intended to unravel inter-OTU (i.e. inter-variable) relationships. These multivariate methods are different in underlying assumptions about data structure and in algorithms [see Shi (1993b) for a review and references therein provided], but are supplementary in the sense that they provide a cross-check for congruency and rigour of the patterns derived individually. Many authors have adopted this integral approach to both palaeoecology (e.g. Bjerstedt, 1988) and palaeobiogeography (e.g. Smith, 1988; Shi and Waterhouse, 1991; Shi and Archbold, 1993, 1995b). The multivariate analyses were carried out using PATN software (Belbin, 1992). UPGMA (unweighted pair-group arithmetic averaging) CA was first used to classify the OGUs into discrete core groups (Fig. 2). The

degree of robustness of the core groups recognized was tested in three different ways. Firstly, the cophenetic correlation coefficient of the cluster analysis, which in this case is valued exceptionally high at 0.9604, is a strong indication that there was little distortion to the underlying grouping structure, if any, of the data set caused by the clustering procedure employed. Secondly, NMDS and PC0 were applied to the same data set for investigating and graphically displaying the interrelationships of the OGUs in the form of a scatter plot (Figs 3,4). However, if the original data set does possess any grouping structure, these potential groups would appear in both NMDS and PC0 scatter plots and they should, in theory, correspond to those recognized by CA.

Mean latitude, longitude

45”3O’S, 168”30’E 42”54’S 147”18’E 25”17’S 150”18’E 31”07’S 150”58’E 3”4O’N 103”31’E 40”19’N 97”12’E 44”41’N 144”27’E 42”2O’N 117”30’E 40”32’N 95”57’E 33”24’N 113”ll’E 28”12’N 109”12’E 18”12’N 113”ll’E 39”43’N 141”08’E 33”5O’N 132”47’E 43”lO’N 151”2O’N 48”24’N

Stations (OGUs)

Takitimu Mts, New Zealand Tasmania, E. Australia Queensland, E. Australia N.S.W., E. Australia E. Peninsula Malaysia Qilian Mt., China Inner Mongolia Yanji, NE China Beishan, NW China Qinling, China South China Hainan Island Kitakami Mountains Shikoku, Japan Primore E. Russia Northeast

33.9”s (7) ?586”N

22”N (6) ?

14.8”N (4) 5.9”s (;)

19”N (3) ?14.8ON (;)

?

27” If: 5”s (1) 80”s (2) 65”s (2) 70”s

Palaeolatitude (Ref.) # Stratigraphic unit

0.10 0 0.2

20 14 15

0.86

0.16 0

37 4

7

0

0.30

2.42 0.71

0.71 0.28

1.4

0.14 0.1

10 30

2

0.28 0.05

?

3.29

1.85

18

0

0.57

0.08

25

1.14

0.57

0.07

14

1

0.28

0

11

2.4

1.57

1.57

0.29 0.71

0.86

1

0.86 0.29

0.71

1

Permian ratio (c)

0.43

0.86

Sampling efficiency (b)

and Briggs (1986)

(1973)

(1973)

Manakov

(1992) continued wet-leaf

Likharev and Kotlyar (1978)

Yanagida

Hayasaka (1964)

Xu et al. (1992)

Jin and Hu (1978)

Ding et al. (1989)

Ding and Qi (1983)

Li, Gu and Su (1980)

Ding et al. (1985)

Yang et al. (1962)

Nakazawa

Dickins (1968)

Waterhouse

Principal references (1982a)

Clarke (1987)

Waterhouse

data and statistics

0.86

0.15

26

0.06

0.18

11

17

0.06

Endemicity (a)

faunas-basic

16

Total genera

Western Pacific brachiopod

Brook St. Mangarewa terrane formation Tasmania Malbina Fm. basin Abels Bay Bowen Flat top basin Fm. Sydney Muree and basin Mulbring Fms “Jenka IndoChina Pass” North Ciaodigou China Fm TMHFB Yihewusu Fm. North Miaoling China Fm. TMHFB Jushitan FM. North Wulipo and China Shuxiakou Yangtze Gufeng and block Maokou Hainan Nanlong block Fm. Kitakami Iwaizaki massif Stage Outer Yamamba Limestone zone Sikhote Upper Alin terr. Chandalaz Siberia Yal’din

Tectonostratigraphic unit

Table I. Kazanian-Midian

w

112”ll’E 52”12’N 113”35’E 63”12’N 143”27’E 68”5l’N 124”08’E

Stations (OGUs)

Mongolia East Zabaikal KolymaOmolon River Lena River, Siberia

Tectonostratigraphic unit

block (6) Siberia ?586”N block (6) 59”N Kolyma block (6) 65.2 + 2”N Verchoyan FB (6)

Palaeolatitude (Ref.)# Horizon Sosuchey complex Omolon and Gijigin H. Omolon and Gijigin H.

Stratigraphic unit

0.14

21 0.43

0.57

0.24

25

Sampling efficiency (b) 0.28

Endemicity (a) 0

7

Total genera

Table 1. continued from page 131

1

1

0.29

Permian ratio (c)

Abramov

and Grigoryeva

(1988)

Stepanov (197 1)

and Popeko (1974)

Zavodovskiy

Kotlyar

Principal references

families found

Although the index may be viewed as a modified diversity statistic, crudely normalized for adequacy of sampling, our study (Shi and Archbold, 1994) shows that it is biased in favour of poorly sampled stations by giving them higher values. To minimize this effect, we propose an alternative version of the index by replacing the denominator with “cosmopolitan families expected”. The amended Permian Ratio seems to give more realistic diversity values when taking into account the belt; Fm-Formation; total number of families found and the Sampling Efficiency (see Shi and Archbold, 1994). Explanation of abbreoiations: FB-fold Gp-Group; cng.-conglomerate; lst.-limestone; mst.-mudstone; sle.-shale; slst.-siltstone; sst.-sandstone. TMHFB-Tianshan-MongoI-Hingan Fold Belt.

cosmopolitan

Stehli and Grant (1971) chiefly based on faunas from the Northern Hemisphere, proposed 16 Permian brachiopod families, named the Cosmopolitan Dominants, which they considered can be expected to occur in most Permian marine localities. This list has been revised and reduced to only seven by Waterhouse (1982) to suit all Permian brachiopod faunas. This reduced list is further amended here. The seven cosmopolitan dominant families are: Streptorhynchidae, Rugosochonetidae, Linoproductidae, Marginiferidae, Athyrididae, Spiriferidae and Dielasmatidae. The new sampling index basically provides a measure of the completeness of the sampling of a fauna1 station. (c): Permian Ratio was also proposed by Stehli (1970) to indicate relative palaeolatitudes. This index was defined by families present-cosmopolitan families found

cosmopolitan families found cosmopolitan families expected.

References for palaeolatitua’es are as foflows: (1)-Haston et al. 1989, (2)-Embleton in Veevers 1984, (3)-Pruner 1987, (4)-Ma and Zhang 1986, (5)-Zhao and Coe 1989, (6)-Khramov and Ustritskiy 1990, (7)_McElhinny et al. 1981. (a)-Endemicity refers to the degree of endemism of a fauna1 station, calculated as the percentage of endemic genera of a station over the station’s total genera. (b): Sampling efficiency index (Stehli, 1970; Stehli and Grant, 1971) is a statistic used to evaluate the quality of sampling of a station, calculated as follows:

Mean latitude, longitude

E

.F

0

KAZANIAN-MIDIAN

WESTERN

PACIFIC

BRACHIOPOD

133

FAUNAS

East Malaya (9) Kitakami Massif (10) Inner Mongolia

(11)

Sikhote Alin (12) Qilian Mts. China (13) Qlnltng Mts. Chtna (14) Yangtze Belshan.

Platform (15) China (16)

t

I

I

0.987

0.699

0.411

Fig. 2. Dendrogram of Kazanian-Midian brachiopod stations derived from UPGMA based on Jaccard coefficient. Cophenetic correlation value of this dendrogram is 0.9604. Three core groups (labelled as A, B and C) are clearly delineated at the chosen division line. Generic associations of the groups are also indicated.

In addition to these two testing procedures, minimum spanning trees (MST) (see Gower and Ross, 1969) were also superimposed on the scatter plots of NMDS and PC0 (Figs 3 and 4). A MST may be viewed as a set of line segments, representing the pair-wise associations of the OGUs. The construction of a MST is constrained by (1) that all OGUs have at least one connection, (2) that there are no circuits or loops, and (3) that the tree is minimal length (Belbin, 1992). A MST thus provides a simple and effective means of displaying the relationships between OGUs in the form of a graph. When superimposed on an ordination plot, a MST also serves to indicate both the within-group cohesiveness and the inter-group relationships. R-mode CA, NMDS and PC0 analyses of the 129 OTUs (genera) were also carried out (see Shi and Archbold, 1994, Appendices S-10 for results and graphs). The R-mode CA demonstrates clear delineation of three major generic associations corresponding to the three core groups of OGUs detected by CA. However,

these same generic associations have not been as clearly identified by NMDS or PCO. A final step in the quantitative analysis was to apply discriminant analysis (DA) to the three outlying OGUs (Yanji of Jilin Province, China, Hainan Island of China and Shikoku of Japan) that have been excluded from CA and NMDS/PCO, and relate them via a quantitative procedure to the core groups recognized previously by the latter methods. This re-allocation procedure has been described in detail by Shi (1995; see also Shi and Archbold, 1993). Applying this method, an outlying OGU is allocated to a pre-defined core group with which it has the highest discriminant score. Mathematically, the discriminant score of an OGU is defined by i= 4,

=

ti

a,,s,,. z=I 2

where dk, is the discriminant score of the kth outlying OGU with the jth core group (i = 1. . N), xik the element of binary data of the ith genus at the kth OGU

0.324r

-0.509

-0.384

-0.010

PC0

0.172

AXIS 1

Fig. 3. A two-dimensional ordination plot of fauna1 stations on the first two principal axes of PC0 (accounting for 43% of total variation). Letters (A, B and C) correspond to those in Fig. 2. Sequential numbers (I-16) indicate fauna1 stations as in Fig. 2. The thick black arrow indicates the inferred palaeo-temperature gradient (see text for more discussion).

134

G. R. SHI and N. W. ARCHBOLD

-0.726

-0.078

1.122

0.568

NMDS AXIS 1 Fig. 4. A two-dimensional ordination plot of fauna1 stations on the principal axes 1 and 3 of NMDS (stress value = 0.1053). Letters (A, B and C) correspond to those in Fig. 2. Sequential numbers (1-16) indicate fauna1 stations as in Fig. 2. The thick black arrow indicates the inferred palaeo-temperature gradient (see text for more discussion).

(i.e. either 0 or 1) and a,- the association value of the ith OTU with the jth group, defined as

reliable among the non-probabilistic indices available (Shi, 1993a, b).

binary similarity

Results and Interpretations

,=I

where .x0 is the number of occurrences of the ith OTU in the jth group, Xxi the total number of occurrences of the ith OTU in the entire dendrogram. The association value index, av, provides a measure of the relative contribution or importance of each OTU to each recognized group; it is equal to 0 if an OTU is entirely absent from a group; conversely, it may be scored to 1 if it is confined to a group. In other words, the higher the a+ the more characteristic of the ith OTU for thejth group. As exemplified by Shi and Archbold (1993) with Asselian-Tastubian (Early Permian) Circum Pacific brachiopod faunas, this index appears effective in identifying character taxa and defining generic associations that characterise particular fauna1 groups. Association values of the 129 genera derived from this analysis are given in Appendix 1. As stressed by Shi (1993b), employment of multivariate techniques is for detecting potential grouping patterns of OGUs or OTUs within a raw data set on the basis of quantitative similarity values calculated using a particular type of similarity coefficient and for creating working hypotheses as to how the various groups were formed. Geological and/or biogeographical interpretations of the groups, if derived, must be sought independently and tested by other data and techniques. In this study, the Jaccard Coefficient was used throughout to calculate mutual fauna1 similarities among the stations and genera. This coefficient appears to be the most

The dendrogram derived from UPGMA CA of the 16 OGUs (Fig. 2) reveals three core groups (labelled A, B and C, respectively) at the chosen division line. The same three core groups also appear distinguishable in both PC0 (Fig. 3) and NMDS (Fig. 4) scatter plots. The core groups are interpreted as representing fauna1 provinces, although we are not entirely clear about the compatibility of such defined provinces with modern zoogeographical provinces. Group A is composed of stations from eastern Australia and New Zealand, and founded by the Wyndhamia-Sulciplica-Glendonia association. Other characteristic genera ,that are scored 1 with this group include Echinaiosia, Birchsella, Capillonia, Filiconcha, Magniplicatina, Saetosina, Aperispirifer, Cyrtella, Notospirifer, Maorielasma and Fletcherithyris (see Appendix 1). The

recognition of this group readily conforms to the establishment of the Austrazean Province by Archbold (1983). This province has a relative low generic diversity and a high endemicity (Table 2), suggesting probably much cooler water temperatures than the other provinces and considerable palaeogeographical separation from the other groups in terms of linear distance and/or climatic barriers. Group B is restricted to northern Mongolia, eastern Zabaikal and northeast Siberia and characterised by the Cancrinelloides-Spitzbergenia-Olgerdia association. Its

Table 2. Genetic diversity and endemicity of recognized core fauna1 groups

Total genera (N) Endemic genera (n) Endemicity (n/N)

Group A (Austrazean Province)

Group B (Verkolyma Province)

Group C (Cathaysia Province)

24 17 0.75

35 20 0.57

86 76 0.89

KAZANIAN-MIDIAN

WESTERN

generic diversity is higher than Group A but substantially lower than Group C, and its endemicity is scored moderate at 0.57, being intermediate between the other two groups (see Table 2). The term Verkolyma Province, proposed by Shi and Archbold (1995b) initially for the Baigendzhinian-Early Kungurian time interval, is herein adopted to accommodate this group (more discussion is provided below on the extension of Verkolyma). Group C is large and geographically dispersed, embracing fauna1 stations from southeast Asia (eastern Peninsular Malaysia), China and Japan. Other stations that are also strongly scored to this group through their discriminant scores are Yanji of northeast China, Hainan Island and Shikoku, Japan. This group is identified by the Cathaysia-Spinomarginifera-Tyloplecta association incorporating numerous genera that are either unique to or most characteristic for this group: Meekella, Enteletes. Pygmochonetes, Tenuichonetes, Catha_vsia, Haydenella, Spinomarginifera, Tvloplecta, Compressoproductus, and Richthofenia. The generic di-

versity of the group is the highest compared to the other groups, coupled with also the highest endemicity value of 0.89 (Table 2) suggesting a lower, presumably palaeotropical, palaeolatitudinal setting and considerable palaeogeographical distances or/and palaeoclimatic barriers from the other two groups. Fang (1985) has recognized the distinctiveness of the group and other Permian marine invertebrate faunas from south China, Japan and Indo-China and advocated the term Cathaysian Province for representing these faunas. As revealed by CA (Fig. 2) and confirmed by the MST (Figs 3 and 4), Group A is closer to Group B with respect to Group C. This linkage is facilitated by a number of genera unique to Groups A and B. These taxa. commonly known as the antitropical or bipolar elements, include Arctitreta, Terrakea, Fusispirifer, Tomiopsis, and Marinurnula. It is also interesting to note that in both PC0 (Fig. 3) and NMDS (Fig. 4) Groups A and B are located on one side of the scatter plots, being separated from Group C. As interpreted above, Groups A and B may represent higher palaeolatitudinal faunas and Group C a palaeotropical fauna, the spatial positioning of the groups in the scatter plots thus appears to indicate a palaeolatitude-related temperature gradient, changing from cool or temperate to warmer palaeo-conditions in the inferred directions (Figs 3 and 4).

Evolution of Permian Marine Provincialism the Western Pacific

in

Although more work is required to complete the stage by stage analysis of Permian western Pacific marine palaeobiogeography, an evolving pattern of marine provincialism through the Permian is already discernible from our current as well as previous studies. To summarize this pattern, we may examine three time slices: early Early Permian (Asselian-Aktastinian), late Early Permian (Baigendzhinian-Kungurian) and Late Permian (Kazanian-Dorashamian) (see Shi and Archbold, 1993, 1995b; and this paper), which demonstrate a dynamic pattern of Permian marine provinciality in the western Pacific. In the Asselian-Tastubian, four biotic provinces are distinguishable in the western Pacific: Indoralian, HiSEAES 17-1 2--J

PACIFIC

BRACHIOPOD

FAUNAS

135

malayan, Cathaysian, and Verkolyma (Fig. 5A). The name Verkolyma Province is herein preferred to the Boreal Province as we originally used (Shi and Archbold, 1993) because the latter seems less precise in definition and is better preserved as a realm rather than a provincial name. Shi and Archbold (1995b) realized this nomenclatural problem and proposed Verkolyma, initially only for the Baigendzhinian-Early Kungurian time slice and restricted it to the Verchoyan Mountains and the Kolyma massif. The usage of this term is now extended to also include east- Zabaikal and northern Mongolia, and temporally also stretched to cover virtually the entire Permian. The province seems to have a persistent identity throughout the Permian and is readily distinguishable from neighbouring southeastern Mongolia-Inner Mongolia and the Russian Far East (see also Durante et al., 1985). Two features are distinct for the Asselian-Tastubian palaeobiogeographical pattern. Firstly, the recognition of the Indoralian Province embracing Peninsular India, Australia and the Shan-Thai terrane signifies the existence of a relatively uniform cold climate over these (micro)continents and their flanking shallow seas, a cold regime apparently associated with the onset of extensive contemporaneous Gondwanan glaciation. The ShanThai terrane is included because it contains a distinct cool-water Gondwana-type marine fauna of Asselian-Tastubian age (Waterhouse, 1982b). A similar coolwater fauna of contemporaneous age or slightly younger has also been recorded from the Baoshan block, western Yunnan (Fang, 1994; Shi et al., in preparation). The second significant feature is that Inner-Mongolia-northeast China and southeast Mongolia are incorporated into the Cathaysian Province during this time, whereas the same regions, together with northeast Japan and the Sikhote Alin terrane of the Russian Far East, formed an independent fauna1 province in the Baigendzhinian-Early Kungurian (see Fig. 5B) and, again, fell into the Cathaysian Province in the KazanianMidian (Fig. SC). For the late Early Permian interval, fundamental restructuring of the early Early Permian western Pacific marine palaeobiogeography appears to have taken place (Fig. 5B). This restructuring process included-( 1) that the Indoralian Province had split into two provinces: Austrazean and Westralian (Archbold, 1983); (2) that two distinct transitional provinces were developed: the Sino-Mongolian Province (Shi and Archbold, 1995b) in the north and the Cimmerian Province (Archbold, 1983) in the south; and (3) that the areal extents of the Cathaysian Province and the Gondwanan Realm appear to have shrunken relatively due to the development of the two transitional provinces. Included in the SinoMongolian Province are the TianshanMongol-Hingan Fold Belt (i.e. southeast Mongolia, Beishan of northwest China, Inner Mongolia, Heilonjiang and Jilin Provinces), the Sikhote Alin Terrane of the Russian Far East, west¢ral and northeast Japan. This roughly east-west striking belt, named by Tazawa (1991) the Inner Mongolian-Japanese transition zone. is distinguished by an admixture of marine invertebrate genera indicative of both the Cathaysian and Verkolyman affinities. Among the Verkolyman representatives are Uraloproductus. Sowerhina, Yakotlleria. .lictinoconchus. Timaniella. and Tumarinia; and among the Cathaysian affinities are included Meekella, Entekefcs. Edriosteges,

136

G. R. SHI and N. W. ARCHBOLD

Verkolymr Pr. Sino-Mongolia 1%. Csthaysian 1%. CimmCriiul

Pr.

Westralian I’r. Austra7fi!an1%.

m

Verkolyma Pr.

Austrazean

Pr.

Fig. 5. Schematic reconstructions of Permian palaeogeography and palaeobiogeography (with base map modified from Scotese and McKerrow, 1990). A-Asselian-Tastubian(early Early Permian) (slightly modified from Shi and Archbold, 1993); B-Baigendzhinian-Early Kungurian (late Early Permian); C-Kazanian-Midian (Late Permian). I-Gondwana Realm, II-Tethys Realm, III-Boreal Realm. Explanation of major tectonic blocks: A-Afghanistan; Bh-Birds Head terrane, Irian Jaya; I-Iran; Id-India; Im-Sa-Inner Mongolia-SikhoteAlin terrane; In-IndoChinaBlock; Jp-Japan; K-Kolyma Block; Kz-Kazkhstan; L-Lhasa terrane; M-Central Mongolian Massif; Q-Qiangtang terrane; S-Songliao Massif; SkSine-Korea Block; Sm-Sumatra; St-Shan-Thai terrane; T-Tarim Basin; V-Verchoyan Mountains; Yz-Yangtze Block.

KAZANIAN-MIDIAN Tyloplecta, Spinomarginifera,

WESTERN

and Compressoproductus.

In addition to these two major components, the SinoMongolian Province is further characterized by endemic genera such as Paramargintjera and genera with antitropical distributions, notably the fusulinid Monodiexo dina, the rugose coral Lytvolasma, and numerous brachiopod genera: Strophalosia, Wimanoconcha, Transennatia, Magniplicatina, Camerisma, Attenuatella, Spiriferella, Spirelytha, and Tomiopsis (Shi et al.,

1995). Likewise, the Cimmerian Province is also typified by an admixture of genera representing both the Cathaysian (e.g. Tenuichonetes, Edriosteges, Neoplicatifera, Tyloplecta, Uncinunellina) and Gondwanan affinities (e.g. Retimargint~era, Stereochia, Costiferina, Globiella), in addition to endemic (e.g. Jipuproductus and Calliomarginatia) and genera that are also characteristic for the Sino-Mongolian Province (e.g. Chonetinella, Cyrtella, Spiriferella, Lytvolasma, Monodiexodina). At this point, it is appropriate to note that a narrower, more specific spatio-temporal definition of the Cimmerian Province is adopted herein, that is, to exclude Peninsular India. Under this revised definition, this province stretches from the middle east through Afghanistan and the Himalaya east to southeastwards to the Shan-Thai terrane, Timor and Irian Jaya. Recognition of subprovinces is possible given the complexity of the faunas embraced and the great geographical extent of the province (see Archbold and Thomas, 1986; Archbold, 1987; Shi and Archbold, 1995b). The palaeobiogeographical picture of the Kazanian-Midian (Late Permian) interval is again different. The two prominent late Early Permian transitional provinces are no longer recognizable as such, but are incorporated into the Cathaysian Province, which, as a result, broadened its geographical extent (Fig. 5C). Clearly, the patterns shown by Fig. 5 point to a highly dynamic nature of the Permian palaeobiogeography of the western Pacific. Centre to this dynamic pattern is the formation and diminution of the Sino-Mongolian and Cimmerian Provinces. It has been suggested (Shi et al., 1995) that the formation of the Sino-Mongolian Province was probably due to the transgression of the Arctic sea during the Baigendzhinian to the Kungurian and the accompanied fauna1 invasion of the Verkolyman elements into the Sino-Mongolian seaway. On the other hand, the origin of the Cimmerian Province is more speculative. Shi and Archbold (1995a) suggested two scenarios: a plate tectonic displacement interpretation and an alternative approach involving climatic amelioration and fauna1 migration across the southern Tethys. A more preferable hypothesis has also been put forward (Shi et al., 1995), which emphasizes the effect of the interplay of tectonic movement and climatic change on marine provincialism. This hypothesis explains that the Cimmerian blocks were probably located in the northern periphery of Gondwana during the Asselian to Tastubian and hence developed a cold to cool-water fauna reminiscent of those of Gondwana. This cold-water phase corresponded to, and hence can be presumed to be a consequence of, the extensive early Early Permian Gondwanan glaciation. A rifting event may have taken place after the Tastubian in response to the opening of the Neo-Tethys (cf. Gaetani and Garzanti, 1991). As a consequence, the climatic amelioration may have been further enhanced by the global

PACIFIC

BRACHIOPOD

FAUNAS

137

warming event that followed the retreat of the Gondwanan glaciation (Dickins, 1985a, b). During the rifting and subsequent drifting process, part of the Cimmerian blocks might have been split off from the others and drifted more rapidly northwards. An example of this rapid-drifting fragment of the Cimmerian blocks may be the Shan-Thai terrane which exhibited a Late Sakmarian mixed type fauna, while other parts of the Cimmerian blocks were still predominantly influenced by cold to cool temperate conditions during the Sakmarian to Early Artinskian interval (Shi and Archbold, 1995a; Shi et al., 1995). By the Late Artinskian, all Cimmerian blocks probably had moved into a lower latitudinal zone, presumably in a position intermediate between the Cathaysian blocks (Yangtze, Indo-China, and Tarim) and Gondwana. It is due to this intermediate position with geographical proximity to both Cathaysian blocks to the north and Gondwana in the south that the Cimmerian blocks developed a transitional/mixed type fauna characterized by both Tethyan and Gondwanan elements. By the Kazanian/Midian, owing to their continuing northwards drift and further climatic amelioration, the Cimmerian blocks probably had moved into the palaeotropical zone and become integrated with the Asian Tethyan biogeographical region (Fig. SC). Palaeomagnetic data are critical in validating this hypothesis. Unfortunately, such data are still limited for most of the Cimmerian blocks, and those that have been published lack precise stratigraphical control and usually do not provide a consistent signature, as to where and how far, the various Cimmerian blocks moved during the Permian. For example, the Early Permian palaeomagnetic result of 17.76”N latitude reported by Klootwijk (1979) for the Helmand block of central Afghanistan is in sharp contrast with that (42”s latitude) of Huang and Opdyke (1991) for the western Yunnan portion of the Shan-Thai terrane although this disparity may be accounted for by the possibility that the Cimmerian continental strip was oriented in a north-south or northwest-southeast direction as it appears today, with central Iran and Helmand blocks crossing over the plaeoequator and Shan-Thai terrane located in the southern palaeotemperate zone.

Conclusions Three marine biotic provinces appear to have existed in the western Pacific region during the KazanianMidian and their distribution seems to have been primarily controlled by variations in palaeolatitudes (Fig. 5C). This Late Permian palaeobiogeographical pattern is notably different from those of the early Early (AsselianTastubian) and late Early Permian (BaigendzhinianKungurian) times. In particular, the two late Early Permian transitional provinces, Sino-Mongolian and Cimmerian, have not been recognized for the KazanianMidian, nor do they appear to be recognizable in the Asselian-Tastubian. The waxing and waning of the Sino-Mongolian Province is assumed to have resulted from the transgression and regression of the Arctic sea into the Sino-Mongolian seaway during the late Early Permian and fauna1 invasions from both the Verkolyma and Cathaysian Provinces. On the other hand, the

G. R. SHI and N. W. ARCHBOLD development and diminution of the Cimmerian Province is probably more likely to have been related to a series of tectonic events that swept the Cimmerian blocks away from northern Gondwana in the Early Permian (Sakmarian-Artinskian). As the Cimmerian blocks advanced north, fauna1 mixing with Cathaysian elements took place, and this process may have been reinforced by contemporaneous global warming. Alternative explanations are also available, either emphasizing facies control on fauna1 differentiation (Jin, 1985) or influence of ocean currents and global warming as the main determinant during the Permian biogeographical (Dickins et al., 1993). Acknowledgements-The faunas and quantitative the Australian Research

authors’ work on Late palaeobiogeography Council (NWA grant

Palaeozoic is supported by no. A391321 12).

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Tazawa J. (1991) Middle Permian brachiopod biogeography of Japan and adjacent regions in East Asia. In Pre-Jurussic Geology qf‘inner Mongolia, China (Edited by Ishii, K.. Liu. X.. Ichikawa K. and Huant B.), pp. 213-230. Matsuya Insatsu. Osaka. Veevers J. J. (1984) (Ed.). Phanero-_oic Eurth Hi.s/ory qf Australia. p, 418. Clarendon Press, Oxford. Xu G.-H., Zhang Z.-Z.. Li Z.-H. er a/. (1992) The Permian system. In Geology qf Hainan Island. Vol. I. Slratigraphy and Palaeonrology. pp. 131-160. Geological Publishing House. Beijing. Waterhouse J. B. (1982a) New Zealand Permian brachiopod systematits. zonation. and palaeoecology. N. 2. Grol. Surr. Paleont. Bull. 48, I-158. Waterhouse J. B. (1982b) An Early Permian cool-water fauna from pebbly mudstones in south Thailand. Grol. Msg. 119, 3377342. Waterhouse J. B. and Briggs D. J. C. (1986) Late Palaeozoic Scyphozoa and Brachiopoda (Inarticulata. Strophommida. Productida and Rhynchonellida) from the southeast Bowen Basin. Australia. Palaeontographica Abt. A 193, I-76. Yanagida J. (1973) Late Permian brachiopods from the Yamamba Limestone in the Sakawa Basin, Shikoku, Japan. Science Report Tohoku University, 2nd series (Geology). special vol. 6. 353-369. Yang Z.-Y.. Ding P.-Z., Yin H.-F. and Fan J.-S. (1962) Carboniferous, Permian and Triassic brachiopod faunas of the Qilian Mountains. Geol. Qilian Mountains 4, l-134. Zavodovskiy V. M. and Stepanov D. I. (1971) Brachiopoda. In Polevoi Atlas Permskoi Fuuny i Flory Serero-costoka SSSR (Field Atlas of Permian Fauna and Flora of Northeast USSR) (Edited by Kulikov M. V.), pp. 70--1X2. Severo- Vostochno Ordena Trudovogo Krasnogo Znameni Geologicheskoe Upravleni, Magadan.

Appendix List of Kuzanian-Midian values

brachiopod genera used for quantitative analysis in this stud]) and their association Association (Au&&an Province)

Genera

I. 2. 3. 4. 5. 6. 7. 8. 9.

10. Il. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25.

Acosarina Cooper and Grant (1969) Actinoconchus McCoy (1844) Alatoproductus Ching and Zhu (1978) Altiplecus Stehli (1954) Amurothyris Koczyrkevicz (1976 Anemonaria Cooper and Grant (1972) Anidanthus Whitehouse (1928) Aperispirifer Waterhouse (1968) Araxathyris Grunt (1965) Arctitreta Whitfield (1908) Arctochonetes Ifanova (1968) Attenuatelfa Stehli (1954) Aulosteges Helmersen (1847) Bujtugania Grunt (1976) Birchsellu Clarke (1987) Brach_ythyrina Fredericks (1929) Callispirina Cooper and Muir-Wood (1951) Cancrinella Fredericks (1928) Cuncrinelioides Ustritskiy (1963) Capillomesolobus Pecar ( 1985/ 1986) Capillonia Waterhouse (1973) Cartorhium Cooper and Grant (1976) Cathuysia Ching (1966) Chaoina Fredericks (1933) Chiliunshania Yang and Ting (1962)

values

(Verktlyma Province) 0

1

1

0 I 1 1 0.5

0

0 0 0.5

0

0.66 0 0 0 0 0 0 0.4 0 0

C (Cathaysian Province)

0.6 0 0 0.33 1 1 0 1 0 1 0 0.6 1 0 0 0 0 0 0

0.4 0 1 0 0 0 1 0 0 0 I 0 (1 I (1 I 1 I I continued orerleaf

140

G. R. SHI and N. W. ARCHBOLD

Appendix-continued List of Kazanian-Midian values

brachiopod genera used for quantitative analysis in this study and their association

Association

Genera Cleiothyridina Buckman (1906) Composita Brown (1849) Compressoproductus Sarycheva (1960) Costispinifera Muir-Wood and Cooper (1960) Crassispirijer Archbold and Thomas (1985) 31. Crenispirifer Stehli (1954) 32. Crurithyris George (193 1) 33. Cryptospirifer Grabau (193 1) 34. Cyrtella Fredericks (1924) 35. Denticufophoria Likharev (1956) 36. Derbyia Waagen (I 884) 37. Dictyoclostoidea Wang and Ching (1964) 38. Dielasma King (1859) 39. Echinalosia Waterhouse (1967) 40. Echinauris Muir-Wood and Cooper (1960) 41. Edriosteges Muir-Wood and Cooper (1960) 42. Enteletes Schellwien (1892) 43. Filiconcha Dear ( 1969) 44. Fletcherithyris Campbell (1965) 45. Fusispirifer Waterhouse (1966) 46. Geyerella Schellwien (1900) 47. Gilledia Stehli (1961) 48. Girlasia Gregorio (1930) 49. Glendonia McClung and Armstrong (1978) 50. Haydenella Reed (1944) 51. Hemiptychina Waagen (1882) 52. Hustedia Hall and Clarke (1893) 53. Isogramma Meek and Worthen (1870) 54. Keyserlingina Chernyschev (1902) 55. Komiella Barchatova (1970) 56. Leptodus Kayser (1883) 57. Lethamia Waterhouse (1973) 58. Limbella Stehli (1954) 59. Lingula Bruguiire (1797) 60. Linoproductus Chao (1927) 61. Liosotella Cooper (1953) 62. Loczyella Frech (190 1) 63. Magadania Ganelin (1977) 64. Magniplicatina Waterhouse (1983) 65. Maorielasma Waterhouse (1964) 66. Marginalosia Waterhouse (1978) 67. Marginifera Waagen (1884) 68. Marinurnula Waterhouse (1964) 69. Martinia McCoy (1844) 70. Meekella White and St. John (1867) 71. Megousia Muir-Wood and Cooper (1960) 72. Mongolosia Manakov and Pavlova (1976) 73. Monticulifera Muir-Wood and Cooper (1960) 74. Neochonetes Muir-Wood (1962) 75. Neoplicatifera Ching, Liao and Fang (1974) 76. Neospirifer Fredericks (1924) 77. Notospirifer Harrington (1955) 78. Notothyris Waagen (1882) 79. Olgerdia Grigor’yeva (1977) 80. “Omolonia” Ganelin (1990) [non Omolonia Alekseeva (1967)] 81. Orbiculoidea d’orbigny (1947) 82. Orthotetina Schellwien (1900) 83. Orthotichia Hall and Clarke (1829) 84. Paramarginifera Fredericks ( 19 16) 85. Paraplicatifera Zhao and Tan (1984) 86. Penzhinaella Solomina (1985)

26. 27. 28. 29. 30.

(Austtazean Province) 0.14 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 I 1 0.66 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1

1 0 0.66 0 0 0 0 0 0 0.33 0 0 1 0 0 0 0 0 0 0 0 0

values

(Verktlyma Province) 0.3 0 0 0

1 0 0 0 0 0 0 0 0.2 0 0 0 0 0 0 0.33 0 0 0 0 0 0 0 0 0

1 0 0 0 0 0.25 0 0 1 0 0 1 0.33 0 0 0

1 1 0 0 0 0.66 0 0

1 1 0.5 0 0 0 0

1

C (Cathaysian Province) 0.56

1 1 1 0

1 1 1 0 1 1 1 0.8 0

1 1 1 0 0 0 1 0

1 0 1 1 1 1 1 0 1 0 1 1 0.75

1 1 0 0 0 0 0 1 1 1 0 0 1 0.66 1 0.33 0

1 0 0 0.5

1 1 1 1 0 continued

KAZANIAN-MIDIAN

WESTERN

PACIFIC

BRACHIOPOD

141

FAUNAS

Appendix-continued List of Kazanian-Midian values

brachiopod genera used for quantitative analysis in this study and their association

Association values B C (Austtazean (Verkolyma (Cathaysian Province) Province) Province)

Genera 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129.

Permicola Koczyrkevicz (1976) Phricodothyris George (1932) Plekonella Campbell (1953) Prorichthofenia King (193 1) Pugnax Hall and Clarke (1893) Punctospirifer North (1920) Pygmochonetes Jin and Hu (1978) Rhombospirifer Duan and Li (1985) Rhynchopora King (1865) Rhynoleichus Abramov and Grigoryeva (1983) Richthofenia Kayser (188 1) Rostranteris Gemmellaro (1899) Saetosina Waterhouse (1986) Scacchinella Gammellaro ( 189 1) Schellwienella Thomas ( 19 10) Schuchertellu Girty (1904) Semibrachythyrina Liao (1979) Spinomarginifera Huang (1932) Spiriferella Chernyshev (1902) Spiriferellina Fredericks (1924) Spiriferina d’orbigny (1847) Spirigerella Waagen (1883) Spitzbergenia Kotlyar (1977) Squamularia Gemmellaro (1899) Stenoscisma Hall (1847) Streptorhynchus King (1850) Strophalosia King (1850) Strophalosiina Likharev (1935) Sulciplica Waterhouse (1968) Tenuichonetes Jin and Hu (1978) Terrakea Booker (193 1) Tomiopsis Benediktova (1956) Transennatia Waterhouse (1976) Tumariniu Solomina and Grigoryeva (1973) Tyloplecta Muir-Wood and Cooper (1960) Uncinunellina Grabau (1932) Uncisteges Jin and Hu (1978) Urushtenoidea Jin and Hu (1978) Vediproductus Sarycheva (1965) Waagenoconcha Chao (1927) Whitspakia Stehli (1964) Wyndhamia Booker (1930) Zhenaniu Ding (1983)

0 0

0 0 0 0 0 0

0 0

0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0

0.66 0.8 0 0 0 0 0 0 0 0 0 0

0.5 1 0 0 0 0 0 0 0 0 0.25 0 0 0 1 0 0 0 1 1 0 0 0.33 0.2 0 1 0 0 0 0 0 0 0 0 0

1 1 0 1 1 1 1 1 0.5 0 1 1 0 1 1 1 1 I 0.75 I 1 1 0 1 1 1 0 0 0 1 0 0 1 0 1 1 1 1 1 1 1 0 1