Palaeobiogeography and Diversification of Tournaisian–Viséan Bryozoans (Lower–Middle Mississippian, Carboniferous) From

Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Palaeobiogeography and diversiﬁcation of Tournaisian–Viséan bryozoans (lower–middle Mississippian, Carboniferous) from Eurasia

Zoya Tolokonnikova a,⁎, Andrej Ernst b, Patrick N. Wyse Jackson c a Kuban State University, Kazan Federal University, 353235, Aphipskij, Post box 30, Krasnodar, Russia b Institut für Geologie, Universität Hamburg, Bundesstr. 55 20146 Hamburg, Germany c Department of Geology, Trinity College, Dublin 2, Ireland article info abstract

Article history: The Mississippian represented the last diversification event of bryozoans in the Palaeozoic which affected all tax- Received 28 May 2014 onomic levels. Within the borders of modern Eurasia Tournaisian–Viséan bryozoans are known from 24 areas, Received in revised form 21 August 2014 with a total of 878 species in 180 genera. Palaeobiogeographical analysis is here made according to the stages Accepted 26 August 2014 and substages of the Tournaisian and Viséan. Six characteristic species were identified for the lower Tournaisian Available online 10 September 2014 which are distributed in more than one region of the continent, and for the upper Tournaisian there are two such species. Eleven species occur in the narrow stratigraphic interval of the lower Viséan that display a wide geo- Keywords: Bryozoans graphic distribution within Eurasia, in the upper Viséan there are 14 such species. Analysis of the generic compo- Tournaisian sition showed the closest similarity between Tournaisian bryozoans of Kazakhstan, the Kuznetsk Basin and Viséan Eastern Transbaikalia. Significant similarity in the generic composition of Germany, Britain, Ireland and France Palaeobiogeography is observed during the Viséan. Two centres of radiation bryozoans were identified: «Eastern» and «Western». Eurasia © 2014 Elsevier B.V. All rights reserved.

1. Introduction than in isolated basins. Buttler et al. (2013) have reported on Early Palaeozoic biogeography, and Ross (1982) and Ross and Ross (1985) Bryozoans are benthic organisms playing important role in marine provided a summary of Late Palaeozoic (Carboniferous–Permian) bio- ecosystems from the Ordovician to the recent. Evolution of the phylum geography, while Ross and Ross (1982) discussed factors that influ- Bryozoa displays several significant diversification and succeeding enced faunal distributional patterns. At the level of the geological extinction events, one falling in the Frasnian/Viséan interval. Rise in ge- period, a number of overviews of bryozoan biogeography throughout neric diversity is observed from the end of the Famennian reaching a the Palaeozoic have been documented either globally or from a more maximum at the end of Viséan and followed by extinction of several restricted viewpoint geographically or temporally: Ordovician (Ross, taxa in the Serpukhovian (Ernst, 2013). During Famennian–Viséan 1982; Tuckey, 1990a), Silurian (Tuckey, 1990b; McCoy and Anstey, ages bryozoans experienced crucial changes in their taxonomic compo- 2010), Devonian (Bigey, 1985; Tolokonnikova and Ernst, 2010), Carbon- sition (e.g., the rapid appearance of new families and suborders; extinc- iferous (discussed below), and Permian (Ross, 1978, 1979; Gilmour and tion of Devonian genera and families; change of dominant order). The Morozova, 1999). Mississippian represented the last diversification event of bryozoans The first overview of the palaeobiogeography of Carboniferous bryo- in the Palaeozoic. zoans was made by Ross (1981). She analysed generic diversity for 11 An analysis of the diversity of orders of Palaeozoic bryozoans, includ- regions of the world according to stages of the Carboniferous and ing those from the Carboniferous, was made by Gorjunova et al. (2004). discussed changes in the composition of families and orders. Ross rea- They came to the conclusion that the diversity depends on the develop- soned for continuity between Devonian, Carboniferous and Permian mental phase of the higher taxon (bloom, decline, extinction or origina- bryozoans, and inferred bryozoan diversification and radiation during tion) under secondary influence of abiotic factors. the Viséan, which was apparently induced by such abiotic factors as General biogeographical analyses have been provided for most of fluctuating sea-levels driven by glaciations in Gondwana, and the loss the Palaeozoic. Naimark et al. (1999) suggested that faunal radiations of shallow-water environments due to the closure of the ocean between across different regions occurred diachronously, and that endemic Euramerica and Gondwana (Ross, 1982). During the Late Carboniferous species developed in areas that promoted bryozoan habitats rather an increased number of new ecological niches developed due to a grad- ual warming of the climate that resulted in the origination of some new bryozoan lineages and a pronounced provincialism (Ross and Ross, ⁎ Corresponding author at: 353235 Aphipskij, Post box 30, Krasnodar Region, Russia. E-mail addresses: [email protected] (Z. Tolokonnikova), [email protected] 1985). However, the second half of the Carboniferous is generally (A. Ernst), [email protected] (P.N. Wyse Jackson). marked by a reduction in taxonomic diversity (Ross, 1981). McKinney

(1993) documented the distribution of the distinctive fenestrate Archi- 1983; Gorjunova, 1985; Popeko, 2000; Ariunchimeg, 2010). This has medes from the Carboniferous of North America and argued that chang- allowed regional correlation between these areas and the establishment es in its life-history during that period resulted in a change in its of biostratigraphical units (beds with bryozoans) for Mongolia and distribution from shallow water shelf areas during the early Carbonifer- bryozoans zones for Eastern Transbaikalia which are utilized for local ous to deeper basinal facies towards the end of the Carboniferous. correlation (Popeko, 2000; Ariunchimeg, 2010). On account of their richness and abundance in marine deposits, it On other regions bryozoans have been described from segmental has been recognised that bryozoans have a considerable potential for stratigraphical levels, and this has resulted in difficulties in their analysis use in stratigraphy (e.g., Nekhoroshev, 1925; Bancroft, 1987). Their and interregional correlation on the coeval deposits. The scheme of the wide distribution is ensured by passive dispersion of larvae with current Mississippian deposits of Eurasia is shown in Table 1. Asterisks mark system or floating objects (for example, seaweeds, pumice). As all living levels for which the exact temporal distribution of the bryozoans is un- organisms, development and diversity of bryozoans depend on abiotic known, or bryozoan records are associated with certain biostratigraphical and biotic factors. The most limiting ones for the development of the zones. Grey shadowing marks intervals from which bryozoans are colonies of modern bryozoans are water temperature, salinity, food unknown. supply, substrate and predation (Taylor and Larwood, 1990). It can be In the lower–middle Mississippian deposits of Eurasia records of 878 inferred that the same factors had similar effects on bryozoans during species belonging to 180 genera have been databased for this study. the Mississippian. They represent 41 families in 4 orders (Table 2). Unfortunately, no During the last few decades a considerable volume of new data on detailed information about the stratigraphical distributions of the ma- Carboniferous bryozoans has been obtained for Eurasia (e.g., Yang jority of species is known. For many species, the range covers a stage et al., 1988; Wyse Jackson, 1996; Ernst, 2005), while for other conti- or even two. Therefore, it was only possible to use the following inter- nents such data are scarce. During this time, bryozoan taxonomy has vals for palaeobiogeographical comparisons: lower Tournaisian, upper been revised significantly and new information on bryozoan distribu- Tournaisian, undivided Tournaisian, lower Viséan, upper Viséan, and tions have been obtained. For other continents such as North America undivided Viséan. a number of monographic and other taxonomic studies have appeared Palaeobiogeographical analysis was carried out using an unweighted (Snyder, 1991a,b; Gilmour and McColloch, 1995) but studies carried pair-group average method (UPGMA) in which clusters are joined out in the late 1890s and first half of the twentieth century require reas- based on the average distance between all members in the two groups sessment. Furthermore, understanding of the stratigraphical framework (Hammer et al., 2001). Similarity was measured using the Jaccard simi- for some regions in Eurasia has been improved. This increase in data has larity coefficient for absence–presence data (Jaccard, 1901). This coeffi- permitted this new compilation to document in considerable detail the cient measures similarity between sample sets, and is defined as the size palaeobiogeography of bryozoans of Eurasia during Tournaisian–Viséan of the intersection divided by the size of the union of the sample sets. time which was an important stage in the development of the phylum Cluster analysis was performed using PAST software version 1.97 Bryozoa. (Hammer et al., 2001). Palaeogeographical distribution of bryozoan genera was shown using of Principal Coordinates analysis performed 2. Geological setting in PAST. Only species with adequate taxonomic description were utilized for the analysis, whereas those species with inadequate descrip- According to the International Stratigraphic Time, the Tournaisian tions, without illustrations or originally left in open nomenclature were and Viséan belong to the Mississippian — the lower part of the Carbon- ignored. iferous system (Cohen et al., 2013). The lower limit of the Tournaisian is dated as 358.9 ± 0.4 Ma, and the Viséan ranges from 346.7 ± 0.4 Ma to 4. Palaeobiogeography of Tournaisian bryozoans 330.9 ± 0.2 Ma; the duration of the Tournaisian is 12.2 Ma and of the Viséan 15.8 Ma. At the end of the Famennian, bryozoans show intensive diversifica- The territory of modern Eurasia is built up of many continental tion at the species and genus levels (Tolokonnikova and Ernst, 2010; blocks and oceanic depressions, which were situated in different Ernst, 2013). A significant number of taxa which appeared at the end hemispheres during the Mississippian (Stampflietal.,2013). The north- of the Devonian continued their existence or even flourished in the ern part of Eurasia belonged to Laurasia, and the southern part to Early Carboniferous. The Devonian/Carboniferous boundary is marked Gondwana. According to existing palaeogeographical reconstructions, by an increase in extinction rates of bryozoans (Ernst, 2013). In continuous climatic cooling and aggregation of the terrains of Laurasia the Early Carboniferous bryozoans showed a marked increase in mor- occurred at the beginning of the Carboniferous (Scotese, 2001). phological specialization, especially among members of the orders Fenestrata and Cryptostomata. A similar trend is also observed in the 3. Materials and methods brachiopods. The Strunian (latest Famennian) brachiopods have a characteristic aspect of a transitional fauna consisting of a minority of Within the area of modern Eurasia, bryozoans of Tournaisian–Viséan Devonian to prevalent Carboniferous forms (Conil et al., 1986). Analyses age are known from 24 areas. Tournaisian bryozoans have been report- of brachiopod fauna from southern China reveal that after a decline ed from Germany, Ireland, Poland, the Donetsk Basin, Nakhichevan in the generic diversity at the Devonian–Carboniferous boundary, (or Transcaucasia in text below), Kazakhstan, Russia (Southern Urals, the Early Tournaisian fauna shows slight impoverishment (Sun and Eastern Transbaikalia, Kurgan area, Russian Platform, Kuznetsk Basin), Baliński, 2011). In the middle Tournaisian brachiopods experienced an Mongolia and China. Viséan bryozoans are known from Ireland, explosive increase in diversity at the generic level. The brachiopods con- Germany, Britain, Spain, France, the Donetsk Basin, Poland, Russia stitute a fully recovered high diversity fauna consisting of forms (Middle and Northern Urals, Eastern Transbaikalia, Kurgan area, representing a wide spectrum of attachment strategies as well as highly Russian Platform, Kuznetsk Basin, Northeast), Afghanistan, Uzbekistan, specialised forms adapted to special kinds of ecological niches. Iran, Turkmenistan, Kazakhstan, Mongolia, China and Japan. Shen et al. (2006) in an analysis of brachiopod diversity of the same The current levels of information on the bryozoan faunas from these region suggest that this recovery occurred slightly later during the territories are disproportional. Successive following one another Serpukhovian. Similar patterns of diversification in the Mississippian complexes were described from the lower Tournaisian–upper Viséan have been noted in Argentina where the number of bivalve genera dou- deposits of Kazakhstan, Mongolia, Eastern Transbaikalia and the bled between the Early and Late Carboniferous while brachiopods have Kuznetsk Basin (Nikiforova, 1948, 1950; Nekhoroshev, 1953, 1956; shown a more than six-fold increase in generic diversity (Sterren and Trizna, 1958; Plamenskaya, 1964; Troitzkaya, 1975; Plamenskaya, Cisterna, 2010). With regard to crinoids, 69 genera are known in the 202 Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

Table 1 Correlation of regional stratigraphical units of Tournaisian and Viséan ages in Eurasia. Shaded parts are units from which no bryozoans have been reported.

Famennian, but following the end-Devonian extinction they diversified by bryozoans–Kurgan region and Ireland (Wyse Jackson and Buttler, rapidly, with 146 genera documented in the Tournaisian and an 1994; Tolokonnikova, 2012). In addition, bryozoans are known from additional 23 genera in the Viséan (Kammer and Ausich, 2006). the undivided Tournaisian deposits of Eurasia in the following regions: According to Simakov (1993) the most important consequence of Germany, the Donetsk Basin, Poland and China (Nekhoroshev, 1932; the D/C event was not a mass extinction of Devonian taxa but the ap- Dunaeva, 1964a,b, 1969; Yang et al., 1988; Lu, 1999; Morozova et al., pearance of new forms of a Carboniferous aspect. Kammer and Ausich 2006). (2006) attribute the increase in crinoid diversity to increased circulation About 80% of the total number of lower–middle Mississippian bryo- and stenohaline conditions on carbonate ramps. Li and Shen (in press) zoan species is endemic. The rest includes species which are known have documented that during the Tournaisian and Viséan brachiopod from several regions of Eurasia. For instance, in the lower Tournaisian faunas exhibited little provincialism, but that major changes in their of Eurasia six species occur in more than one region, but in the upper distribution occurred during the Serpukhovian. Tournaisian only two such species occur. For the undivided Tournaisian Tournaisian deposits of Eurasia contain 362 bryozoan species belong- deposits 11 species occur in more than one region. There are 19 species ing to 98 genera, of which 51 species occur in undivided Tournaisian– in the Tournaisian occurring in 2–3 regions which are now situated Viséan sediments. rather far apart geographically on the modern globe (Table 3, Fig. 2). Bryozoans have been described from the lower Tournaisian deposits They can be used for dating the oldest deposits of the Carboniferous. of Kazakhstan, Mongolia, Transcaucasia, the Russian Platform, Southern For the Tournaisian a significant similarity of species complexes from Urals, Kuznetsk Basin and Eastern Transbaikalia (Nekhoroshev, 1956; Kazakhstan, Mongolia, the Kuznetsk Basin and Eastern Transbaikalia, Trizna, 1958; Lavrentjeva, 1970, 1974; Popeko, 2000; Gorjunova and which are established at the level of substages, is noted. This allows Lavrentjeva, 2007; Ariunchimeg, 2010; Tolokonnikova, 2011)(Fig. 1). the use of bryozoans for detailed regional correlation. In the second half of the Tournaisian, bryozoans are unknown from At the generic level, significant similarity is observed between the the Russian Platform and Southern Urals. At this time, besides the bryozoan faunas of Kazakhstan, the Kuznetsk Basin and Transbaikalia. existing areas of the lower Tournaisian, new regions became occupied Similarity between regions of Eurasia is shown in Fig. 3. A major similarity Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211 203

Table 2 Table 2 (continued) Distribution of bryozoan genera in the Tournaisian and Viséan. I — Tournaisian, II — Viséan. Abbreviations: A — ﬁrst appearance; E — extinction; D — decline; + − presence. Order Genera I II Remarks Megacanthopora +A Order Genera I II Remarks Nicklesopora ++ Cystoporata Family Fistuliporidae Paranicklesopora ++ Cyclotrypa ++ Pseudorhabdomeson +A Fistulipora ++ Family Nicklesoporidae Eridopora ++A Europora + Cystiramus +E Family Rhomboporidae Dybowskiella + Rhombopora ++ Family Cheilotrypidae Saffordotaxis ++ Cheilotrypa ++E Klaucena + Family Cystodictyonidae Shishoviclema + Sulcoretepora ++ Primorella ++ Acrogenia + Megacanthoporina ++ Family Goniocladiidae Family Streblascoporidae Goniocladia + Ipmorella + Goniocladiella + Strebloascopora + Ramiporidra +A Family Hyphasmoporidae Family Hexagonellidae Hyphasmopora + Glyptopora + Streblotrypa ++ Galtopora +A Family Arthrostylidae Dichotrypa ++A Pseudonematopora ++ Prismopora + Nematopora ++ Evactinopora + Family Nikiforovellidae Fistulamina ++ Nikiforovella ++ Meekopora ++ Streblotrypella ++ Meekoporella + Family Nematotrypidae Trepostomata Family Eridotrypellidae Clausotrypa ++ Eostenopora +E Family Nematoporinae Eridotrypella ++E Hexites ++ Family Aisenvergiidae Family Mediaporidae Volvovachia +A Mediapora ++ Aisenvergia +A Cryptostomata Family unknown Family Anisotrypidae Taeniodictya ++ Anisotrypa ++ Cryptostomata Family Intraporidae Family Dyscritellidae (suborder Ptilodictyina) Intrapora + Eodyscritella + Family Rhinidyctyidae Dyscritella ++ Jurggarotrypa + Pseudobatostomella ++ Family Worthenoporidae Family Batostomellidae Worthenopora +A Batostomella + Family Mysticellidae Family Atactotoechidae Mysticella +E Leptotrypa +D Family Timanodictyidae Leptotrypella +E Timanodictya + Family Trematoporidae Fenestrata Family Fenestellidae Neotrematopora ++E (suborder Fenestellina) Penniretepora ++ Raissiella + Fenestella ++ Minussina +E Permofenestella +A Family Stenoporidae Parafenestella + А Tabulipora ++A Lyropora + А Parastenodiscus + Ispanayella +A Stenophragmidium ++ Paraseptopora ++ Stenodiscus ++A Minilya ++ Stenopora ++A Rectifenestella ++ Nikiforopora ++ Spinofenestella ++ Coeloclemis + Laxifenestella ++ Family Amplexoporidae Levifenestella +A Triznotrypa ++ Fabifenestella +A Family Ulrichotrypellidae Ptylopora ++ Petalotrypa +E Hemitrypa ++E Ulrichotrypella +A Pseudounitrypa +A Family Heterotrypidae Exfenestella ++ Leioclema + + D/E Lyrocladia +A Family Crustoporidae Flexifenestella ++ Tabuliporella ++ Ptilofenestella +A Crustopora ++ Filites ++ Hinaclema +A Narynella +A Nipponostenopora +A Lunofenestella +A Hunanopora + Ptiloporella + Family Coelotubuliporidae Diploporaria ++A Coelotubulipora + Family Fenestraliidae Cryptostomata Family Rhabdomesidae Fenestralia +A (suborder Rhabdomesina) Ascopora ++A Parafenestralia ++A Orthopora + Family Septoporidae Euthyrhombopora + Septopora ++A Rhabdomeson ++A Ignotifenestella +A

(continued on next page) (continued on next page) 204 Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

Table 2 (continued) Viséan deposits. Lower Viséan bryozoans are known from eight regions

Order Genera I II Remarks of the palaeocontinent (Fig. 1). Most diverse are the assemblages of Kazakhstan, Uzbekistan, the Kuznetsk Basin, Transbaikalia, and Fenestrata Family Acanthocladiidae Mongolia (Nikiforova, 1948; Nekhoroshev, 1953, 1956; Trizna, 1958; (suborder Fenestellina) Acanthocladia ++APlamenskaya, 1964; Michno and Balakin, 1975; Plamenskaya, 1983; Thamniscus + Gorjunova, 1996; Ernst, 1998; Popeko, 2000; Ariunchimeg, 2010). Less in- Baculopora +A formation exists about lower Viséan bryozoans of the Russian Platform, Arborocladia ++A Kurgan region and Iran (Schulga-Nesterenko, 1951; Lavrentjeva, 1985, Polyporella ++ Fenestrellina +A 1997; Tolokonnikova, 2012; Tolokonnikova and Yazdi-Moghadam, 2013). Mackinneyella ++A Upper Viséan bryozoans are known from 11 areas of Eurasia. Bryozoan Anastomopora + assemblages from Ireland, Kazakhstan, Mongolia, the Kuznetsk Basin Protoretepora +A and Transbaikalia have been studied in detail. Several species have Eulyra +A been identified from the upper Viséan deposits of Japan, the Donetsk Kazarchimedes +A Polypora ++ Basin, Russian Platform, Northern Urals and Spain (Schulga-Nesterenko, Pseudopolypora ++A 1951, 1955; Trizna, 1961; Sakagami, 1962, 1963; Dunaeva, 1964a,b; Gorjunopora +A Ernst and Rodriguez, 2013). A diverse bryozoan fauna occurs in the Family Semicosciniidae upper Viséan of France (Ernst et al., submitted for publication). Semicoscinium +D Quadrisemicoscinium +D In deposits of Viséan age without precise stratigraphical information Eosemicoscinium +D bryozoans are known also from China, Poland, Kirgizia, Turkmenistan, Neoreteporina +A northeastern Russia, Germany, Britain, Ireland and Afghanistan Reteporina ++D (Nekhoroshev, 1932; Nikiforova, 1933, 1950; Termier and Termier, Family Admiratellidae 1971; Morozova, 1981; Xia, 1986; Wyse Jackson, 1988; Ernst, 2005; Admiratella +A Admiranda +A Weber and Wyse Jackson, 2006). Fenestrata Family Chasmatoporidae All of these areas contain predominantly endemic species. However, (suborder Phylloporinina) Bashkirella + some species were recorded, which appear in two or four areas in the Family Chainodictyonidae same time interval (Table 4, Fig. 2). In lower Viséan deposits 11 shared Chainodictyon +A Rhombocladia +A species were found, in upper Viséan 14. In undivided Viséan deposits 32 shared species occur. In total, 57 species shared by more than one region is observed between generic assemblages of Transbaikalia, Kazakhstan in Eurasia are known. A similarity of bryozoan faunas from Kazakhstan, and the Kuznetsk Basin, which also resemble the cluster China–Ireland. Transbaikalia, Mongolia and Uzbekistan at the species level is recogniz- All these regions share genera with Mongolia, the Kurgan region and able. These «Eastern» regions do not contain common taxa found within Afghanistan. Germany, Kirgizia, the Donetsk basin and the Russian the «Western» regions. Among European countries, Germany, Ireland Platform form a separate cluster. Shared taxa occur between the Southern and Britain contain assemblages of bryozoans which show no similari- Urals and Britain; however, they are distant from other groups. ties to the «Eastern» taxa. Transcaucasia and Poland contain the most endemic bryozoans. For the undivided upper Tournaisian–lower Viséan deposits three shared taxa were evident which occur in three to five regions of Eurasia. Species which are known in five to ten regions, show relatively wide 5. Palaeobiogeography of Viséan bryozoans stratigraphical distributions from the lower Tournaisian or even the Famennian to the Serpukhovian inclusively. There are 18 such species, Viséan deposits of Eurasia contain 563 bryozoan species belonging to but revision and re-study of type material are probably necessary to 111 genera, of which 51 species occur in the undivided Tournaisian– confirm their identification.

Fig. 1. Palaeogeography and locations of known lower Mississippian bryozoan faunas (circles — Tournaisian; triangles — Viséan). Palaeogeographical and palaeoclimatic reconstructions after Scotese (2001).1— Kazakhstan, 2 — Urals (Middle, Northern and Southern Urals), 3 — Kirgizia, 4 — Kurgan region, 5 — Uzbekistan, 6 — Kuznetsk Basin, 7 — Eastern Transbaikalia, 8 — Northeast of Russia, 9 — Mongolia, 10 — China, 11 — Japan, 12 — Turkmenistan, 13 — Russian Platform, 14 — Transcaucasia, 15 — Donetsk Basin, 16 — Germany, Belgium and Poland, 17 — Ireland, 18 — Britain, 19 — Spain, 20 — France, 21 — Iran, and 22 — Afghanistan. Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211 205

Table 3 Stratigraphical distribution of common bryozoan species by regions in the Tournaisian. + − presence.

Stage Substage No. Species Number Kuznetsk Kazakhstan Eastern Kurgan Mongolia Kirgizia China areas basin Transbaikalia region

Tournaisian Undivided 1 Klaucena aculeus 3+ + + 2 Fenestella quadrulla 2++ 3 Sulcoretepora nurensis 2+ + 4 Sulcoretepora tomiensis 2++ 5 Fenestella pseudoirregularis 2+ + 6 Fenestella sergunkovae 2+ + 7 Rectifenestella glabra 2++ 8 Triznotrypa tenuilignata 4+ + + + 9 Penniretepora subangulata 2++ 10 Parafenestralia bukhtarmensis 3++ + 11 Tabulipora incrustans 3++ + Upper 12 Rhombopora ﬂoriformis 2+ + 13 Rectifenestella analoga 2+ + Lower 14 Raissiella tabulata 2++ 15 Anastomopora ovalifenestra 2++ 16 Neoreteporina altaica 2++ 17 Hemitrypa altaica 2++ 18 Pseudobatostomella minima 2++ 19 Leioclema tubulosa 3++ +

Generic similarity between regions of Eurasia is shown in Fig. 4. Uzbekistan; in the second and smaller one France, Germany, Belgium, The greatest similarity is observed between European bryozoan Britain and Ireland are included. These faunas are unique and do not faunas (Germany, Britain, Ireland, and France). Another large cluster is contain common species (except for the cosmopolitan species). They formed by regions in the Asian part of the continent (Kazakhstan, are endemic at the genus level at 90.8% in the Tournaisian and 88.8% Uzbekistan, Mongolia, Transbaikalia and northeastern Russia). The in the Viséan. At the start of the Tournaisian the «Eastern» radiation cen- most endemic are faunas from the clusters Iran–Spain and Kirgizia– tre comprised the Palaeothetys, whereas the European centre of radia- Afghanistan. Other regions are indirectly related to «Eastern» and tion belonged to the Rheic Ocean. Separate parts of the centres show «Western» centres. close similarities with each other. In Mississippian «Eastern» centre have been located in 20°–40° palaeolatitudes of northern hemisphere, 6. Discussion and «Western» centre — 5°–10° palaeolatitudes of southern hemisphere. They are located in different climatic conditions (the first centre in During the Tournaisian–Viséan phase of the development of the warm climate, second centre in arid climate according to reconstruction phylum Bryozoa there is an increase in species richness (362 in the in Scotese, 2001). Probably, that factor affected on separate bryozoan Tournaisian vs. 563 in the Viséan), and a decrease of species endemism species between radiation centres. (19 shared species occurring in 2–3 territories in the Tournaisian vs. 57 Tournaisian–Viséan deposits of Eurasia contain 193 species in total shared species in 2–4 territories in the Viséan). Two centres of radiation which occur in more than one region. The majority of the species are can be identified within Eurasia in the lower Mississippian, which are distributed in 2 to 3 regions of the continent. Most of the shared species here called «Eastern» and «Western». The first and larger one includes occur in Kazakhstan — 116 of 193 species (60%) — followed by Mongolia Kazakhstan, Mongolia, the Kuznetsk Basin, Eastern Transbaikalia and —60 species (31%), the Kuznetsk Basin and Eastern Transbaikalia — 62 and 54 species, respectively (32% and 28%) and Uzbekistan — 52 species (27%). Ireland contains 26 shared species (13%), Germany and the Kurgan region contain 25 and 29 species, respectively (13% and 15%) and France 14 species (7%). Only a few shared species are known for Afghanistan, Poland, the Donetsk Basin, Southern Urals and Japan. Obviously, this fact can be explained by the low level of study of these regions or by a high level of endemism (as in the case of Japan and the Donetsk Basin). At the level of orders a decrease of richness can be observed in Trepostomata (reduction of 8 genera in two families), rapid diversification in Fenestrata (appearance of 31 genera), and a moderate renewal of the generic composition in Cystoporata.

7. Conclusions

Results of the analysis of compiled data showed the following changes in the taxonomic composition of the phylum Bryozoa from the beginning of the Tournaisian to the end of the Viséan in Eurasia:

- Species richness increased 1.5 times and generic richness increased slightly (by 1.1 times). - The number of areas containing bryozoans increased from 14 to 22. Fig. 2. Histogram showing palaeogeography of bryozoan species known in more than one - Endemism decreased and palaeobiogeographical connections in- region. creased. 206 Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

Fig. 3. Palaeobiogeographical afﬁnities of bryozoan assemblages of Eurasia in Tournaisian performed by: 3a — Cluster analysis, 3b — Principal coordinates analysis. All analyses performed with PAST (Hammer et al., 2001). Abbreviations: Kaz — Kazakhstan, Kirg — Kirgizia, Kurg — Kurgan region, Uzb — Uzbekistan, Kuzb — Kuznetsk Basin, Tr Bai — Eastern Transbaikalia, NE — Northeast of Russia, Mong — Mongolia, Jap — Japan, Turk — Turkmenistan, Rus Pl — Russian Platform, Tr Cau — Transcaucasia, Don — Donetsk Basin, Germ — Germany, Pol — Poland, Ire — Ireland, Brit — Britain, Spa — Spain, Fran — France, and Afg — Afghanistan.

- Many new genera appeared within the Order Fenestrata, and there was a Buttler, C.J., Wyse Jackson, P.N., Ernst, A., McKinney, F.K., 2013. A review of the early Palaeozoic biogeography of bryozoans. In: Harper, D., Servais, T. (Eds.), Early reduction of genera in the Order Trepostomata. Palaeozoic Palaeobiogeography and Palaeogeography. Geological Society 38, pp. - Characteristic species of bryozoans can be identified for substages and 145–155. stages of the lower–middle Mississippian, which occur in more than one re- Cohen, K.M., Finney, S.C., Gibbard, P.L., Fan, J.-X., 2013. The ICS International Chronostratigraphic Chart. Episodes 36 (3), 199–204. gion (e.g. species Rhabdomeson progracile, Pseudonematopora planatus, Conil, R., Dreesen, R., Lentz, M.A., Lys, M., Plodowski, G., 1986. The Devono-Carboniferous Tabulipora howsii distributed in Viséan deposits; Penniretepora pluma — transition in the Franco–Belgian Basin with reference to Foraminifera and brachio- upper Viséan, Sulcoretepora minor, Narynella narynica — lower Viséan; pods. In: Bleass, J.M., Streel, M. (Eds.), Late Devonian Events around the Old Red. – Triznotrypa tenuilignata and Klaucena aculeus characterized for Tournaisian, Continent. Annales de la Société géologique de Belgique 109, pp. 19 26. Dunaeva, N.N., 1964a. Novye mshanki otryada Trepostomata iz nizhnego karbona Leioclema tubulosa — lower Tournaisian) (Table 5) Donetskogo bassejna (New bryozoans of the order Trepostomata from the Lower - Two centres of radiation can be identified: «Eastern» and «Western». Carboniferous of Donets Basin). Paleontologicheskij Zh. 2, 39–44 (in Russian). Dunaeva, N.N., 1964b. K faune nizhnekamennougol'nykh trepostomat donetzkogo basseina (The Early Carboniferous trepostome fauna from the Donetsk Basin). Acknowledgements Trudy Instituta Geologii Akademii Nauk Ukrainskoi SSR, seriya stratigraphiya i paleontologiya 48 (2), pp. 104–141, (in Russian). Dunaeva, N.N., 1969. Znachenie mshanok dlya stratigraphii karbona Donetskogo Zoya Tolokonnikova thanks the Deutscher Akademischer Austauschdienst baseina (Importance of Bryozoa for the Carboniferous stratigraphy of the Do- – (DAAD), grant PKZ A/13/00100, for the financial support of fieldtripintheRhe- netsk Basin). Geologicheskiy Zh. 29, 71 80 (in Russian). Ernst, A., 1998. Two unusual trepostome bryozoans from the Lower Carboniferous of the nishMassifin2013.ThestudyofMississippian bryozoans of Russia was con- Karatau (south Kazakhstan). Paläontol. Z. 72 (1/2), 89–97. ducted with financial support from the Paleontological Society (Sepkoski Ernst, A., 2005. Lower Carboniferous Bryozoa from some localities in Sauerland, Grants 2008, 2010, and 2012) (Grants RUG1-1648-XX-06, RUG0-33032-XX- Germany. In: Moyano, G.I., Cancino, J.M., Wyse Jackson, P.N. (Eds.), Bryozoan Studies 2004 — Proceedings of the 13th International Bryozoology Association 10, RUG1-33062-XX-12). Part of this work was performed according to the Conference. Balkema Publishing House, Concepcion, Chile, pp. 49–62. Russian Government Programme of Competitive Growth of Kazan Federal Uni- Ernst, A., 2013. Diversity dynamics and evolutionary patterns of the Palaeozoic versity. Andrej Ernst thanks the Deutsche Forschungsgemeinschaft (DFG) for fi- stenolaemate Bryozoa. Habilitationschrift (unpublished), Christian-Albrechts- Universität zu Kiel. 435p. nancial support (project ER 278/6.1). Reviewers are thanked for the helpful Ernst, A., Rodriguez, S., 2013. Stenolaemate bryozoan fauna from the Missis- comments that improved this paper. This study is a contribution to the sippian of Guadiato Area, southwestern Spain. Span. J. Palaeontol. 28 IGCP 596 “Mid-Palaeozoic climate and biodiversity”. (2), 173–192. Ernst, A., Wyse Jackson, P.N., Aretz, M., 2014. Bryozoan fauna from the Mississippian (Viséan) of Roque Redonde (Montagne Noire, southern France). Geodiversitas References (submitted for publication). Gilmour, E.H., McColloch, M.E., 1995. Fenestrida and Rhabdomesida (Bryozoa) of Ariunchimeg, Ya, 2010. Paleozoiskie mschanki Mongolii (Palaeozoic bryozoans of the Otter Formation (Visean), Central Montana. J. Paleontol. 69, 813–830. Mongolia). (Thesis of doctoral dissertation) , (Moscow, 54 pp. (in Russian)). Gilmour, E.H., Morozova, I.P., 1999. Biogeography of Late Permian bryozoans. Paleontol. J. Bancroft, A.J., 1987. Biostratigraphical potential of Carboniferous Bryozoa. Cour. 33, 36–51. Forschungsinstitut Senckenberg 98, 193–197. Gorjunova, R.V., 1985. Morfologiya, sistema i filogeniya mshanok (otryad Rhabdomesida) Bigey, F.P., 1985. Biogeography of Devonian Bryozoa. In: Nielsen, C., Larwood, G.P. (Eds.), (Morphology, system and phylogeny of Bryozoa (Order Rhabdomesida)). Tr. Bryozoa: Ordovician to Recent. Olsen and Olsen, Fredensborg, pp. 9–23. Paleontologicheskogo Inst. Akad. Nauk. SSSR 208, 1–152 (in Russian). Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211 207

Table 4 Stratigraphical distribution of common bryozoan species by regions in the Viséan. + − presence.

Stage Substage No. Species Number areas Kuznetsk basin Kazakhstan Transbaikalia Mongolia Iran China

Visean Undivided 1 Stenodiscus tumida 3 2 Fenestella alexinensis 2 + 3 Nematopora hibernica 2 4 Goniocladia cellulifera 3 5 Stenophragmidium incrustans 3 6 Streblotrypa pectinata 3 7 Rhabdomeson progracile 5 8 Rhabdomeson regulare 3 9 Pseudonematopora planatus 4 10 Baculopora megastoma 3 11 Dyploporaria tenella 3 12 Thamniscus colei 3 13 Tabulipora howsii 4 14 Penniretepora ﬂexicarinata 3 15 Tabulipora urii 3 + 16 Fenestella sokolskayae 2 17 Fenestella turkestanica 2+ 18 Fenestella daubabaensis 2+ 19 Paraseptopora talasskyensis 3+ 20 Rectifenestella kalbensis 2++ 21 Polyporella kaindiensis 2++ 22 Acanthocladia antiqua 2++ 23 Polypora maccoyaniformis 2++ 24 Nikiforopora intermedia 2+ + 25 Dychotrypa djaltyrensis 2+ 26 Spinofenestella major 3 27 Tabuliporella nalivkini 2+ 28 Clausotrypa ramosa 3 29 Fistulipora proliﬁca 2+ 30 Polypora cesteriensis 2+ + 31 Diploporaria marginalis 2 32 Fenestella angustata 2 Upper 33 Sulcoretepora mergensis 2++ 34 Dyscritella mergensis 2++ 35 Spinofenestella galinae 2++ 36 Fabifenestella fabalis 2++ 37 Penniretepora tschironensis 2++ 38 Lyrocladia mariae 2++ 39 Lyrocladia tschironensis 2++ 40 Nikiforovella vachromeevi 2++ 41 Lanopora mongolica 2++ 42 Rectifenestella mergensis 2++ 43 Rectifenestella invulgata 2++ 44 Polypora narymensis 2++ 45 Penniretepora pluma 3 46 Ptilofenestella carrickensis 2 Lower 47 Sulcoretepora minor 3+++ 48 Fenestella mirabilis 2+ 49 Fenestella confusa 2+ 50 Fenestella terectiensis 2+ 51 Polyporella actashensis 2+ 52 Admiratella prima 2+ 53 Rectifenestella cecikensis 2++ 54 Narynella narynica 3+ 55 Hemitrypa composita 2+ 56 Fistulipora ugamica 2+ 57 Pseudopolypora pseudospininodata 2++

Gorjunova, R.V., 1996. Phylogeniya paleozoiskich mschanok (Phylogeny of the Paleozoic of the genus Leptotrypa of the Russian platform). In: Astrova, G.G., Chudinova, I.I. Bryozoa). Tr. Paleontologicheskogo Inst. Akad. Nauk. SSSR 267, 1–161 (in Russian). (Eds.), New species of Paleozoic Bryozoa and corals. Nauka, Moscow, pp. 47–50 Gorjunova, R.V., Lavrentjeva, V.D., 2007. New bryozoans from the Devonian–Carboniferous (in Russian). boundary beds of Transcaucasia. Paleontol. J. 81 (2), 146–155. Lavrentjeva, V.D., 1974. Late Devonian and Early Tournaisian bryozoans of the central part Gorjunova, R.V., Markov, A.B., Naimark, E.B., 2004. Evolutsiya i biogeographiya of the Russian Platform. Paleontol. J. 8 (2), 157–166. paleozoiskich mshanok: rezultaty kolichestvennogo analiza (Evolution and biogeog- Lavrentjeva, V.D., 1985. Verchnedevonskie mshanki Zakavkaz'ya (Upper Devonian bryo- raphy of the Palaeozoic Bryozoa: results of the numerical analysis). GEOS, Moscow zoans of Transcaucasia). Izvestiya Vysschikh Uchebnykh ZavedeniiGeologiya i (in Russian). Razvedka 8, pp. 12–18, (in Russian). Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001. PAST: paleontological statistics soft- Lavrentjeva, V.D., 1997. Novye dvysloino-simmetrichnye mshanki otryada Trepostomida ware package for education and data analysis. Palaeontol. Electron. 4 (1) (9 pp. (New bifoliate bryozoans of the order Trepostomida). Paleontologicheskij Zh. 4, http://palaeo-electronica.org/2001_1/past/issue1_01.htm). 22–26 (in Russian). Jaccard, P., 1901. Étude comparative de la distribution ﬂorale dans une portion des Alpes Li, Q., Shen, S.-Z., 2014. Global paleobiogeography of brachiopods during the Mississippian — et des Jura. Bull. Soc. Vaudoise Sci. Nat. 37, 547–579. response to the global tectonic reconﬁguration, ocean circulation, and climate changes. Kammer, T.W., Ausich, W.I., 2006. The “Age of Crinoids”: a Mississippian biodiversity Gondwana Res. (in press). spike coincident with widespread carbonate ramps. Palaios 21, 238–248. Lu, Lin-Huang, 1999. Famennian–Tournaisian bryozoans of the Aergati Mt., NW Xinjiang. Lavrentjeva, V.D., 1970. Novye pozdnedevonskie i rannekamennoygol'nye mshanki roda Palaeozoic Fossils of northern Xinjiang, China 37–47. Nanjing Institute of Geology and Leptotrypa Russkoi platformy (New Late Devonian and Early Carboniferous bryozoans Palaeontology, Academia Sinica, Nanjing, pp. 142–186 (in Chinese). 208 Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

Table 4 Stratigraphical distribution of common bryozoan species by regions in the Viséan.

Germany Ireland Britain Belgium Poland Spain Turkmenistan Russian Platform Urals Uzbekistan Kirgizia France

+++ + + + +++ +++ +++ ++++ + +++ ++ + + +++ ++ + ++ + +++ + ++ + ++ ++ + + ++

+ ++ + + ++ + +

++ ++

++ + + +

+ + + + +

++ + +

McCoy, V.E., Anstey, R.L., 2010. Biogeographic associations of Silurian bryozoans genera in Nekhoroshev, V.P., 1925. Geologicheskoe stroenie Priirtyschskich gor vblizi ustjya reki North America, Baltica and Siberia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 297, Bychtarmu v Rudnom Altai (Geological structure of Irthysh mountains around the 420–427. outfall river Buchtarma in Rudnyi Altai). Izv. Geologicheskogo Komiteta 43 (6), McKinney, F.K., 1993. Carboniferous biogeography of the bryozoan Archimedes in North 767–786 (in Russian). America. Hist. Biol. 7, 71–90. Nekhoroshev, V.P., 1932. Die Bryozoen des deutschen Unterkarbons. Abhandlungen der Michno, N.M., Balakin, G.V., 1975. Foraminifery i mshanki nizhnego karbona Chatkal'skich Preussischen Geologischen Landesanstalt N. S. 141, pp. 1–74. gor (Foraminifers and bryozoans from lower Carboniferous of Chatkal’skich Moun- Nekhoroshev, V.P., 1953. Nizhnekamennougol'nye mshanki Kazakhstana (Lower Carbon- tains). FAN Uzkekskoi SSR, Tashkent (in Russian). iferous Bryozoa of Kazakhstan). Trudy Vsesoyuznogo Nauchno-Issledovatelskogo Morozova, I.P., 1981. Kamennougol'nue mshanki severo-vostoka SSSR (Carboniferous Instituta (VSEGEI). pp. 1–183, (in Russian). Bryozoa of North Eastern USSR). Tr. Paleontologicheskogo Inst. Akad. Nauk. SSSR Nekhoroshev, V.P., 1956. Nizhnekamennoygol'nyi mshanki Altaya i Sibiri (Lower Carbon- 188, 1–119 (in Russian). iferous Bryozoa of Altai and Siberia). VSEGEI, Leningrad (in Russian). Morozova, I.P., Weiss, O.B., Racki, G., 2006. Novye devonskie I kamennougol'nue mshanki Nikiforova, A.I., 1927. Materialy k poznaniu nizhne-kamennougolnych mshanok Sventokschirskich gor (Pol'sha) (New Devonian and Carboniferous bryozoans from Donetskogo bassejna [Materials for knowledge Lower Carboniferous bryozoans of Sventokschirsky Mountains (Poland)). Paleontologicheskij Zh. 5, 58–67 (in Russian). Donetsk Basin]. Izvestiya Geologicheskogo Komiteta 46 (3), 245–268 (in Russian). Naimark, E.B., Markov, A.V., Gorjunova, R.V., 1999. Biogegraphiya paleozoiskich mshanok: Nikiforova, A.I., 1933. Kamennougolnye otlozheniya Sredrei Azii. Materialy k poznaniu kolichestvennui analiz (Biogeography of Paleozoic bryozoans: quantitative analysis). mshanok Turkestana (Carboniferous deposits of the Middle Asia. Materials for Paleontol. Zh. 3, 50–59 (in Russian). knowledge bryozoans from Turkestan). Tr. VGRO 207, 1–77 (in Russian). Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211 209

Fig. 4. Palaeobiogeographical afﬁnities of bryozoan assemblages of Eurasia in the Viséan revealed by: 4a — Cluster analysis, 4b — Principal coordinates analysis. All analyses performed with PAST (Hammer et al., 2001). Abbreviations as for Fig. 3.

Nikiforova, A.I., 1948. Nizhne-kamennougolnye mshanki Karatau (Lower Carboniferous Simakov, K.V., 1993. Biochronological aspects of the Devonian–Carboniferous crisis in the Bryozoa of Karatau). AN KazSSR, Alma-ata (in Russian). regions of the former USSR. Palaeogeogr. Palaeoclimatol. Palaeoecol. 104, 127–137. Nikiforova, A.I., 1950. Nizhne-kamennougolnye mshanki zapadnoi okonechnosti chrebta Snyder, E.M., 1991a. Revised taxonomic procedures and paleoecological implications for Talasskogo Alatau (Tyan'-Schan') (Lower Carboniferous Bryozoa of the western end some North American Mississippian Fenestellidae and Polyporidae. Palaeontogr. of the Talas Ala-Tau Range (Tien Shan)). Tr. Inst. Geol. AN Uzbekskoi SSR 5, 90–157 Am. 57, 1–275. (in Russian). Snyder, E.M., 1991b. Revised taxonomic approach to acanthocladiid Bryozoa. In: Bigey, F.P. Plamenskaya, A.G., 1964. O nachodkach predstavitelei rodov Chainodictyon i (Ed.), Bryozoaires actuels et fossil: Bryozoa living and fossil. Bulletin de la Société des Kazarchimedes v nizhnem karbone Kazakhstana (About discovery of representa- Sciences Naturelles de l'Quest de la France, Mèmore HS 1, pp. 431–445. tives of genera Chainodictyon and Kazarchimedes in the Lower Carboniferous of Stampfli, G.M., Hochard, C., Verard, C., Wilhem, C., von Raumer, J., 2013. The formation of Kazakhstan). Paleontologicheskij Zh. 2, 45–48 (in Russian). Pangea. Tectonophysics 593, 1–19. Plamenskaya, A.G., 1983. O novom rode mshanok iz niznego karbona Kazakhstana (About Sterren, A.F., Cisterna, G.A., 2010. Bivalves and brachiopods in the Carboniferous — Early new bryozoan genus from the Lower Carboniferous of Kazakhstan). Paleontologicheskij Permian of Argentine Precordillera: diversification and faunal turnover in Southwest- Zh. 1, 54–59 (in Russian). ern Gondwana. Geol. Acta 8, 501–517. Popeko, L.I., 2000. Karbon Mongolo-Ochotskogo orogennogo poyasa (Carboniferous of Sun, Y., Baliński, A., 2011. 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Palaeontology 24, stan Documents des Laboratories de Geologie de la Faculte des Sciences de 313–341. Lyon. 47, 1–52. Ross, J.R.P., 1982. Biogeography of Ordovician ectoproct (bryozoan) faunas. In: Larwood, Tolokonnikova, Z., 2011. The first data on bryozoans from Devonian–Carbonifer- G.P., Nielsen, C. (Eds.), Recent and Fossil Bryozoa. Olsen and Olsen, Fredensborg, ous boundary beds of the Southern Urals (Zigan and Sikaza sections). Biostra- pp. 213–220. tigraphy, Paleogeography and Events in Devonian and Lower Carboniferous Ross, C.A., Ross, J.R.P., 1982. Biogeographical influences on Late Palaeozoic faunal distribu- (SDS/IGCP 596 Joint Field Meeting). Publishing House of SB RAS, Novosibirsk, tions. In: Larwood, G.P., Nielsen, C. (Eds.), Recent and Fossil Bryozoa. Olsen and Olsen, pp. 156–158. Fredensborg, pp. 199–212. Tolokonnikova, Z., 2012. Early Carboniferous bryozoans from Western Siberia, Russia. In: Ross, C.A., Ross, J.R.P., 1985. Carboniferous and Early Permian biogeography. 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Main features of bryozoans in Central Kazakhstan at the boundary University of Texas, Arlington (http://www.scotese.com,PALEOMAPwebsite). between the Devonian and the Carboniferous. Paleontol. J. 9, 323–339. Shen, S.-Z., Zhang, H., Li, W.-Z., Mu, L., Xia, J.-F., 2006. Brachiopod diversity patterns from Tuckey, M.E., 1990a. Biogeography of Ordovician bryozoans. Palaeogeogr. Palaeoclimatol. Carboniferous to Triassic in South China. Geol. J. 41, 345–361. Palaeoecol. 77, 91–126. 210 Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211

Table 5 Some typical bryozoans species from Mississippian of Eurasia. Fig. A–C. Nikiforopora intermedia (Nikiforova, 1950). Palaeontology Department of the National Iranian Oil Company Exploration Directorate: A — tangential section, MZY 3568; B — longitudinal section, MZY 3566b; C — longitudinal section, MZY 3566b. Iran, Howz-e-Dorah cut; Carboniferous, Mississippian, Viséan, lower Viséan, Shishtu 2 Member of the Shishtu Formation. Fig. D, E. Triznotrypa tenuilignata (Trizna, 1958). SibGIU: D — longitudinal section, No. 12/3; E — tangential section, No. 12/3. Russia, Kurgan region, borehole Kurgan–Uspenskaya 1, depth 1860.0–1865.8 m; Carboniferous, Mississippian, Tournaisian, upper Tournaisian. Fig. F, G. Spinofenestella major (Nikiforova, 1927). Senckenberg Museum: F — tangential section, SMF 21.899; G — tangential section, SMF 21.899. France, Roque Redonde; Carboniferous, Mississippian, Viséan, upper Viséan, Roque Redonde Formation. Fig. H–J. Rhabdomeson progracile (Wyse Jackson and Bancroft, 1995). Senckenberg Museum: H — tangential section, SMF 21.872; I — longitudinal section, SMF 21.882; and J — branch transverse section, SMF 21.875. France, Roque Redonde; Carboniferous, Mississippian, Viséan, upper Viséan, Roque Redonde Formation. Z. Tolokonnikova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 414 (2014) 200–211 211

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