Global paleobiogeography of brachiopods during the ...

Gondwana Research 26 (2014) 1173–1185

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Global paleobiogeography of brachiopods during the Mississippian—Response to the global tectonic reconfiguration, ocean circulation, and climate changes Li Qiao a,b,⁎, Shu-zhong Shen a a b

State Key Laboratory of Paleobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Science, 39 East Beijing Road, Nanjing 210008, China Geological Survey of Jiangsu Province, 700 Zhujiang Road, Nanjing 210018, China

a r t i c l e

i n f o

Article history: Received 5 May 2013 Received in revised form 5 September 2013 Accepted 8 September 2013 Available online 12 October 2013 Handling Editor: J.G. Meert Keywords: Brachiopods Paleobiogeography Mississippian Rheic Ocean Latitudinal diversity gradient

a b s t r a c t Changes in Mississippian global paleogeography derived from the reconfiguration of the continents, a reversal in ocean currents and global cooling. Although the tectonic and climatic changes are well-documented, their effects on the distribution of brachiopod fauna are poorly documented. Here we present systematic quantitative analyses on global paleobiogeography based on a global brachiopod database from the Mississippian (i.e., Tournaisian, Visean, and Serpukhovian). The dataset consists of 2123 species of 344 brachiopod genera from 1156 localities. Our results reveal that global provincialism was not evident during the Tournaisian and Visean Stages. Two realms, i.e., the Gondwanan and Paleoequatorial Realms, are recognized during the Tournaisian. The Paleoequatorial Realm dominates during the Visean Stage, whereas the Gondwanan Realm is not documented due to the absence of data points. In contrast to the early and middle Mississippian stages, faunal provincialism is greatly enhanced in the Serpukhovian Stage with Paleotethyan and North American realms easily distinguished. This indicates that the Rheic Ocean was closed before the Serpukhovian due to the collision between Gondwana and Laurussia, that disrupted faunal interchange between the Paleotethys and North America. In addition, the paleolatitude-related thermal gradient was enhanced and the Boreal Realm was distinguished from the Paleotethyan Realm during the onset of the Late Palaeozoic Ice Age (LPIA) in the Serpukhovian. The paleolatitude diversity gradient pattern further shows a distinct shift of diversity center from the southern tropic zone in the Tournaisian and Visean to the northern tropic zone in the Serpukhovian. © 2013 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

1. Introduction The Mississippian was one of the most critical transitions during the earth history. First of all, two major continents, Gondwana in the south and Laurussia in the north, approached each other. This resulted in the closure of the Rheic Ocean, which had been an extremely important seaway for biotic interchange between Gondwana and Laurussia during the Late Palaeozoic (Mckerrow et al., 2000; Scotese, 2001; Franke, 2006; Nance, 2010; Meert, 2012). It is natural to suppose that this kind of major geographic changes will dramatically alter the global oceanic currents and faunal distributions. However, previous studies on this topic were mostly from a tectonic perspective (e.g., Bozkurt et al., 2008; Sintubin et al., 2008; Nance, 2010; Nance et al., 2010; Romer and Hahne, 2010; Jastrzębski et al., 2013; Klootwijk, 2013), and relatively little has been done from a perspective of faunal distribution. Second, the Mississippian also witnessed notable climate changes between greenhouse and icehouse climates (Veevers, 2004; Shi and Waterhouse, 2010), that led to substantial global sea-level changes (Stanley and Powell, 2003; Bambach et al., 2004; Powell, 2007; ⁎ Corresponding author. E-mail address: [email protected] (L. Qiao).

Isaacson et al., 2008). Third, the Mississippian is sandwiched between the Hangenberg event at the Devonian–Carboniferous boundary and the Mid-Carboniferous event (Caplan and Bustin, 1999; Kaiser et al., 2006; Shen et al., 2006; Wang et al., 2006; McGhee et al., 2012). Throughout geological history marine faunas have responded to external forcing such as changes in climate and tectonism though evolutionary change. Examples include bryozoans (Ross and Ross, 1985, 1990), foraminifers (Kalvoda, 2002), and corals (Wang et al., 2003; Aretz, 2010, 2011). It has also been noted that the gradually increasing provinciality during the Mississippian was closely related to the global tectonic reconfiguration (Ross and Ross, 1985, 1990; Korn, 1997). A recent study on the Mississippian pelagic ammonoid associations using quantitative methods has demonstrated that provinciality became pronounced in the latest Visean and Serpukhovian due to a major reconfiguration of shelf area (Korn et al., 2012). However, so far no study has been systematically done based on brachiopods. Brachiopods are one of the dominant benthic fossil groups during the Carboniferous and Permian with a very short swimming larvae stage, inhabiting on oceanic shelf from the tidal zone down to ocean floor. Brachiopods are strongly affected by their environment, including sediment composition, water depth, temperature and water salinity (Zezina, 1997, 2008). Brachiopods are mostly sensitive to the

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L. Qiao, S. Shen / Gondwana Research 26 (2014) 1173–1185

latitude-related thermal gradient, but relatively less sensitive to longitude change unless natural barriers like oceans and mountains separate them. Therefore, brachiopods have important implications for paleogeographical reconstruction. Quantitative methods on paleobiogeography based on the Permian brachiopods (Shi, 1993; Shi and Archbold, 1995, 1996; Shen and Shi, 2000, 2004; Shen et al., 2009, 2013; Angiolini, 2013) demonstrated that this fossil group is one of the most useful in order to reconstruct global paleogeography and faunal distributions. Our present work aims to carry out a quantitative study of the Mississippian brachiopods. We first focus on the spatial and temporal paleobiogeographical patterns within the three consecutive Mississippian stages (i.e., Tournaisian, Visean, and Serpukhovian), based on a recentlyestablished global brachiopod database. We further analyze and discuss the major factors controlling the distribution of brachiopods including climate and paleogeography. 2. Data and methods Global brachiopod occurrences in three consecutive stages of the Mississippian (i.e., Tournaisian, Visean and Serpukhovian) all over the world (Table 1) are compiled using the software Filemaker Pro. The dataset consists of 4778 occurrences involving 2123 species of 344 brachiopod genera from 1156 localities (Fig. 1). The items entered into the brachiopod database include the systematic, biostratigraphic, modern coordinates, and paleo-latitude data that were calculated using Track Point 7.0 designed by Christopher Scotese, as well as the lithological, and tectonic data from published literatures for each collection. All the brachiopod species and higher taxonomic assignments were revised and updated in light of recent developments in brachiopod taxonomy. We take recorded taxa that have been systematically attributed to certain genera and species. Records with uncertain taxonomic assignments are excluded, such as overtoniini gen. and sp. indet., orthotetid? indet., productacean indet., Spiriferidina gen. et sp. indet., dielasmatid, choristitinid, and choristitinid. Some species with questionable occurrences are revised, such as Spiriferella sp. recorded in the Visean in Sichuan, South China. Species with qualified names such as aff., cf. etc. are treated as the species without the qualifiers. Age determinations of different brachiopod faunas from literature have been updated as well based on the global stratigraphic and chronostratigraphic timescale (Davydov et al., 2012), whereas the records without stage-level age determination are excluded. We note that the durations of the three stages of the Mississippian are quite different (i.e. the Visean is twice as long as either the Tournaisian or the Serpukhovian). While somewhat of an issue, we note that further sub-division is difficult because many of the fauna do not have biostratigraphic control below the stage level. All localities with brachiopods are assigned to different paleogeographical analysis units or operational geographic units (OGUs) following the principles suggested by Crovello (1981). Decisions related to the choice of OGUs include size and shape of OGUs, each of the OGUs has to be uniform to be delimited, and it is critical that the area of an OGU must be geographically and tectonically contiguous. However, it is inevitable that sampling intensity or data quality may affect the analysis results. To reduce the unevenness in different regions, we divided intensively studied regions (e.g., South China, North China, Western Europe etc.) into smaller units. In addition, stations with limited genera (less than 5; e.g., some stations in South America) are not included. Thus, a total of 58 OGUs (see Table 1) are used in our analyses. Initial genus occurrence and binary (presence/absence, “1/0”) data matrices for the three stages from 58 different paleogeographical analysis units are derived from the Mississippian brachiopod database using fossil occurrences (see Supplementary data). Four similarity coefficients, i.e., Jaccard, Dice, SI and Yule's Y, are used in this paper. The Jaccard (Jaccard, 1901) and Dice (Dice, 1945) coefficients have

been preferentially recommended and extensively used in many biogeographical studies. They are preferable when the shared taxa between two regions are demonstrated (Shi, 1993; Shen and Shi, 2000). Because of the Yule's Y coefficient's emphasis on the taxon that is included in the total sample but absent in the observed small regional samples (Yule and Hendall, 1950), it is adopted in this paper with the purpose of measuring the similarity or dissimilarity when the data contain a relatively high proportion of endemic taxa, as suggested by Huang et al. (2012). To test the possible shortcomings derived from the binary coefficients, the probabilistic index (SI) developed by Raup and Crick (1979) was also used. The SI index uses a randomization (“Monte Carlo”) procedure to compare taxon in common in the observed paleogeographic regions. It considers statistical significance in determining similarities; therefore, the biogeographical data are weighted on the basis of frequency of occurrence by this method so that widespread genera do not have a disproportionate effect on measurement of similarity (Schmachtenberg, 2008; Shen et al., 2013). When we interpret the data we will see whether the results derived from those different approaches and coefficients are consistent or not. If the results generally agree with one another based on different coefficients, then we think that the results are robust. Hierarchical Cluster Analysis (CA) is employed to calculate brachiopod fauna's distance based on Jaccard, Dice, SI, and Yule's Y similarity coefficients using PAST v. 2.12 software (Hammer et al., 2001). The Non-metric Multidimensional Scaling (NMDS) which superimposed the Minimum spanning tree (MST) was performed by PAST with the purposes of cross-verifying the results derived from the cluster analysis. NMDS is a multivariate statistical method that reduces the taxonomical space into two or three dimensions based on fauna similarities and therefore builds the linkage between the distance in plots of faunal stations (equal to OGUs in this paper) and fauna distance (Shi, 1993). Similar plots based on Jaccard, Dice, SI, and Yule's Y similarity coefficients are showed respectively, and only results based on the Jaccard coefficient are given in Fig. 6 for controlling the article in suitable length. MST usually is attached to NMDS and provides perspicuous mind about the fauna distance by finding the smallest distance between plots (Pettie and Ramachandran, 2002). Sheppard diagram and “stress value” are simultaneously calculated to quantitatively weigh the level of “goodness of fit” between the original input data matrix and the ultrametric matrix of the resultant NMDS scatter plots (Kruskal and Wish, 1978; Babcock, 1994; Shen and Shi, 2000, 2004). 3. Results and interpretations 3.1. Tournaisian The Tournaisian brachiopod faunas recorded from 34 stations worldwide are generally grouped into two distinct supergroups (Supergroups A and B) based on all coefficients (Fig. 2). Supergroup A consists of two stations only (the San Juan Basin and Sierra de Almeida, Argentina), both of which were situated in the southwestern margin of Gondwana. All other stations are grouped into Supergroup B. Further division at group level is poorly delineated. However, three groups (Groups B1, B2, and B3) can be roughly separated based on their binary coefficients. Group B1 is defined by two North American stations, i.e., Appalachian Basin and San Andres Mts., New Mexico. The Missouri River Basin is occasionally clustered with Group B1 based on the SI index. Group B2 is a geographically dispersed large complex group, embracing the OGUs in North America, Europe, China, Cimmeria, Australia, and Siberia. Clusters within Group B2 are very weak although a few vague clusters of several geographically close stations are grouped, such as stations from South China or eastern Australia. Group B3 is represented by Algeria Sahara and the Murzuq Basin, both of which were situated in northern Africa. This group, however, occasionally merged with Group B2 based on the SI and Yule's Y coefficients.


The large complex supergroup (Supergroup B) consists of stations in low latitudinal equatorial oceans and the northern hemispherical shelf regions (Fig. 3A). The relatively weak groupings among different stations at group/subgroup level apparently indicate that most of the Tournaisian brachiopods in the large paleoequatorial and north hemispherical shelf regions could easily migrate or exchange with their

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neighboring faunas and therefore obscured the paleogeographical outlines at province/subprovince level. Brachiopod faunas from stations around the shelf regions of the Paleotethys Ocean generally show consistent results within all coefficients, including South and North China, Europe (e.g., England, the Namur-Dinant Basin, and the Moscow Basin) and regions from peri-Gondwanan region (e.g., Iran,

Table 1 Mississippian brachiopod faunas of the world-basic data and stations (OGUs) for the paleobiogeographical analysis. Number

Stations/OGUs

Abb.

Tectonic units

Stratigraphic units Tournaisian

1 2 3

Alabama Alberta, Canada Algeria Sahara

Ala Ab AS

4

Amazon Basin, Brazil Appalachian Basin

AmB Amazon Basin ApB

North America Platform

6

Arctic Alaska terrane

Ak

Arctic Alaska Terrane

7

Bb

Eastern margin of Australia

8 9

Babbinboon, NSW Baoshan Block Bonaparte Basin

Ba BpB

Baoshan Block Bonaparte Basin

10 11 12 13

California Canning Basin Carnarvon Basin Cathaysian Block

Ca CB CvB Ct

North America Platform Canning Basin Carnarvon Basin South China Block

Laurel Fm. Moogooree Limestone Nanbiancun Fm.; Chuanbutou Fm.

14 15

Caucasus central and north Iran central Texas Ciudad Victoria, Mexico Daba Mts., Shaanxi E YunnanGuizhou England

Cu Ir

Cimmeria Belt Cimmeria Belt

Geran-Kalasi Mountain section Sardar Fm.

CT CV

North America Platform North America Platform

Chappel Limestone Lower–middle Vicente Guerrero Fm.

Da

South China Block

EY

South China Block

Eng

European Platform

Fermanagh Basin, N Ireland Great Basin

FB

5

North America Platform North America Platform northern Africa

Visean

Serpukhovian Bangor Fm.

Banff Fm. Gres du Khenig Fm.; Iridet Fm.; Argiles de Teguentour Fm.

Uppermost Iridet Fm. Itaituba Fm.

Lower Ramp Creek Fm.; Edwardsville Fm.; Harrodsburg Limestone Upper Woodhurst Member of Lodgepole Limestone, uppermost Kayak shale; upper Kayak shale Top of lower Burindi group

Fayetteville Shale; Pennington Fm., St. Louis Limestone; Keokuk Limestone; Upper Ramp Creek Fm.; Pride Mt. Fm. Warsaw Fm.; Salem Limestone Lisburne Group; Alapah Limestone Lisburne Group; Alapah member

Yudong Fm.; Qingshuigou Fm.; Septimus Limestone; Enga Sandstone; Ningbing Limestone; Burt Range Fm.

Shihuadong Fm.; Yunruijie Fm. Milligans Beds; Burvill Beds; Waagen Creek breccia; Utting Calcarenite Baird shale

Upper Huangjin Fm.; lower Luochen Fm.; Simen Fm.

Baizuo Fm.

Middle Zhanpo Fm.

Upper Zhanpo Fm. Zhaojiashan Fm.

European Platform

Jiusi Fm.; Shangsi Fm.; Yazitang Member of Caohai Fm. High Tor Limestone; upper Grey Limestone Subreefal Limestone

GB


Diamond Peak Fm.; Brazer Fm.

Chainman Fm; Ely Lm.; Yellow Pine Lm.

HC

European Platform

Paprotnia series

24 25 26

Holy Cross Mts., Poland Iberian Peninsula India Kashmir Japan

Ib IK Ja

Armorica (South Europe) Indian Plate Kitakami Massif

27

Jungar terrane

Jg

Jungar Terrane

Heishantou Fm.; Kuangou Fm.

28 29

Kuznetsk Basin Lhasa Block

Ku Lh

Kuznetsk Basin Lhasa Block

30

Longmen Mts.

Lm

South China Block

31

Missouri River Basin

Mo


Unnamed Yali Fm.; Luogong Fm.; Diyagxia Fm. Majiaoba Fm.; Beichuan Fm.; Changtanzi Fm. Chouteau limestone; Burlington Limestone; Boone Fm.; upper Hampton Fm.

Alba or Genicera Fm. Syringothyris Limestone top Mano Fm.; Omi Limestone; Nagaiwa Fm.; Arisu Fm. Yamansu Fm.; Nanmingshui Fm.; Aqialehe Fm.; Xibeikulas Fm. Unnamed Bariadong Fm.; Yongzhu Fm.; Yali Fm.; Lower Naxing Group Zongchanggou Fm.

32 33

Moesian Platform Me Mongolia Mn

European Platform Mongolian Continent

34

Moscow Basin

MsB

East Europe

35 36

Murzuq Basin Namur–Dinant Basin

MrB Nm

Northern Africa European Platform

16 17 18 19 20 21 22 23

Tangbagou Fm.; Muhua Fm.; Yanguan Fm. Unnamed

Upinsky; Cherepetsky; Malevsky; Upinsky Basal Mrar Fm.; Unnamed

Baogutu Fm.; Nanmingshui Fm.; Hongshanzui Group Unnamed Naxing Group Zhangbagou Fm.

Moorefield shale; Keokuk Limestone; Pella Fm.; lower Warsaw Fm. Valea Idegului Fm. Bayansair Horizon; Tsagaankhaalga Fm.; Urmugteiuul Fm. Tulsky; Aleksinsky Tarussan; Steshevsky Upper Mrar Fm. Unnamed (continued on next page)

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Table 1 (continued) Number

Stations/OGUs

Abb.

Tectonic units

Stratigraphic units Tournaisian

Visean

Serpukhovian Yeyungou Fm.; Akeshayi Group; Nanmingshui Fm.; Nugushayi Group

37

northern Tianshan

NT

Tianshan Fold Belt

Yeyungou Fm.; Meilukahe Fm., Aqialehe Fm.; Akeshayi Group

38 39

Nova Scotia Oklahoma

NS Ok

North America Platform North America Platform

Lower of Windsor series Moorefield Limestone; Boone Fm.

40 41

Pyrenees, France Qaidam Block

Py Qd

Variscan Belt Qaidam Block

42 43 44

Qiangtang Block Qilian Mountains Rocky Mts.

Qt Ql Rc

Qiangtang Block Qilian Fold Belt North America Platform

45

San Andres Mt., New Mexico San Juan Basin

SA


SJ

South America

Sct SrA

European Platform South America

49 50

Scotland Sierra de Almeida, Chile southern Ireland southern Urals

SI SU

European Platform East Europe (Baltica)

51

Tarim Block

Ta

Tarim Block

52 53

Turkey Verkhoyan Region

Tu Ve

Cimmeria Belt Eastern Siberian Platform

54

West Qinling Fold Belt

56

West Qinling Fold WQ Belt Xin'an-Songliao XS Basin Yangtze Block Yz

South China Block

Yiwa Fm.; Shimentang Fm.; Luodongke Fm. Beitongqigou Fm.; Honghutuhe Fm.; Hongshuiquan Fm. Jinling Fm.,Wanghucun Fm.

57

Yarrol Basin

YB

Yarrol Basin

58

Yukon Territory

YT

Western margin of North American Platform

46 47 48

55

Total stations

North China Block

Chengqianggou Fm.; Middle Member of Chuanshangou Fm.

Machala Fm.; Dagangou Fm.; Chengqianggou Fm.; Huaitoutala Fm. Riwanchaka Fm. Badou Fm.; Chouniugou Fm.

Sappington Fm.; lower part of Lodgepole Fm. Caballero Fm.; Lake Valley Fm.

Redoak Hollow Fm.; Fayetteville Shales Ardengost Limestone Top part of Lower Carboniferous

Dagangou Fm. Chouniugou Fm.; Huaitoutala Fm. Arco Hills Fm.

Volcan Fm.; Maliman Fm.; Agua de Lucho Fm.; La Punilla Fm. Unnamed Upper member of Zorritas Fm.

Bachu Fm.; Xiaohaizi Fm.; Gancaohu Fm.

Carboniferous Limestone Kizilskoy Fm.; Verkhoyaya Toma Horizon Heshilafu Fm.; Nugusibulake Fm.; Gancaohu Fm. Kokaksu section Chuguchanskian Horizon; Kuranakhskian Horizon; Sokol'skian Horizon; Tikinskian Horizon

Kizilskoy Fm. Kongtaiaikengou Fm.; Heshilafu Fm.; Shaquanzi Fm. Anomonanskaya Suite; Ovlachan Fm.; Tikinskian Horizon; Nudymiiskian Series Zhatiemen Fm.; Malu Fm.

Luquantun Fm.; Honghutu Fm. Datang Fm.; Yankou Fm.

Tellebang Fm.; Pond Fm.

Zimenqiao Fm.; Shidengzi Fm.; Tzemenchiao Fm. Baywulla Fm.; Woolooma Fm.; Flagstaff sandstone Hart River Fm.

34

41

25

Caucasus, Baoshan Block, and Australia). The global faunal exchange in the whole paleoequatorial region is further displayed by the statistical relationship between brachiopod faunas from western North America (e.g., the Missouri River Basin, Central Texas) and northern and eastern Australia (e.g., the Bonaparte Basin, Yarrol Basin) based on Jaccard, Dice, and SI coefficients. The Gondwanan Realm (Supergroup A) was situated in the southern high latitudes and it is represented by endemic brachiopod genera such as Azurduya, Chilenochonetes, Paurorhyncha, Sanjuania and Septosyringothyris. This characteristic brachiopod association is generally of low diversity and lacks warm-water genera. They have been usually assumed to reflect the biogeographical differentiation of the western margin of Gondwana from contemporaneous periGondwanan and Northern Hemisphere regions (Isaacson and Dutro, 1999; Taboada and Shi, 2009; Taboada, 2010). The large complex Supergroup B represents a widespread brachiopod realm in the Tournaisian, i.e., the Paleoequatorial Realm, as indicated by the calcareous foraminifers and bryozoans (Ross and Ross, 1985, 1990) and ammonoids (Korn et al., 2012). This realm includes all stations in Paleotethys, North America, and southern part of the Siberian Platform. The brachiopod faunas in these regions contain diverse and abundant productids, spiriferids, and rhynchonellids. Relatively closer relationships among the stations in the eastern and southern North America (Group B1) as well as the northern Africa (Group B3) are displayed in the dendrograms.

Brachiopod faunas from Group B1 are represented by Rhytiophora, Ericiatia, Torynifer and Voiseyella. Meanwhile those from Group B3 are characterized by containing Saharonetes, Coveenia, Mouydirhynchus and Prospira (Wendt et al., 2009; Mottequin and Legrand-Blain, 2010). This indicates that certain distance between brachiopod faunas existed in the western part of Rheic Ocean. Further subdivision within Supergroup B is virtually impossible. Very weak faunal links between North America (Group B1) and Western Europe (Group B2), as well as links between Western Europe (Group B2) and northern Africa (Group B3) are shown in the diagrams. The data suggest viable migration routes for brachiopods in the marginal shelf of Laurussia or eastern Rheic Ocean during Tournaisian.

3.2. Visean Brachiopods from 41 stations worldwide are analyzed using four coefficients during the Visean Stage. Results based on cluster analysis among brachiopod faunas reveal a generally similar pattern to that of the Tournaisian Stage with some minor changes (Fig. 4). The widely recognized Supergroup A represented by two southern hemisphere stations (the San Juan Basin and Sierra de Almeida) during the Tournaisian is absent in the Visean analysis because no brachiopods have been recorded from the southern part of the Gondwana during the Visean Stage.


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Fig. 1. Location of the selected faunal stations on modern map showing the present-day distribution (station numbers same as in Table 1).

The large complex, geographically dispersed supergroup (Supergroup B) recognized during the Tournaisian Stage dominates during the Visean Stage, that implies faunal exchange in the Paleoequatorial Realm. Further subdivision at group level can be roughly recognized (i.e., Groups B1 and B2) based on all coefficients. Group B1 that represents the North American brachiopod provinciality during the Tournaisian continued to be seen in the Visean. It consists of two stations in the southern and western North America, i.e., Oklahoma and the Great Basin, Nevada. The Verkhoyan region situated in the Boreal region is occasionally clustered with Group B1 based on the SI and Yule's Y coefficients. The complex group (Group B2) persisted in the Visean, including stations in Europe, Siberia, North and South China, Australia, and those in the peri-Gondwana regions. However, the northern Africa group (Group B3) recognized in the Tournaisian has been incorporated into Group B2 in the Visean based on all coefficients. Subdivisions within Group B2 are very weak although a vague subgroup (B2–1) is recognized based on Yule's Y coefficient. It consists of the stations around the Siberian Platform (i.e., Kuznetsk Basin, Mongolia, Yukon Territory). The large Paleoequatorial Realm represented by Supergroup B still dominated the low equatorial regions and the northern hemisphere in the Visean Stage. This realm includes stations in Europe, Siberia, China, Australia, North America, and those in the peri-Gondwanan regions (Fig. 3B). Further subdivisions at group/subgroup level within this realm are still weak although several vague clusters are recognized. The southern and western North American group (Group B1) is represented by stations in Oklahoma and the Great Basin (Fig. 4). Endemic genera such as Keokukia, Marginirugus, Diaphragmus, Marginovatia, Auloprotonia and Heteralosia in Group B1 were diversified in the Visean. They can be readily differentiated from those in the paleoequatorial region (e.g. Group B3) at group/subgroup level, indicating the incipient separation of the North American faunas. By contrast, links between the southern and western North America areas and the Boreal Verkhoyan region revealed by the SI and Yule's Y indices indicate that these regions had a close relationship in view of the absence of some common taxa in the Paleotethys Ocean. Moreover, the former Group B3 became unrecognizable and faunas from northern Africa largely grouped with Group B2 (Western Europe) in the Visean, suggesting close distance in the eastern Rheic Ocean. In addition, the differentiation of Subgroup B2–1 indicated by the Yule's Y coefficient implies the incipient paleobiogeographical separation of Siberia platform from the low latitudinal region at subprovince level. This statistical result is well explained by the absence of the characteristic brachiopod genera

outside of the Paleotethys Ocean, such as Gigantoproductus, Striatifera, Latiproductus, Linoprotonia and Overtonia. 3.3. Serpukhovian Brachiopods in the Serpukhovian Stage were relatively less diverse than those in the Tournaisian or Visean Stages in terms of total genera recorded. Most brachiopods among the 25 stations are distributed primarily in Europe, Asia and American midcontinent, along with a few genera recorded in the Amazon Basin, South America. Three supergroups (i.e., Supergroups B, C, and D) (Fig. 5) representing three brachiopod realms in the paleoequatorial region and the northern hemisphere are well recognized based on both CA and NMDS analyses. During the Serpukhovian Stage, biogeographical relationships among brachiopod faunas had drastically changed. The former huge Paleoequatorial Realm in the Tournaisian and Visean is split into two distinct realms (Fig. 3C). Supergroup B consists mainly of a large number of stations around the Paleotethys Ocean, embracing South China, North China, Tarim, Qaidam blocks and neighboring terrenes distributed in Asia, as well as the Moscow Basin, southern Urals, the Pyrenees and the eastern marginal shelf of Laurussia. Strong faunal links among this supergroup are demonstrated by tight convergence of stations between Asia and Europe (Fig. 5). Two OGUs situated in the northern margin of Gondwana, i.e., the Lhasa Block and Himalaya Tethys Zone, are also grouped with the stations in the Paleotethys Ocean based on all coefficients, suggesting close paleobiogeographical relationships between the peri-Gondwanan region and central Paleotethyan areas. Further subdivision at group/subgroup level is not recognized within Supergroup B. This large supergroup represents the Paleotethyan Realm that dominated the low latitudinal regions across the Paleotethys Ocean. It embraces diverse brachiopods which are absent outside this realm, such as Gigantoproductus, Plicatifera, Krotovia, Latiproductus and Pugilis. Supergroup C consists mainly of the North American stations, along with the Amazon Basin in the northern part of South America, which is tightly grouped and readily distinguished from other equatorial stations during the Serpukhovian. Close faunal relationships within this supergroup suggest increasing faunal connections between South and North America. Two subdivisional small groups are recognized based on the binary coefficients (Groups C1 and C2). Group C1 is represented by stations in the eastern part of North America, i.e., the Appalachian

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Fig. 2. Dendrograms of the Tournaisian brachiopods stations derived from analysis by PAST based on the four coefficients. Two supergroups (A, B) represent two distinct realms (Gondwanan and Paleoequatorial Realms) are recognized. The relatively weak grouping within Supergroup B indicates good faunal exchange, although three groups (Groups B1, B2 and B3) can be roughly recognized.

Basin and Alabama. The Great Basin and Rocky Mts. distributed in the western North America are closely clustered based on the dendrograms (Fig. 4). They are grouped with the Amazon Basin in the northern South American and readily recognized as Group C2. The convergence of these two groups implies a possible faunal migration between North and South America along the western marginal shelf areas of the American midcontinent. Supergroup C is characterized by abundant endemic taxa such as Diaphragmus, Eumetria, Reticulariina, Anthracospirifer, Marginovatia and Flexaria. Moreover, the North American Realm can be differentiated from the Paleotethyan Realm in terms of the absence of the highly diverse large productids, which are also supported by coral (Fedorowski, 1981) and foraminifer (Kalvoda, 2002) data.

Supergroup D is easily recognized in the dendrograms for close convergence of two stations, i.e., Kuznetsk Basin and Verkhoyan Region situated in the Boreal region (Fig. 5). They are tightly grouped and readily distinguished based on all four coefficients, suggesting the very high brachiopod endemism of Boreal regions and the distinct faunal separation from other two supergroups in the paleoequatorial region. The Boreal Realm as indicated by Supergroup D has been demonstrated by many marine faunas for common occurrences of cold-water genera (Bambach, 1990; Yang, 1994). The brachiopods within this realm are evidently different from those in the paleoequatorial region, with comparatively low diverse genera such as Balkhashiconcha, Lanipustula, Tomiopsis and Bailliena.


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Fig. 3. Global brachiopod paleobiogeography in the Tournaisian (A), Visean (B) and Serpukhovian (C) (with base maps modified from Scotese, 2001), together with the expected Serpukhovian (D) oceanic surface circulation pattern modified from Smith and Read (2000) (abbreviation of stations same as in Table 1).

The Mississippian paleogeography recognized by the CA analyses is supported by the NMDS analysis superimposed on MST. As shown in Fig. 6, during the Tournaisian, the two southern American stations are well separated from all other stations in the middle of the diagram (Fig. 6A). There is no distinct grouping during the Visean (Fig. 6B). In contrast, three distinct supergroups can be well separated during the Serpukhovian (Fig. 6C). Based on distribution of stations in the diagram, it can be inferred that Coordinate 1 is related to longitude because all stations in the Paleotethyan Realm are located in the right part of the diagram, whereas all other stations in the same Paleoequatorial zone are located in the left part of the figure. On the other hand, Coordinate 2 is highly likely related to the paleolatitude change as it is displayed that the high-latitude stations are distributed in the lower part of the diagram whereas all other stations in the paleoequatorial zone are in the upper part of the diagram. Thus, geographic barrier and paleolatitude-related thermal gradient are two major factors to affect the distribution of brachiopods based on the NMDS and MST analyses. In summary, provinciality was clearly enhanced during the Serpukhovian compared to the Tournaisian and Visean, with three distinct paleobiogeographical realms readily recognizable. 4. Discussions 4.1. Closure of the Rheic Ocean drastically changed the Mississippian global paleobiogeography The Mississippian is a period in Earth history during which the various landmasses were assembling into the Pangean supercontinent. The main landmass Laurussia straddled the equator, whereas Gondwana was near the South Pole and moved progressively northward throughout the Late Devonian to Mississippian (e.g., Veevers, 2004; Stampfli and Kozur, 2006; Sintubin et al., 2008; Nance et al., 2010). It is generally believed from a tectonic viewpoint that an initial episode of continental collision between the two landmasses during the Variscan Orogeny had resulted in the closure of the Rheic seaway and therefore

formed an geographic barrier for various marine faunas between the west and east (Ross and Ross, 1985; Lane and Sevastopulo, 1990; Ross and Ross, 1990; Wang et al., 2003; Aretz, 2010, 2011). The precise timing of Rheic Ocean closure, however, is hotly debated (Bozkurt et al., 2008; Nance, 2010; Nance et al., 2010; Romer and Hahne, 2010; Jastrzębski et al., 2013). Based on our study of brachiopods outlined above, we believe that the closure of the Rheic Ocean took place during the Visean. This is based on the transition from a weak provincialism in the Tournaisian to a strong provincialism in the Serpukhovian. Many warm-water paleoequatorial genera, such as Latiproductus, Pugilis, Antiquatonia, and Krotovia, occurred in Western Europe or Cathaysia in the Tournaisian other than in the peri-Gondwanan region, and mostly spread to the Tarim, Junggar terrenes in the northern mediate latitudinal areas. Moreover, the faunal interchange between east and west in the Paleoequatorial Realm was yet present because no distinct paleobiogeographical province can be recognized among the Paleoequatorial zone during the Tournaisian based on our analyses (Fig. 2), as well as the ammonoid data (Korn et al., 2012). Thus, it can be inferred that the Rheic seaway between the Central Saharan basins to NW Europe and eastern North America was still present during the Tournaisian, which connected the Paleotethys and western Panthalassan Ocean (Fig. 3A) (Golonka, 2000; Stampfli and Borel, 2002; Scotese, 2004; Stampfli and Kozur, 2006; Cocks and Torsvik, 2011). Nevertheless, as Gondwana moved progressively northward and approached to Laurussia, the Rheic Ocean became narrower, the paleolatitude-related thermal gradient was reduced, and the connection of brachiopod faunas in the Variscan Sea was increased as suggested by the cluster of northern Africa with Western Europe in the Visean Stage. In addition, the viable migration route from Europe via North Africa into the Appalachia, known as the “Afro-Appalachian link” (House, 1973; Becker and Mapes, 2010) was cut off for brachiopods during Visean and Serpukhovian. Both of the above two phenomena are most easily explained by the closure of the Rheic Ocean between Gondwana and Laurussia during late Visean and Serpukhovian. This

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Fig. 4. Dendrograms of the Visean brachiopods stations derived from analysis by PAST based on four coefficients. An incipient paleobiogeographical separation within Supergroup B is faintly indicated by the convergence of geographic close stations.

interpretation is also supported by the coral data. As documented by Aretz (2010), the Visean coral fauna found from NE Morocco comprises many taxa, which are well-known in NW Europe north to the Variscan Orogen. Its composition clearly indicates that it belongs to the Western European faunal province. The relatively weak provincialism during the Tournaisian can be explained by global ocean circulation models. As documented by many previous studies, a warm equatorial current spilled from the northeast low latitudes through the equatorial Rheic seaway, and flowed into the Panthalassan Ocean in western Pangea (Ross and Ross, 1985; Horne, 1999; Smith and Read, 2000). This narrow oceanic seaway between Laurussia and Gondwana was confirmed by the presence of many marine organisms such as corals and bryozoans

(Mckerrow et al., 2000; Aretz, 2010) and brachiopods distributed along both sides of the seaway in northern Africa and SW Europe. The oceanic seaway played an extremely important role in connecting the faunas among central Europe, the Appalachian Basin, and northern Africa, and Paleotethys during the Tournaisian. In contrast to the Tournaisian and Visean Stages, brachiopod paleobiogeography became much more distinct during the Serpukhovian Stage: three striking paleobiogeographical realms can be readily recognized, and in particular the Paleotethyan Realm can be well differentiated from the North American Realm, which strongly suggests that brachiopod exchanges between east and west had been disrupted while Gondwana and Laurussia merged together during the late Mississippian (Golonka, 2002; Nance et al., 2010; Shi and Waterhouse, 2010).


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Fig. 5. Dendrograms of the Serpukhovian brachiopods stations derived from analysis by PAST based on four coefficients. The increasing fauna provinciality is evidently indicated by three distinct supergroups (B–D), which represent three paleobiogeographical realms (Boreal, Paleotethyan, and North American realms).

After the collision, the paleoequatorial east–west oceanic current was deflected to the north and south respectively along the western coast of Paleotethys (Fig. 3D). According to the climate-biome model simulations (Poulsen et al., 2007), the closure of the Rheic Ocean would have strongly deflected the warm water equatorial currents and tropical warmth to higher latitudes along the western coastlines of the Paleotethys, reaching as far north as to northeast Asia (Kolyma regions), and as far south as to northern peri-Gondwanan region, that greatly changed the faunal distribution. Circum-Paleotethys circulation enhanced faunal similarity within the Paleotethys toward the end of Mississippian as indicated by the cluster analysis and NMDS. This

seems comparable with the distributions of extant brachiopods in the Atlantic Ocean where brachiopod species spread in the west and east margins of modern Atlantic Ocean with the Atlantic warm water circulation (Zezina, 1997; Logan, 2007; Zezina, 2008). The benthic brachiopod paleobiogeography in the Serpukhovian is generally consistent with the pelagic ammonoid paleobiogeography in which an increasing provincialism occurred from the Visean to Serpukhovian (Korn, 1997; Korn et al., 2012). Therefore, it is clear that the closure of the Rheic seaway affected both benthic and pelagic faunas and caused the disruption of the faunal exchange between the Paleotethyan areas and the North America.

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Fig. 6. Plots of fauna stations in the Tournaisian, Visean and Serpukhovian on the first two dimensions (Coordinates 1 and 2) of NMDS based on the binary Jaccard coefficient, superimposed with MST and Shepard plots (upper right) (abbreviation of stations same as in Table 1).


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Tectonic motion and ocean circulation as major controlling factors of brachiopods have been demonstrated by both extant brachiopods (Zezina, 1997; Logan, 2007; Zezina, 2008) and Permian brachiopods (Shi and Archbold, 1995, 1996; Shen and Shi, 2000, 2004; Powell, 2007; Shen et al., 2009, 2013). Tectonic and oceanic circulation played a major role in regulating marine fauna provincialism and exerted more profound influence on the Middle and Late Devonian paleobiogeography of brachiopods and bivalves (Stigall Rode and Lieberman, 2004). In addition to the geographic barrier the paleolatitude-related thermal gradient appeared to be enhanced from the Visean to Serpukhovian. As shown in Figs. 2–6, the Boreal Realm was not recognized based on our analyses, but it became well distinguished from those in the Paleotethys, which are dominated by abundant warm-water genera such as Gigantoproductus, Striatifera, Plicatifera, Linoprotonia, Latiproductus, Pugilis, Krotovia and Overtonia. Paleogeographically, the Siberian platform was situated in the northern high-latitude Boreal region where it was affected mainly by northern polar cold currents (Fig. 3D). Although oceanic connection between the Paleotethys and northern polar region was present, the brachiopods around the Siberian platform are obviously different from those in the Paleotethys, with mono- or pauci-specific genera such as Azurduya, Paurorhyncha, Sanjuania, Septosyringothyris, Balkhashiconcha, Lanipustula and Bailliena. This increasing provincialism related to the thermal gradient could be enhanced by the onset of the Late Palaeozoic Ice Age (Isbell et al., 2003; Fielding et al., 2008; Isbell et al., 2012), but further analyses are necessary to confirm this causal-effect link.

and in the southern hemisphere in Mississippian, habitable area was relatively less for brachiopods in the higher latitudes of the northern hemisphere. Our brachiopod diversity gradient pattern shown in Fig. 7 generally matches the Hubbell's neutral theory of biodiversity and biogeography (Hubbell, 2001) that is characterized by the dramatic increase in number of species and higher taxa from the poles to the tropics (Turner, 2004; Turner and Hawkins, 2004). However, it does not strictly show a consensus with the neutral theory in detail (cf., Jablonski et al., 2006). The Mississippian latitude diversity gradient indicates that most brachiopods distributed in the southern tropical zone during the Tournaisian and Visean, which covered eastern North America, Western Europe and South China. However, in Serpukhovian, most brachiopods diversified in the northern tropical zone and distributed in Western Europe, North and South China. This is apparently related to the northward movement of Gondwana during the Mississippian (Witzke, 1990). As Raymond et al. (1989) and Kelley et al. (1990) documented, most brachiopod genera moved northward during a temporary warming of the high-latitudes due to the northward deflection of the circumequatorial currents as the collision of Laurussia and Gondwana progressed during the middle and late Mississippian. On the other hand, narrowly distributed, largely highly diversified genera in the tropical zone were eliminated when glaciation began in southern hemisphere in Serpukhovian (Powell, 2005, 2007). Therefore, diversity pattern as well as biogeographical structure of the brachiopod fauna was affected by the climate change associated with the onset of the LPIA as well.

4.2. Paleolatitude diversity gradients

5. Conclusions

The Mississippian latitudinal diversity gradient of brachiopods is evident. As shown in Fig. 7, more than 70% of overall brachiopods were recorded in the paleoequatorial region and constricted within 30° during the Mississippian. Less than 1/3 of total genera are found in higher latitude than 30°. This is partially resulted from the available habitats. Since the continents are concentrated around the equator

The Mississippian global paleobiogeographical analysis of brachiopod fauna suggests that provincialism was weak during the Tournaisian and Visean. Two realms, i.e., Gondwanan and Paleoequatorial realms can be recognized during the Tournaisian, and the Paleoequatorial Realm was not differentiated between east and west. A dramatic change in global paleobiogeography and latitudinal diversity gradient occurred from the

Fig. 7. Paleolatitude gradient of brachiopod genus diversity during three successive stages of the Mississippian. Arrows indicate migration of brachiopod diversity center within paleolatitude belts from the Tournaisian to Serpukhovian.

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late Visean to Serpukhovian. The Paleotethyan and North American realms in the paleoequatorial region were well differentiated during the Serpukhovian owing to the closure of the east–west Rheic Ocean, that disrupted the faunal interchanges between the Paleotethys in the east and North America in the west. In addition, the closure of Rheic Ocean deflected the warm water equatorial currents and tropical warmth to higher latitudes along the western coastlines of the Paleotethys and formed circum-Paleotethys currents, which greatly enhanced faunal similarity within the Paleotethys and global thermal gradient in the Serpukhovian. Thus, the Paleotethyan Realm was well differentiated from the Boreal Realm during the Serpukhovian. Further latitudinal diversity gradients indicate a distinct northward shift of diversity center from the Visean to Serpukhovian. The shift of diversity center was controlled by both collision between Gondwana and Laurussia and the beginning of LPIA during the Visean and Serpukhovian. Thus, the plate tectonics in the late Mississippian not only caused the closure of the Rheic Ocean and the formation of the supercontinent, but also drastically changed the global paleobiogeography and faunal latitudinal diversity pattern. Acknowledgments Many thanks to Dr. Zhang Hua (NIGPAS) for the help on the buildup of the primary brachiopod database platform, and to Pan Jianxin for the assistance on compiling the dataset. We thank Dr. Miao Desui from Kansas University and Dr. Joseph G. Meert from the University of Florida for the help in improving the English grammar. We also thank Lucia Angiolini from the University of Milan and Markus Aretz from the University of Toulouse for constructive comments on the manuscript. This paper is supported by the National Natural Science Foundation of China (41290260). Appendix A. Supplementary data Supplementary data to this article can be found online at http://dx. doi.org/10.1016/j.gr.2013.09.013. References Angiolini, L., 2013. Guadalupian (Middle Permian) paleobiogeography of the Neotethys Ocean. Gondwana Research 24, 173–184. Aretz, M., 2010. Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif (NE Morocco): taxonomy, biostratigraphy, facies and palaeobiogeography. Palaeontologische Zeitschrift 84, 323–344. Aretz, M., 2011. Corals from the Carboniferous of the central Sahara (Algeria): the collection “Marie Legrand-Blain”. Geodiversitas 33, 581–624. Babcock, L.E., 1994. Biostratigraphic significance and paleogeographic implications of Cambrian fossils from a Deep Core, Warren County, Ohio. Journal of Paleontology 68, 24–30. Bambach, R.K., 1990. Late Palaeozoic provinciality in the marine realm. In: McKerrow, W.S., Scotese, C.R. (Eds.), Palaeozoic Palaeogeography and Biogeography. The Geological Society Memoir, 12. Geological Society of London, London, pp. 307–324. Bambach, R.K., Knoll, A.H., Wang, S.C., 2004. Origination, extinction, and mass depletions of marine diversity. Paleobiology 30, 522–542. Becker, R.T., Mapes, R.H., 2010. Uppermost Devonian ammonoids from Oklahoma and their palaeobiogeographic significance. Acta Geologica Polonica 60, 139–163. Bozkurt, E., Pereira, M.F., Strachan, R., Quesada, C., 2008. Evolution of the Rheic Ocean. Tectonophysics 461, 1–8. Caplan, M.L., Bustin, R.M., 1999. Devonian–Carboniferous Hangenberg mass extinction event, widespread organic-rich mudrock and anoxia: causes and consequences. Palaeogeography, Palaeoclimatology, Palaeoecology 148, 187–207. Cocks, L.R.M., Torsvik, T.H., 2011. The Palaeozoic geography of Laurentia and western Laurussia: a stable craton with mobile margins. Earth-Science Reviews 106, 1–51. Crovello, T.J., 1981. Quantitative biogeography: an overview. Taxon 30, 563–575. Davydov, V.I., Korn, D., Schmitz, M.D., Gradstein, F.M., Hammer, O., 2012. The Carboniferous Period. Chapter 23 The Geologic Time Scale.Elsevier, Boston 603–651. Dice, L.R., 1945. Measures of the amount of ecological association between species. Ecology 26, 297–302. Fedorowski, J., 1981. Carboniferous corals: distribution and sequence. Acta Palaeontologica Polonica 26, 87–160. Fielding, C.R., Frank, T.D., Birgenheier, L.P., Rygel, M.C., Jones, A.T., Roberts, J., 2008. Stratigraphic imprint of the Late Palaeozoic Ice Age in eastern Australia: a record of alternating glacial and nonglacial climate regime. Journal of the Geological Society 165, 129–140.

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