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12-15-2018 Precaspian Isthmus Emergence Triggered the Early Sakmarian Glaciation (Paleontologic, Sedimentologic and Geochemical Proxies) Vladimir I. Davydov Boise State University

Publication Information Davydov, Vladimir I. (2018). "Precaspian Isthmus Emergence Triggered the Early Sakmarian Glaciation (Paleontologic, Sedimentologic and Geochemical Proxies)". Palaeogeography, Palaeoclimatology, Palaeoecology, 511, 403-418. http://dx.doi.org/ 10.1016/j.palaeo.2018.09.007

This is an author-produced, peer-reviewed version of this article. © 2018, Elsevier. Licensed under the Creative Commons Attribution- NonCommercial-NoDerivatives 4.0 license. The final, definitive version of this document can be found online at Palaeogeography, Palaeoclimatology, Palaeoecology, doi: 10.1016/j.palaeo.2018.09.007 Precise timing of the Precaspian- Tethys Seaway closure (Precaspian Isthmus) is established

The emergence of the Isthmus occurs at the Asselian-Sakmarian transition

The biotic and sedimentological evidences support the emergence of the Isthmus

The event changed the global oceanic circulation and is the major driver of the glaciation 295-290 Ma ? 310-295 Ma Uralian Foredeep ? 11 12 11 10 10 8 8 5 12 7 9 6-7 5 9 6 4 4 2

3 1 2 Center of originations

1 Center of originations

WWBF Realms: Tethyan Boreal North American Direction of oceanic currents 1 Precaspian Isthmus emergence triggered the Early Sakmarian glaciation

2 (paleontologic, sedimentologic and geochemical proxies)

3 Vladimir I. Davydov a,b,с,*

4 a Permian Research Institute, Boise State University, 1910 University Drive, Boise, ID, 83725,

5 USA

6 b Kazan Federal University, Kremlevskaya St., 4/5, Kazan’, Tatarstan Republic, Russia

7 c North-East Interdisciplinary Scientific Research Institute n. a. N.A. Shilo Far East Branch of the

8 Russian Academy of Sciences, Magadan.

9 * Corresponding author: [email protected]

10ABSTRACT

11 The sub-meridional seaway that connected Paleo-Arctic and Paleo-Tethys basins was one of the

12 most important geographical attributes of the Late Paleozoic Pangea landscape,

13 paleogeography and paleoclimate. Existing models about the timing of the disconnection of

14 the Paleo-Arctic and the Paleo-Tethyan oceans is very controversial and poorly documented.

15 Warm-water benthic foraminifera (WWBF) were utilized to establish the precise timing of the

16 closure of the Urals-Precaspian-Paleo-Tethys Seaway (UPTS) during Cisuralian time. The WWBF

17 of Paleo-Tethys and those of the Ural—Precaspian Basins during the Gzhelian-Asselian, display a

18 considerably high level of similarity. Beginning from the Sakmarian, the faunas of these two

19 regions became dissimilar, suggesting a break in the connection between the Paleo-Tethys and

20 Ural-Precaspian Basins. The sedimentological evidence (olistostromes and seismites) of the final

21 collision of the Eastern Ural, Kazakhstania, Scythian-Turan plates with the southeastern part of

22 the Russian Platform during Late Paleozoic also support the emergence of the Precaspian

23 Isthmus at the Asselian-Sakmarian transition. The oceanic currents in the Precaspian and the

24 Southern Ural Basins before the Sakmarian were directed northward and later changed to the 25 south. These biotic and physical changes are consistent with the proposed timing of the cutoff

26 of the UPTS. The biotic and sedimentologic features clearly suggest the UPTS closure and the

27 origination of Precaspian Isthmus during the Asselian-Sakmarian transition. The abrupt changes

28 in the oceanic circulation triggered changes in atmospheric CO2, atmospheric circulation and,

29 possibly, albedo feedback. The emergence of the Precaspian Isthmus induced an increase in the

30 poleward salt and heat transport towards mid- to lower latitude Gondwana and Cathasia

31 margins. The warm water currents and moisture along the margins of Gondwana caused a rapid

32 increase in the precipitation necessary to build significant ice sheets during the early-middle

33 Sakmarian.

34 Keywords: Oceanic gateway; Isthmus; Late Paleozoic glaciation; benthic foraminifera;

35 continents configuration.

36 1. Introduction

37 The Late Paleozoic glaciation is a very intriguing and controversial matter and is the penultimate

38 icehouse-greenhouse transition on Earth (Crowell, 1999; Montanez and Poulsen, 2013; Smith and

39 Read, 2000). Our understanding of the processes associated with the glaciation, and particularly

40 the factors that caused the icehouse to greenhouse transition, may help us better understand the

41 changes to recent climate perturbations. The level of atmospheric CO 2 is considered the major

42 factor that drives climate change along with the other less important, such as tectonics,

43 continents configuration, variations of the orbital and spin axis of the Earth and other

44 extraterrestrial events (Montanez and Poulsen, 2013; Smith and Read, 2000). However, the

45 factors behind CO2 fluctuations in the past are unclear. The role of paleogeographic configuration

46 and solar irradiation is considered of secondary importance (Lowry et al., 2014).

47 The sub-meridional seaway that connected Paleo-Arctic and Paleo-Tethys basins was

48 one of the most important geographical attributes of the Late Paleozoic Pangea landscape,

49 paleogeography and paleoclimate (Scotese, 2015). The seaway which connected the shelves

50 along the East-European Craton, the Ural and the Paleo-Tethys was the major oceanic gateway

51 between the oceans in the Paleo-Tethys and Paleo-Arctic (Fig. 1). The final collision of

52 Kazakhstania and Siberian continents with the sutured Laurentia and Gondwana is usually

53 considered to have caused the closure of the Uralian foredeep and UPTS sometime in latest

54 Cisuralian time (Puchkov, 2010). Most tectonic models of the development of the Ural and the

55 surrounding areas during Paleozoic-Mesozoic time imply the existence of the UPTS until the

56 Kungurian because of the early Permian marine sedimentation in the Precaspian and the

57 Southern Ural. According to these models, the end of the marine sedimentation and the

58 accumulation of thick sabkha evaporites in the Precaspian and along the Ural during the

59 Kungurian denote the complete closure of the UPTS (Brown et al., 1997; Cocks and Torsvik,

60 2007; Golonka, 2007; Nikishin et al., 1996; Puchkov, 1997, 2009, 2010; Snyder et al., 1994;

61 Ziegler, 1989; Zonenshain et al., 1990). Nevertheless, some paleogeographic models suggest

62 the existence of the seaway between the Paleo-Arctic and the Paleo -Tethys until the Triassic

63 (Blakey, 2013; Chumakov and Zharkov, 2002; Domeier and Torsvik, 2014 Kaz'min and and

64 Natapov, 1998; Scotese, 2015). Other models suggest that the closing of the Uralian—Paleo-

65 Tethys connection happened in the Moscovian (Lawver et al., 2011) or sometime within the

66 Bashkirian - Kasimovian (Cavazza et al., 2004; Stampfli et al., 2013). Golonka (2011) proposed

67 that the seaway was closed in the Asselian-Artinskian and reopened in the Guadalupian. A

68 somewhat exotic idea by Sengor and Atayaman (2009) proposed that the Paleo-Arctic—Tethys

69 was connected through the hypothetical Carapelit rift during latest Kungurian-Wuchiapingian

70 time.

71 Existing models about the timing of the disconnection of the Paleo-Arctic and the Paleo-

72 Tethyan oceans is very controversial and poorly documented and one of the goals of this study

73 was to establish the precise timing of the event. The exceptionally accurate, quantitative

74 biostratigraphic and radioisotopic calibration of the Pennsylvanian–early Permian global time

75 scale, developed in the type sections in the Southern Ural was employed to establish the

76 precise timing of the geological events in the Late Paleozoic, including those in the Ural-

77 Precaspian regions ( Davydov et al., 2012; Davydov and Cozar, 2018, in press; Schmitz and

78 Davydov, 2012). The tropical-subtropical paleogeographic distribution and the well-known

79 sensitivity of WWBF to paleoenvironments, coupled with their application in development of

80 high-resolution spatial and temporal framework, provides the basis for the study presented

81 here. The taxonomic changes and the evolutionary divergence of the foraminifera in the Uralian

82 and West Tethyan oceans through the Pennsylvanian and early Permian time definitively

83 indicate the emergence of the Precaspian Isthmus and the closure of the connection between

84 the Uralian and Paleo-Tethys oceans on both the northern and mid-east sides of Pangea (Fig. 1).

85 The mid-late Artinskian closure of the connection between the Uralian--Paleo-Tethyan

86 oceans utilizing fusulinids data has been proposed previously (Chuvashov, 1998; Leven, 2004;

87 Ross, 1967b;). Shi and Waterhouse (2010) interpreted this as a significant event that enhanced

88 the cooling of the paleo-Arctic ocean which had already been occurring due to Pangaea's

89 northward drift throughout the Permian. However, this closure was considered insignificant in

90 paleoclimatic models because of the lack of the temporal connection with any known climatic

91 event ( Lowry et al., 2014; Montanez and Poulsen, 2013). In this paper we propose and discuss

92 the causal link between the development of the Precaspian Isthmus in-between the Uralian and

93 West Tethyan basins, the profound biotic transformations in the oceans, sea-level changes, the

94 decline of the atmospheric CO2 concentration, and the expansion of the Gondwanan ice sheet

95 during the early-middle Sakmarian.

96 2. Data

97 In this study the foraminiferal data was compiled from 252 literature sources (see supplemental

98 references list) covering the regions along the Ural ocean and the northern margins of the

99 Paleo-Tethys ocean (Fig. 1). The data from the author’s foraminifera collections was also used.

100 Taxonomy was carefully considered and publications with paleontological plates were

101 evaluated. Some publications without plates were also included in the database where the

102 taxonomy appeared reliable. The main goal of the taxonomic evaluation was to develop the

103 database with internally consistent taxonomy for each analyzed region and chronostratigraphic

104 time slice.

105 The data at species level and at generic level was analyzed separately. Foraminifera are

106 very sensitive to environmental changes, particularly at species level. The cutoff of the Ural-

107 Tethys passage would undoubtedly have been expressed in the dramatic changes of the

108 environments and therefore WWBF taxonomy (Davydov and Cozar, 2018, in press). At genus

109 level, those changes are not that apparent as many genera existed at that time and were

110 evolving independently in completely isolated basins for a long time (Collins et al., 1996;

111 Davydov, 2014; Mamet, 1977; Ross, 1995;).

112 Although some of the paleogeographic models suggest a very early UPTS closure in

113 Early-Middle Pennsylvanian time (Lawver et al., 2011; Stampfli et al., 2013), we started to

114 compile the data from the late Gzhelian-Asselian to the Kungurian, as the Bashkirian-Gzhelian

115 fusulinids within the UPTS regions are clearly very similar (Davydov and Cozar, 2018, in press).

116 Most of the existing models propose the UPTS closure during the Kungurian. Some

117 paleogeographic models show the existence of a permanent UPTS during the entire Permian

118 (Blakey, 2013; Domeier and Torsvik, 2014; Golonka, 2011; Scotese, 2015;). These regions during

119 Cisuralian were located within the tropics—subtropics (Blakey, 2013; Davydov, 2014; Domeier

120 and Torsvik, 2014; Golonka, 2011; Scotese, 2015). However, no typical warm-water

121 foraminifera such as fusulinids have ever been found in Kungurian and Roadian seas in the

122 Precaspian, in the Southern Ural, and in the Russian Platform ( Filimonova et al., 2015; Gorsky

123 and Kalmykova, 1986; Sukhov, 2003). The youngest WWBF in these regions are known only

124 from the lower Kungurian in the north-central Preural, where they are rare and of very low

125 diversity (Zolotova and Baryshnikov, 1978).

126 The WWBF data have been compiled along the Ural-Tethyan passage (Fig. 1) which

127 includes the Carnic Alps (CA) (1), Taurids (2), Donets Basin (3), Darvaz (4), South Tian-Shan’ and

128 the northern margin of Tarim microcontinent (5), northern and eastern Precaspian (6-7), the

129 Southern Ural (8), Russian Platform (9), the Central Ural (10) and the Timan-Pechora Basin (11).

130 3. Methods

131 The Late Paleozoic foraminifera FODs (first occurrence datum) and LODs (last occurrence

132 datum) were calibrated using the exceptionally accurate, quantitative biostratigraphic and

133 radioisotopic scale that has been developed in the Ural and surrounding regions (Schmitz and

134 Davydov, 2012). Therefore, the accurate timing of the taxonomic changes associated with the

135 acme of the Precaspian Isthmus closure can be established. All the regions involved in the

136 analyses, occur within the paleo-tropics sub-tropics and therefore the latitudinal differences

137 played no role or minor role in controlling of the distribution of foraminifera in the studied

138 regions. The highest plausible resolution to resolve the time calibration of the foraminiferal

139 events has been employed in this study. The author is confident that the position of the stages

140 and zonal boundaries are constrained with a 0.1-0.2 Myr resolution. The number and precision

141 of the radiometric ages, the number of the bioevents in the composite section (CONOP9), and a

142 direct link to an eccentricity-tuned cyclostratigraphy result in a ca. 0.1–Myr resolution for the

143 chronostratigraphic model through most of this time period (Schmitz and Davydov, 2012).

144 The foraminiferal taxonomic data was analyzed using the PAST v.3.0 software (Hammer

145 et al., 2001). The binary (i.e., presence/absence) matrices for each region include two types: 1)

146 species only; 2) genera only. Probabilistic Raup-Crick similarity/dissimilarity indices were used

147 to calculate the matrices (Hammer and Harper, 2006). More details on the advantages and

148 disadvantages of these different types of the indices applied to WWBF can be found in an

149 earlier published paper (Davydov and Cozar, 2018, in press).

150 4. Results

151 Results of the Raup and Crick probabilistic similarity index (PSI) at species level are given in

152 Table 1, and the data at genus level can be found in supplementary Table 2.

153 The PSI is sensitive to the lack of taxa in any of compared regions and thus, taxa that are

154 geographically widespread do not have a disproportionate effect on the measurement of

155 similarity. The PSI provides the opportunity to measure statistically significant similarity and

156 dissimilarity at the 95% confidence level and is less affected by sampling bias.

157 Two clusters are recognized among WWBF at both species and genus levels during the

158 early-middle Asselian (Figs. 2-4, Tables 1-2). The Paleo-Arctic cluster (Boreal Realm) includes the

159 Russian Platform (9), the central and northern Ural (10) and Timan-Pechora region (11). The

160 Tethyan cluster is comprised of the Carnic Alps (1), Taurids (2), Donets Basin (3), Darvaz (4),

161 South Tian-Shan (5), Precaspian (6-7) and the Southern Ural (8). The Donets Basin was involved

162 in the analyses only for early-middle and late Asselian time-slices. By the early Sakmarian, the

163 Precaspian and the Southern Ural regions became part of the Paleo-Arctic cluster.

164 In the early-middle Asselian the PSI of Paleo-Arctic and Tethyan clusters at species level

165 is 0.2 and increases up to 0.45-0.48 in the late Asselian (Fig. 5). Beginning with the early

166 Sakmarian, the PSI between the Boreal and Tethyan clusters at species level is zero, i.e. the

167 WWBF of these clusters are dissimilar in the post-Asselian time (Fig. 5). The PSI at genus level

168 shows similar tendencies, except the absolute values of the PSI are significantly higher. During

169 early-middle Asselian the PSI at genus level is 0.48, during the late Asselian it increases to 0.7-

170 0.72 (Table 2). Starting in the early Sakmarian the PSI drops drastically to 0.25 and after that,

171 the PSI progressively goes down from 0.25 in the early Sakmarian to 0.1 towards the end of the

172 Artinskian (Fig. 5). Some similarity between the two clusters at genus level can be explained by

173 the fact that the genera ranges are much more prolonged than the ranges of species.

174 Beginning in the early Sakmarian, the Precaspian and the Southern Ural regions became

175 a part of the Paleo-Arctic cluster, the PSI of the latter and the rest of the Paleo-Arctic regions

176 reaching 1.0 (Fig. 5, table1-2). The PSI value remains the same during the entire Sakmarian-

177 Artinskian time.

178 5. Discussion

179 5.1. Paleo-Tethys—Paleo-Arctic Seaway closure

180 5.1.1 Dynamics of taxonomic similarity changes in Paleo-Tethyan and Paleo-Arctic through the

181 Cisuralian time.

182 Four major realms are recognized in the Carboniferous-Permian within the distribution pattern

183 of the Late Paleozoic foraminifera (Ross, 1995) (Fig. 1). The largest realm is the Tethyan, which

184 extended from the tropics in the China blocks to the tropics of the eastern margin of Pangea

185 and the northern margin of Gondwana. The second realm, Boreal or Ural-Franklinian, embraces

186 the subtropics in the northern margins of Pangea. The third realm, commonly called ‘North

187 American’ or ‘Midcontinent-Andean’, is positioned in the tropics and subtropics along the

188 western margin of Pangea. The fourth realm, named ‘Panthalassan’ or ‘Sonomia’, is not well

189 defined. It includes a series of terranes and sea-mounts within Paleo-Pacific, i.e. parts of Japan,

190 the Russian Far East, Cache-Creek, British Columbia, Koryakia, New Zealand, and S. Peru (Fig. 1).

191 The main purpose of this study is the comparison of the Boreal and Tethyan realms. The

192 taxonomic composition of these realms was quite similar during most of the Late Paleozoic, due

193 to the free connection of Paleo-Arctic and Paleo-Tethys oceans (Fig. 6). Although, the WWBF of

194 Russian Platform, the Ural and Timan Pechora formed a separate cluster (Figs. 2-4), they reveal

195 considerable similarity with the WWBF of the Tethyan realm regions during Asselian time,

196 especially at genus level (Figs. 3,5). At that time the WWBF in the Donets Basin, the Precaspian

197 and the Southern Ural belonged to the faunas of Tethyan affinity. The taxonomic diversity of

198 the WWBF within the Gzhelian-Asselian transition reached the maximum recorded for the

199 entire Late Paleozoic and remained very high during the Asselian, except for a small decrease in

200 the diversity in the middle Asselian (Fig. 7) (Davydov, 2014; Groves and Wang, 2009).

201 The major event that drastically changed the similarity among the analyzed regions over

202 an interval of less than 0.5 Myr occurred within the Asselian-Sakmarian transition (Fig. 7). These

203 changes were reported earlier in the Russian literature and some Russian scientists (Bensh,

204 1962; Rauser-Chernousova, 1965) considered the position of the Carboniferous-Permian

205 boundary at the base of the Sakmarian because of these severe changes. Marine

206 sedimentation ceased in the Donets Basin at the end of the Asselian.

207 During the Sakmarian and through the end of the Artinskian the species and genera of

208 the WWBF of the Precaspian and the Southern Ural had a high degree of similarity with those of

209 the Central Ural, Russian Platform and Timan Pechora (Fig. 5). At the same time, the similarity

210 between the WWBF of the Tethyan and Boreal realms dropped twice at genus level and almost

211 to zero at species level (Fig. 5). At the Asselian-Sakmarian transition the global diversity of

212 WWBF at both species and genus levels dropped threefold (Davydov, 2014; Groves and Wang,

213 2009). During the Sakmarian and Artinskian, the similarity between the Tethyan and Boreal

214 WWBF at species level remained close to zero. At genus level, the similarity index had been

215 progressively falling from 0.25 to 0.1 through Sakmarian-Artinskian boundary, and towards the

216 end of the Artinskian dropped below 0.1 (Fig. 5).

217 It is obvious from the discussion above that the distribution of the WWBF throughout

218 the Asselian within the Paleo-Artic and the Ural-Precaspian shelves was not limited by any

219 physical or climatic factors. Starting in the Sakmarian, the WWBF of the Paleo-Tethys shelves

220 were isolated from those of the Paleo-Artic and the faunas of Paleo-Artic and the Ural-

221 Precaspian, developed as a single unified biome. Such a sudden event in the separation of the

222 Paleo-Arctic and Paleo-Tethyan faunas in the subtropics was evidently caused by the

223 appearance of the physical barrier between the Precaspian Basin, the southernmost region of

224 the Paleo-Arctic, and the Paleo-Tethys regions. The presence of this physical barrier is well

225 expressed in the records of the WWBF from the regions closest to the Precaspian-Southern

226 Ural, such as Darvaz, S. Tian-Shan’ and Taurides (Figs. 1, 6). The data from Turan Platform,

227 which was a part of the Precaspian Basin until the late Asselian, is consistent with this

228 observation (Akramkhodjaev et al., 1981; Uzakov, 1996; Dronov et al., 1997; Pronin et al., 2011;

229 Volozh et al., 2011, 2011).

230 5.2 Evolutionary and ecological proxies of the WWBF of the Uralian-Tethyan passage

231 closure.

232 The UPTS passage was the only way for the exchange of the WWBF between the Paleo-

233 Arctic and Paleo-Tethys during the Late Paleozoic (Fig. 1A). Thus, the timing of the

234 disappearance of the Tethyan taxa in the Precaspian and in the Ural and the shift in the

235 provinciality from the Tethyan affinity to the Boreal would be the signs of the emergence of the

236 Precaspian Isthmus (Fig. 6)

237 The migration of the WWBF between the Paleo-Tethyan and Paleo-Arctic oceans

238 occurred in only one direction, from south to north. This is consistent with the distribution of

239 the marine faunas along the temperature latitudinal gradient from the high-diversity

240 assemblages in the Paleo-Tethys to the low-diversity assemblages in the shelves of higher

241 latitudes (Davydov, 1997; Davydov and Arefifard, 2013; Jablonski et al., 2006). Most of the

242 fusulinid genera and many species, which dispersed into the Paleo-Arctic ocean, originated in

243 the Paleo-Tethys (Davydov, 1986; Davydov et al., 2001; Izotova, 1985; Izotova and Vevel, 1998;

244 Nilsson and Davydov, 1997). The distribution and provincialism of the WWBF were largely

245 controlled by climate along the temperature latitudinal gradient (Davydov, 2014; Rind, 1998).

246 WWBF provide a particularly sensitive index of climate changes within the marginal conditions

247 at the transition from subtropics to temperate mid-latitudes, such as the shelves of the Paleo-

248 Arctic ocean (Davydov and Arefifard, 2013). During the Late Paleozoic there were several waves

249 of change in the biodiversity and provinciality of the WWBF, associated with global climate

250 fluctuations (Fig. 7). During episodes of global warming (climatic optimums), the WWBF for the

251 given latitude and depth reached their maximum diversity and world-wide distribution, with

252 tropical taxa migrating to the higher latitudes. For example, Eofusulina originated in the

253 Vereiyan (the early Moscovian) in the Paleo-Tethys and migrated into the Paleo-Arctic

254 (Greenland, Canadian Arctic) during the Kashirian (the late early Moscovian). The late Gzhelian-

255 early Asselian Likharevites and Asselian Sphaeroschwagerina and Biwaella migrated in the late

256 Asselian as far as Spitsbergen, North Greenland and the Canadian Arctic (Nilsson and Davydov,

257 1992, 1997; Rui Lin et al., 1994); and personal data of the author). By contrast, global cooling

258 led to the disappearance of the foraminiferal fauna starting at higher and moving to lower

259 latitudes, their stepwise extinction, increasing provincialism, and to the preferential survival of

260 euryfacial faunas (Davydov and Arefifard, 2013).

261 During the Moscovian-Gzhelian and the entire Asselian the WWBF in the Precaspian,

262 Donets Basin and the Southern Ural were generally of Tethyan affinity with many common

263 species and genera known from the other Tethyan regions ( Akhmetshina et al., 2013; Davydov,

264 1986, 1992; Izotova, 1985; Nikolaev, 2011; Leven and Scherbovich, 1978; Scherbovich, 1969).

265 The WWBF taxa of the Boreal affinity in the Precaspian, Donets Basin and the Southern Ural

266 were very rare during this time. From the beginning of the Sakmarian, the fauna of the Boreal

267 affinity predominated in the Precaspian and the Southern Ural. Only a small number of species

268 of Tethyan affinity, i.e., few species of Sakmarella, Darvasites and Rugosofusulina, described

269 from Darvaz (Leven and Shcherbovich, 1980), were found only in the earliest Sakmarian in the

270 very southern part of the Southern Ural (Davydov, 1986). The Tethyan Sakmarella, Darvasites

271 and Rugosofusulina were found in only one (the lowermost) horizon of the Sakmarian, in the

272 Southern Ural (Davydov, 1986). From the beginning of Sakmarian the WWBF of the Precaspian

273 and southern Ural were exclusively of Boreal affinity. And again, these sharp changes in

274 biogeography signify the physical separation of the Precaspian and Southern Ural regions from

275 the Tethyan shelves at the Asselian-Sakmarian transition.

276 5.3 Olistostrome and seismite deposits and the oceanic currents within the Late Paleozoic in

277 the Southern Ural.

278 Olistostromes and seismites occur widely in ancient orogenic belts and in exhumed subduction–

279 accretion complexes. In latter type of orogenic belt, the temporal and spatial distribution of

280 different types of olistostromes commonly indicate different tectonic stages over period of 10+

281 Myrs from the early rift–drift to the later subduction, collision, and orogenic exhumation

282 (Ettensohn et al., 2002; Festa et al., 2016). Generally, olistostromes are the fundamental

283 markers of tectonic events and their study is a powerful tool for the basin analysis at various

284 scales (Festa et al., 2016). Hence, the evaluation of distribution of the olistostromes and

285 seismites in the area of the Precaspian Isthmus may help in understanding of the dynamics of

286 the closure. Unfortunately, only one area, i.e., Southern Ural, is available for this evaluation.

287 The scarсe published records of the lithostratigraphy from the subsurface of the Precaspian

288 have prevented the analyses of the olistostromes there. The regions immediately to the south

289 from the Precaspian Isthmus are either not exposed or occur far from the collision area

290 (Figs.1,6,8).

291 An olistostrome is a sedimentary deposit composed of a chaotic mass of heterogeneous

292 material, such as blocks of consolidated and unconsolidated sediments with siliciclastic or

293 carbonate matrix mud, that accumulates as a semifluid body by submarine gravity sliding or

294 slumping of the unconsolidated sediments. Olistostromes are well preserved in paleo-active

295 margins such as Southern Ural foredeep. Causative mechanisms for gravitational instability in

296 collisional and intra-collisional settings are mostly seismic shocks originating from thrust

297 faulting (Festa et al., 2016). The seismite, as opposed to olistostromes, referred only to

298 stratigraphic units containing sedimentary structures produced by seismic shaking and is an in

299 situ breccia (Ettensohn et al., 2002; McCalpin, 2009). Thus, the occurrence of olistostromes and

300 seismites is indicative of the seismic activity and dynamic orogeny that are concomitant to the

301 collision processes in the Southern Ural and could be projected to the Precaspian Basin (Mizens,

302 1997; Puchkov, 2010).

303 The olistostromes are broadly distributed and well known in the Southern Ural, where

304 they have been thoroughly studied and described (Keller, 1949; Khvorova, 1961; Mizens, 1997,

305 2002). Two types of the olistostromes are designated in the Southern Ural. The first one occurs

306 as proximal facies of the poorly sorted mass of consolidated, sharp or poorly rounded boulders

307 and blocks up to 100-150 meters in size, conglomerate-breccia and conglomerates. These clasts

308 consisting of shallow-water limestone touch each other. A small amount of the silty-

309 carbonaceous matrix fills the space between the clasts (Khvorova, 1961; Mizens, 1997). This

310 type of olistostrome lacks fossils in the matrix and, perhaps, belongs to the olistostromal carpet

311 in the classification of Festa et al (2016). More distal olistostromes consist of matrix-supported,

312 unconsolidated carbonate and siliciclastics material with very large to small, sharp to rounded

313 blocks, pebbles, and unconsolidated carbonate or siliciclastic clasts of different sizes. The matrix

314 and clasts are generally of the same composition and both contain abundance of fossils of the

315 same taxa (Khvorova, 1961). This type of olistostrome belongs to the category of precursory

316 olistostromes (Festa et al., 2016).

317 Horizons of both precursory olistostromes and olistostromal carpet are unevenly

318 distributed within the Late Paleozoic succession in the Southern Ural (Fig. 7). The olistostromes

319 first appear there in the upper Moscovian (Khvorova, 1961), which is consistent with the initial

320 stage of rigid (hard) collision of Kazakhstania and the Ural and the initiation of the Uralian

321 orogeny (Puchkov, 2010). The olistostromes are frequent and sometimes thick in the upper

322 Moscovian, Kasimovian and early-middle Gzhelian (Khvorova, 1961; Mizens, 1997). The

323 uppermost horizon of the precursory olistostrome in the Carboniferous is recorded at the

324 beginning of the Orenburgian stage (the late Gzhelian in the International Scale) (Davydov,

325 1986; Khvorova, 1961). This late Gzhelian horizon can be traced in the distance of over 600 km

326 from Aidaralash Creek in Aqtobe region in Kazakhstan to Usolka river in the northern part of the

327 Southern Ural in Bashkortostan (Mizens, 1997). The end of the Gzhelian and early-middle

328 Asselian was quiet, with no olistostromes recorded. The beginning of the late Asselian is again

329 associates with the massive deposition of olistostromes, which were especially abundant in the

330 southernmost exposed part of the Southern Ural (Khvorova, 1961; Mizens, 1997). In

331 Kondurovsky section, which is the type of the Sakmarian Stage, two horizons of specific

332 sediments, named Kurmaya breccia, are described in many publications (Keller, 1949;

333 Khvorova, 1961; Mizens, 2002). These distal facies formed from the micritic limestone matrix

334 with the submerged clasts of micritic limestone (1-5 cm in size) of different color from very dark

335 to light-grey (Fig. 8). This type of rock, perhaps, could be interpreted as seismites, as both the

336 matrix and clasts are of the local origin and of same composition (Ettensohn et al., 2002). The

337 occurrence of tsunami deposits in the Late Paleozoic of Zalair Zone of the Ural has been

338 proposed as well (Keller, 1949; Khvorova, 1961).

339 During the Sakmarian and through the Artinskian and Kungurian, no olistostromes have

340 been recognized in the Southern Ural (Khvorova, 1961; Mizens, 1997). The olistostromes of this

341 age were documented only in the Central and Northern Ural, which is consistent with the

342 observed diachronous shift in the granitic magmatism and the collision area of Kazakhstania

343 and the Ural towards the north (Puchkov, 2010). In the southern periphery of the Precaspian

344 Basin, in the Tengiz-Karaton Dom structure and other micro-platforms, the carbonate and

345 mixed carbonate-siliciclastic succession extended up to the Bashkirian. In the surrounding

346 platforms areas the successions extended up to the Asselian (Abilkhasimov, 2016). In the very

347 south of the Emba zone (Buzachi Peninsula) marine sedimentation ended in the late Asselian

348 (Pronin et al., 1997). Pyroclastic material and volcanic tuffs are recognized and widely

349 distributed in the lower Permian there (Abilkhasimov, 2016). From the beginning of the

350 Sakmarian neither seismic activities nor orogeny occur in the area of the Southern Ural and

351 Emba zone in the southern periphery of the Precaspian Basin (Fig. 6) (Abilkhasimov, 2016).

352 Beginning in the Artinskian in the southern periphery of the Precaspian Basins, fluvial fan

353 deposits are documented (Bakirov et al., 1978; Fedorenko and Miletenko, 2002; Rogova and

354 and Pugacheva, 1983), suggesting the development of the weak orogeny along the border

355 between the southern Precaspian Basin and Scythian-Turan plates (Fig. 6).

356 The pattern of oceanic currents in the Southern Ural and Precaspian Basin may also help

357 clarify when the emergence of the Precaspian Isthmus occurred. The northern direction of the

358 oceanic currents in the Southern Ural has been reported in several publications (Bezhaev, 1968;

359 Keller, 1949; Khvorova, 1961; Mizens, 1997, 2002). During the Late Paleozoic, the clastic

360 sources in the Preuralian foredeep were to the east (in the present day coordinates) from the

361 Uraltau and Magnitogorsk Arc (Mizens, 1997; Puchkov, 2010). This material was delivered into

362 the foredeep by a series of paleo-rivers (Khvorova, 1961). The submarine deltaic fan deposits

363 along the mountains are supposedly oriented perpendicular to the mountains. In the Ural this

364 type of the submarine fan conglomerates in the shallow part are oriented perpendicular to the

365 mountains, i.e. from east to west (in the present day coordinates), but in the deeper part of the

366 foredeep they curve and are oriented towards the north-western and northern direction (in the

367 present day coordinates) (Fig. 6) (Khvorova, 1961; Mizens, 1997). Besides, the data on the

368 orientation of the deep-water currents in the foredeep has been obtained from the analyses of

369 the shape and distribution of the ocean paleocurrents features, such as small-scale cross-

370 bedding and ripples in the Late Paleozoic flysh successions. The cross-bedding and ripples also

371 suggest the northern direction of the oceanic currents (in the present day coordinates) during

372 Moscovian-Asselian time ( Bezhaev, 1968; Keller, 1949; Khvorova, 1961; Mizens, 1997).

373 However, beginning in the Sakmarian time the orientation of the ripples and cross-bedded

374 lamination of the sediments suggest southern direction of the oceanic currents within the

375 Uralian foredeep (Mizens, 1997)

376 5.4 The Precaspian and Scythian-Turan plate paleotectonics

377 According to the majority of publications (Cocks and Torsvik, 2007; Golonka, 2007; Nikishin et

378 al., 1996; Puchkov, 2000, 2010; Ziegler, 1989; Zonenshain et al., 1990), the Paleo-Tethys—

379 Paleo-Arctic active passage was closed in Kungurian. The other model suggests that a shallow-

380 water connection existed throughout the entire Permian (Blakey, 2013; Domeier and Torsvik,

381 2014; Chumakov and Semikhatov, 2004; Fedorenko and Miletenko, 2002; Kaz'min and and

382 Natapov, 1998; Scotese, 2015;). The biota the WWBF in the Paleo-Tethys, and Paleo-Arctic

383 basins and key sedimentological features in the Southern Ural suggest that the Paleo-Tethys-

384 Paleo-Arctic connection through the Uralian foredeep lasted through the Asselian with the

385 cutoff of the connection during the Asselian-Sakmarian transition.

386 The data of the subsurface of the areas along the transitional zone between the

387 southern periphery of Precaspian Basin and Scythian-Turan plate is very poor however some

388 assumptions can be made. Most of the models suggest the connection between the Paleo-

389 Arctic and Paleo-Tethys through the Donets—Tuarkyr and the Usturt rift systems

390 (Abilkhasimov, 2016; Afanasenkov et al., 2008; Fedorenko and Miletenko, 2002; Gavrilov, 2011;

391 Khain and Popkov, 2009; Nikishin et al., 2012;). Marine sedimentation in Donets Basin ended in

392 the latest Asselian and the remnants of the basin were filled with the thick succession of salt

393 and other evaporites (Korenevsky et al., 1968). The evaporites of the Kramatorsk formation lack

394 fossils and tentative age proposals vary from the Sakmarian to Kungurian (Volodin, 1993).

395 In the Tuarkyr and Ustyurt rift systems the Upper Paleozoic rocks have been recovered

396 only from the subsurface. The youngest known marine sediments in these regions are Asselian

397 limestone ( Akramkhodjaev et al., 1979; Akramkhodjaev et al., 1981; Maslov et al., 2016). The

398 Asselian carbonates in the region are overlain by the continental late Permian-Triassic

399 sequences (Akramkhodjaev et al., 1979; Gavrilov, 2011). In the area of the connection of the

400 Turan plate and the southern periphery of Precaspian Basin, in the Caspian Sea, the youngest

401 recorded carbonates are Asselian limestone that are unconformably overlain by Mesozoic

402 siliciclastics (Maslov et al., 2016; Pronin et al., 1997). This succession is quite similar to the one

403 in the Buzachi and South Emba regions immediately to the east of the Caspian Sea (Pronin et

404 al., 2011).

405 The most recent reconstructions of the Precaspian Basin and surrounding areas suggest

406 that a narrow rift system within the Turan plate provided a marine connection between the

407 Precaspian Basin and Paleo-Tethys until Kungurian time (Abilkhasimov, 2016) or even later

408 (Fedorenko and Miletenko, 2002). The rift system along the southern slope of the Karpinsky

409 swell that introduced carbonate sedimentation to the Pre-Caucasia and North Caucasus (Kaz'min et

410 al., 2008), emerged much later in the latest Permian time (Letavin A.I. et al., 1994). Although

411 the Precaspian, Southern Ural and the southern part of Russian Platform were still within the

412 subtropics at that time, only temperate foraminifera were found in the Kungurian and post-

413 Kungurian in these regions ( Filimonova et al., 2015; Gusev et al., 1968; Pronina, 1999; Sukhov,

414 2003). This unambiguously suggests direct connection of these regions with the temperate seas

415 of the Paleo-Arctic during the Guadalupian-Lopingian and the lack of a connection with the

416 Paleo-Tethys in post-Sakmarian time.

417 As in the case with the Alleghanian Isthmus (Davydov and Cozar, 2018, in press), the

418 closure of the Paleo-Tethys—Paleo-Arctic passage happened rather quickly, most probably,

419 during an interval of less than 0.5 Myr (Fig. 5). Some elements of the WWBF of the Tethyan

420 affinity still occurred in the earliest Tastubian in the south of the Southern Urals (Aidaralash,

421 Aktobe province) (Davydov, 1986), but rapidly disappeared almost immediately. The fauna of

422 Boreal affinity dominated in the region from Sakmarian through Kungurian and no elements of

423 Tethyan affinity are known in the Precaspian and the Southern Ural or anywhere in the Paleo-

424 Arctic (see the Results chapter for details). This conclusion in accord with the frequent

425 occurrence of olistoliths and seismites in the Uralian foredeep, which suggests the presence of

426 an active tectonic regime until the end of the Asselian in the Southern Ural and the reduction of

427 tectonic activity in the post-Asselian time (Khvorova, 1961; Mizens, 2002). Carbonate

428 sedimentation ended in the late Asselian in the areas to the south and southeast of the

429 southern periphery of the Precaspian Basin in Turan. The widely developed Artinskian

430 submarine fan deposits along the southern periphery of the Precaspian Basin (Abilkhasimov,

431 2016; Fedorenko and Miletenko, 2002) suggest the existence of the Precaspian Isthmus. In the

432 Artinskian the Isthmus was already quite high and delivered significant amount of clastic

433 material to the Precaspian basin (Fig. 6).

434 5.5 Sea-level fluctuations within the Asselian-Sakmarian transition.

435 According to the widely accepted model of the global sea-level fluctuations, the sea-

436 level at the Asselian-Sakmarian transition had dropped and the boundary coincided with a

437 global sequence boundary (Ross and Ross, 1987a). This interpretation is accepted in some

438 subsequent publications ( Golonka and Kiessling, 2002; Haq and Schutter, 2008; Rygel et al.,

439 2008). However, other authors proposed the coincidence of the Asselian-Sakmarian boundary

440 with a global flooding surface ( Boardman, Darwin R., II et al., 2009; Davydov et al., 1997a;).

441 The model of Ross and Ross (1987) was developed from the integration of the Southern

442 Ural and West Texas data based on the commonly accepted fusulinid-ammonid correlation of

443 these regions ( Furnish and Glenister, 1971; Jin et al., 1994, 1994; Ross, 1967). In this

444 correlation the middle Asselian of the Southern Ural corresponds to the Neal Ranch Formation

445 of Texas and the upper Asselian (Shikhanian) corresponds to the gap between the Neal Ranch

446 and Lenox Hill Formations of Texas. The latter formation was correlated with the entire

447 Sakmarian (Ross and Ross, 1987a). The most recent updates on the correlation of these

448 successions, involving new fusulinid and conodont biostratigraphic and taxonomic data the

449 from Ural, Nevada and Texas ( Chernykh, 2006; Chuvashov et al., 2002; Davydov et al., 1997a,

450 b; Wardlaw and Davydov, 2000;), provided a quite different correlation. The Neal Ranch

451 Formation is correlated with the upper Asselian and lower Sakmarian of the Southern Ural and

452 the Lenox Hill Formation – with the entire Artinskian, probably, including the lower Kungurian.

453 Moreover, it was clarified that the Asselian-Sakmarian boundary in both Texas and the

454 Southern Ural coincided with a flooding surface (Davydov et al., 1997a,b; Davydov et al., 2003;

455 Wardlaw and Davydov, 2000). It is now clear that the interpretation of the Kurmaya seismites

456 horizons as a conglomerate associated with sequence boundary within the Asselian-Sakmarian

457 transition at the Sakmarian type, Kondurovsky section in the Southern Ural (Ross and Ross,

458 1987), mislead the latter authors with the global correlation and sequence stratigraphy.

459 Refining the biostratigraphy of the conodonts and fusulinids in the Southern Ural and

460 the other regions provided the precise correlation of the Asselian-Sakmarian transition of the

461 Southern Ural with that of the Canadian Arctic, Iran, Nevada, New Mexico, Oklahoma, Texas,

462 Spitsbergen, Barents Sea, North and South China. The drop in global sea-level and the

463 occurrence of the associated sequence boundary occurred sometime within the early to middle

464 Tastubian (early Sakmarian) ( Beauchamp et al., 2009; Boardman, Darwin R., II et al., 2009;

465 Chernykh, 2006; Davydov, 1995, 1997a, b; Kozur and Le Mone, 1995; Leven and Gorgij, 2011;

466 Ehrenberg et al., 2000; Nilsson and Davydov, 1997; Wardlaw and Davydov, 2000).

467 The beginning of the Sakmarian was associated with a global sea level high stand. That is

468 opposite to what happened at the beginning of the Bashkirian glaciation, where the emergence

469 of the Alleghanian Isthmus corresponds with the one of the largest global sea level drops in the

470 Phanerozoic (Davydov and Cozar, 2018, in press; Ross and Ross, 1987b) Therefore, the global

471 sea-level fluctuations within the Asselian-Sakmarian transition did not coincide with the

472 emergence and development of the Precaspian Isthmus. The UPTS closure was essentially

473 caused by a tectonic event, associated with the active tectonic collisions of Kazakhstania, the

474 East Ural and Magnitogorsk Arc with the southern part of Russian Platform and Karpinsky Swell

475 at that time (Fig. 6). The event is best documented by the occurrence of the olistolith and

476 seismite horizons in the Southern Ural and by the termination of marine sedimentation at the

477 end of the Asselian in the Donets Basin and the southern periphery of the Precaspian Basin and

478 in the surrounding areas in the Turan and Scythian Platforms ( Afanasenkov et al., 2008;

479 Nikishin et al., 1996; Pronin et al., 1997). The connection between the Paleo-Tethys and the

480 Paleo-Arctic oceans during the rest of the Permian time was never re-established.

481 5.6 Seaway closure and climate

482 The chronostratigraphic constraints on the age of glacial events in the Permian are poorly

483 established and still in a state of flux. The new CA-IDTIMS data (Metcalfe et al., 2015)

484 significantly changes the chronostratigraphic framework of the Guadalupian-Lopingian glacial

485 episodes (P3 and P4 glaciations). The chronostratigraphic records from the high-latitude regions

486 of Gondwana produce quite a controversial picture concerning the glacial episodes in the

487 Gzhelian, Asselian and Sakmarian. The early Permian time is considered the apex of the late

488 Paleozoic Ice Age with the glaciations extending from the Gzhelian through the Asselian and

489 Sakmarian (Isbell et al., 2013). But the chronostratigraphy of the glacial episodes remains vague

490 given the paucity of the paleontological and geochronological constraints (Birgenheier et al.,

491 2009; Fielding et al., 2008; Mory et al., 2008). The latest paleontological and paleoecological

492 proxies link the early Permian glaciation (P1) in Australia to only the early Sakmarian (Haig et

493 al., 2014). That is consistent with the low value of the atmospheric carbon dioxide at this time

494 (Foster et al., 2017), which corresponds with the glacial episode in the early Sakmarian. The

495 significant rise of the CO2 and the corresponding global warming episode (Montanez and

496 Poulsen, 2013), occurred in the late Sakmarian (Fig. 7). The data on the carbon and oxygen

497 stable isotopes has been also obtained recently in the Southern Ural, from the type section of

498 the Sakmarian, the Kondurovsky section, and from the GGSP of the Sakmarian Stage at the

499 Usolka section (Zeng et al., 2012). The data reveals a significant shift of δ13C towards the

500 positive values in the early Sakmarian and back to the lighter isotopes in the late Sakmarian.

501 The data on the geographic extension of the Late Paleozoic glaciations (Isbell et al., 2012)

502 implies that the early Sakmarian glaciation was the largest within the Permian (Fielding et al.,

503 2008; Isbell et al., 2012).

504 The onset of Sakmarian P1 glaciation (Haig et al., 2014) coincided with the appearance

505 of the Precaspian Isthmus (Fig. 11). The emergence of the isthmus drastically changed the

506 pattern of the global ocean circulation similar to one that occurred after the appearance of the

507 Alleghanian Isthmus (Davydov and Cozar, 2018, in press; Saltzman, 2003). The changes

508 associated with the appearance of the Precaspian Isthmus are consistent with the scenario

509 proposed in the model of Rheic-Paleo-Tethys passage closure during the Serpukhovian-

510 Bashkirian transition (Smith and Read, 2000; Davydov and Cozar, 2018, in press; Montanez and

511 Poulsen, 2013). Evidently, the development of Precaspian Isthmus lead to the major

512 modification of the oceanic currents within the northern Paleo-Tethys, reorganization of overall

513 thermal regime of the atmosphere, the evolution of the climate and biota of the Earth.

514 The global oceanic circulation is often considered as one of the major factors controlling

515 the global climate (Rahmstorf, 2002). The final closure of the Central American Seaway and the

516 appearance of Panama Isthmus, according to some models, induced an increased poleward salt

517 and heat transport and intensified moisture supply to the northern high latitudes, which

518 resulted in the build of the Northern Hemisphere glaciation (Bartoli et al., 2011; Haug and

519 Tiedemann, 1998; Bartoli et al., 2005). The opening of the Drake Passage in the late Eocene

520 lead to the establishment of the Antarctic Circumpolar Current, that changed the ocean

521 circulation, caused water warming and enhanced precipitation in the Northern Hemisphere,

522 which in turn enhanced silicate weathering and CO2 drawdown and resulted in the

523 development of a large-scale ice sheet in Antarctica (Elsworth et al., 2017). Perhaps, the latter

524 model of the change of the ocean water circulation in the Paleo-Tethys is applicable to the case

525 of the circulation of the oceanic currents between the Paleo-Arctic and Paleo-Tethys. The

526 change of the ocean circulation in the Paleo-Arctic and Paleo-Tethys oceans would have caused

527 the re-direction of the oceanic warm currents from eastern Tethys toward the Pangea and

528 Gondwana margins. This may have enhanced precipitation and consequently silicate

529 weathering and CO2 drawdown around the Paleo-Tethys and the northern Gondwana shelves.

530 This model is consistent with the CO2 data in the Ural (Zeng et al., 2012), Nevada (Tierney et al.,

531 2008) and the global scale (Montanez et al., 2007). Starting in the Sakmarian, the global

532 biogeographic provincialism progressively increased, with the culmination at the end of the

533 Permian.

534 Recent modeling indicates that orbital forcing alone is not sufficient to cause the waxing

535 and waning of a massive Gondwanan ice sheet (Horton & Poulsen 2009, Horton et al. 2010).

536 The coincidence in the closure of the two main gateways, the Rheic-Paleo-Tethys gateway

537 during the Serpukhovian-Bashkirian transition and the Paleo-Tethys—Paleo-Arctic gateway

538 during the latest Asselian—earliest Sakmarian transition, with the onset of the two major

539 glacial events (Bashkirian and early Sakmarian) in the Late Paleozoic suggests that the closures

540 of the gateways, the appearances of the isthmus and the changes of the global oceanic currents

541 were the major drivers, along with the other less significant factors, in controlling the late

542 Paleozoic climate.

543 6. Conclusions

544 This paper has provided precise age constraints for the closure of the Paleo-Arctic—

545 Paleo-Tethys gateway by utilizing warm-water benthic foraminifera taxonomy and statistical

546 analysis. The major event, that drastically changed WWBF similarity among the analyzed

547 regions, occurred within the Asselian-Sakmarian transition during an interval of less than 0.5

548 Myr. The similarity index between the Tethyan and Boreal WWBF at the species level during the

549 Sakmarian and Artinskian time progressively decreased from 0.25 to 0.1 across the Sakmarian-

550 Artinskian boundary and was less than 0.1 towards the end of the Artinskian. From the

551 beginning of the Sakmarian, the fauna of the Boreal affinity predominated in the Precaspian

552 and the Southern Ural.

553 The temporal distribution of the olistostromes and seismites in the Southern Ural

554 suggests that major orogenic activity, and hence seismicity ceased at the Asselian-Sakmarian

555 transition in the Southern Ural and Scythian-Turan plate along the southern periphery of

556 Precaspian Basin. The fluvial fan deposits in the southern periphery of the Precaspian Basin

557 slope, suggesting the development of a weak orogeny along the border between Precaspian

558 Basin and Scythian-Turan plate.

559 The closure of the Paleo-Arctic--Paleo-Tethys seaway was caused by a tectonic event.

560 The event was associated with the collision of the southeastern part of Russian Platform,

561 Karpinsky Swell, the Southern Ural, Kazakhstania and Scythian-Turan plates along the southern

562 periphery of Precaspian Basin. The connection between the Paleo-Tethys and Paleo-Arctic

563 oceans in the post-Sakmarian time was never re-established.

564 The onset of P1 (early Sakmarian) glaciation coincided with the appearance of the

565 Precaspian Isthmus at the Asselian-Sakmarian transition (Fig. 1). The occurrence of this isthmus

566 was the second major tectonic event in the late Paleozoic, that radically changed the pattern of

567 the circulation of the global ocean, similar to one that occurred after the appearance of the

568 Alleghenian Isthmus. The development of the Precaspian Isthmus lead to the major

569 modification of the oceanic currents within the northern Paleo-Tethys, forcing changes in global

570 oceanic circulation, reorganization of the overall thermal regime of the atmosphere, and

571 consequently impacted climate and biota evolution of the Earth. The coincidence in the closure

572 of the two main gateways, the Rheic-Paleo-Tethys gateway during Serpukhovian-Bashkirian

573 transition and the Paleo-Tethys—Paleo-Arctic gateway during latest Asselian—earliest

574 Sakmarian transition, with the onset of the two major glacial events (Bashkirian and early

575 Sakmarian) in the late Paleozoic suggests that the closures of the gateways, the appearances of

576 the isthmus and the changes of the global oceanic currents were the major drivers for the

577 glaciation along with the other less significant factors controlling the Late Paleozoic climate.

578 Acknowledgments

579 The work was supported by the Ministry of Education and Science of the Russian Federation

580 contract No. 14.Y26.31.0029 in the framework of the Resolution No.220 of the Government of

581 the Russian Federation. The funds from the subsidy allocated to Kazan Federal University for

582 the state assignment #5.2192.2017/4.6 in the sphere of scientific activities and in part from the

583 subsidy of the Russian Government to support the Program of Competitive Growth of Kazan

584 Federal University among the World Leading Academic Centers are highly appreciated. I express

585 gratitude for the help of my great colleague and friend Professor Walter S. Snyder, who

586 carefully edited the text and made useful suggestions. Both W.S. Snyder and G.A. Mizens are

587 thanked for providing the photos of olistolith and seismite from the Southern Ural. The libraries

588 at Boise State University and Florida International University provided access to the literature

589 sources. The edits and suggestions of the reviewers greatly improved the manuscript.

590

591 Figures caption

592 Figure 1. Gzhelian-Asselian (A) and Sakmarian-Artinskian (B) paleogeography and foraminiferal

593 paleobiogeography (modified from Blakey, 2013). In pre-Sakmarian, warm-water benthic

594 foraminifera were freely dispersed from the Paleo-Tethys to Paleo-Arctic through the Uralian-

595 Tethyan Seaway. The emergence of the Precaspian Isthmus at the Asselian-Sakmarian

596 transition completely isolated the Boreal and the Tethyan foraminifera.

597 Figure 2. Multivariate cluster analyses with Raup-Crick probabilistic similarity index (PSI) at the

598 species level. During the early-middle Asselian and late Asselian the WWBF of Donets Basin, the

599 Precaspian and the Southern Ural belonged to the Tethyan Realm and had strong similarity with

600 the WWBF of the other Tethyan regions. From the early Sakmarian through the late Artinskian

601 the WWBF of the Precaspian and the Southern Ural belonged to Boreal affinity and got

602 dissimilar with the fauna of the Tethyan realm.

603 Figure 3. Multivariate cluster analyses performed with the Raup-Crick probabilistic similarity

604 index (PSI) at the genus level. The tendencies in the similarity/dissimilarity of the WWBF of

605 Donets Basin, the Precaspian and the Southern Ural regions and the other regions of Boreal and

606 Tethyan Realms are generally the same as for the PSI at the species level.

607 Figure 4. Non-metric multivariate ordination analysis with the Raup-Crick probabilistic

608 similarity index (PSI) at the species level. The taxonomic composition of the WWBF of the

609 Paleo-Arctic and Tethys significantly overlap during the Asselian, but the taxonomy becomes

610 completely dissimilar during the early Sakmarian through the late Artinskian time.

611 Figure 5. The summary of the similarities dynamics of the Boreal-dominated and Tethyan-

612 dominated WWBF at the species and generic levels from the late Gzhelian through Cisuralian

613 time. PSI - Raup-Creek probabilistic similarity index between Tethyan and Paleo-Arctic basins;

614 PSI A_U - Raup-Crick probabilistic similarity index between the WWBF of paleo-Arctic and Ural-

615 Precaspian. Gzhel., Gzhelian; Kung., Kungurian (part); Earl.—Midl., Early—Middle. The

616 separation of Paleo-Tethyan and Paleo-Arctic oceans occurred at the Asselian--Sakmarian

617 transition due to the collisions of Scythian and Turan microplates with southern Russian

618 Platform, Kazakhstania and the Ural. The similarity between WWBF of the Ural-Paleo-Arctic and

619 Paleo-Tethys increased through the Asselian and dropped drastically at the Asselian-Sakmarian

620 transition. After that, the WWBF of Ural-Paleo-Arctic and Paleo-Tethys oceans become

621 dissimilar. At the same time, the similarity of the WWBF of the Southern Ural--Precaspian and

622 Paleo-Arctic had been slowly increasing during the Asselian and had increased quite sharply at

623 the beginning of the Sakmarian, the fauna of these two areas became completely similar.

624 During the Kungurian the WWBF in the Southern Ural-Precaspian and Paleo-Arctic basins went

625 extinct.

626 Figure 6. Paleogeography of the Ural-Precaspian region and surrounding areas during

627 Moscovian-Asselian and Sakmarian (modified from Fedorenko, O.A. and Miletenko, N.V., 2002).

628 Free oceanic connection between Paleotethys and Paleo-Arctic during Moscovian-Asselian

629 provides extensions of fusulinid’s Tethyan domain over the Precaspian, the Southern Ural, and

630 Donets Basins. At the Asselian-Sakmarian transition after the emergence of Precaspian Isthmus,

631 the fusulinid’s Boreal domain extended into the Southern Ural and Precaspian but were not

632 able to migrate to Tethys. Both areas in Sakmarian through Kungurian were still located within

633 the tropics. Fluvial fans deposits during Moscovian-Asselian exist only along Southern Ural

634 foldbelt and their shape indicate northern direction of the Urals oceanic currents. In Sakmarian

635 and Artinskian the fluvial fans developed along South Ural foldbelt as well as in southern

636 periphery of the Precaspian Basin along the Precaspian Isthmus.

637 Figure 7. Temporal distribution of the warming and cooling events along the North American

638 shelves for the Serpukhovian through the Artinskian, relative diversity of the WWBF of the

639 Western Tethys, sea-water surface temperature, sea-level oscillation, fluctuations of the

640 atmospheric carbon dioxide and the records of the glaciations from Eastern Australia. Green

641 arrows indicate migration events of the WWBF from Tethys to North America, which are

642 associated with the global warming events and an increase of the diversity of foraminifera. The

643 immigration of the WWBF to North America occurred rapidly and abruptly and is associated

644 with the warming events. The FOD’s of many important genera in North America are always

645 delayed in respect to the appearance of these taxa elsewhere. Global cooling events are

646 associated with the decrease of the diversity of the foraminifera, intensive sea-level

647 oscillations, and low atmospheric carbon dioxide value. P2 glacial event in Australia is divided

648 into two glacial episodes that are not recognized as separate yet. The late Artinskian warming

649 spike according to the fusulinid data from Timor and Thailand (Haig et al., 2014; Ueno et al.,

650 2015). US Midcontinent sea-level fluctuation data from (Ross and Ross, 1988). Oxygen δ18O (‰

651 V-SWOM) sea-surface temperature data from (Chen et al., 2016). The question mark in the sea-

652 surface temperature curve represents the potential break in the sedimentation at the Naqing

653 section in South China, that is recognized from fusulinids record (Anonymous, 1994).

654 Atmosphere carbon dioxide data from Foster et al., (2017).

655 The scale is calibrated with the conodont—foraminiferal zonation and U-Pb ages from

656 the Carboniferous-Permian Composite (Davydov et al., 2012). Abbreviation: Desmoin. –

657 Desmoinesian; Msr. – Missourian. Fusulinid zonation: 1, Eostaffella tenebrosa; 2,

658 Pseudoendothyra globosa-Neoarchaediscus parvus; 3, Eostaffella mirifica-Eostaffellina protvae;

659 4, Monotaxinoides transitorius- Plectostaffella acuminulata; 5, Plectostaffella bogdanovkensis;

660 6, Semistaffella variabilis; 7, Pseudostaffella antiqua; 8, Pseudostaffella praegorskyi-

661 Staffellaeformis staffellaeformis; 9, Profusulinella parva-Ozawainella pararhomboidalis; 10,

662 Verella spicata- Aljutovella tikhonovichi; 11, Aljutovella aljutovica; 12, Aljutovella priscoidea; 13,

663 Moellerites lopasnensis- Fusulinella subpulchra; 14, Fusulinella colaniae-Beedeina kamensis; 15,

664 Fusulinella bocki-Fusulina cylindrica; 16, Protriticites ovoides-Praeobsoletes burkemensis; 17,

665 Protriticites pseudomontiparus; 18, Montiparus paramontiparus; 19, Montiparus subcrassulus;

666 20, Rauserites quasiarcticus; 21, Rauserites ofive; 22, Rauserites rossicus; 23, Rauserites

667 stuckenbergi; 24, Jigulites jigulensis; 25, Daixina sokensis; 26, Ultradaixina bosbytauensis-

668 Schwagerina robusta; 27, Sphaeroschwagerina vulgaris aktjubensis; 28, Pseudoschwagerina

669 robusta; 29, Sphaeroschwagerina gigas; 30, Sakmarella moelleri; 31, Schwagerina verneuili; 32,

670 Pseudofusulina urdalensis-Ps. plicatissima; 33, Pseudofusulina concavuta-Ps. pedisequa; 34,

671 Pseudofusulina solida-Parafusulina lutugini; 35, Parafusulina solidissima-P. jenkinsi; 36,

672 Chalaroschwagerina vulgaris; 37, FAD Pamirina; 38, Misellina minor; 39, Misellina parvicostata;

673 40, Armenina pamirensis, FAD Pseudodoliolina.

674 Figure 8. The examples of olistolith and seismite from the Southern Ural.

675 Table 1. Multivariate similarity and distance indexes, Raup-Crick coefficient at species level.

676 Table 2. Multivariate similarity and distance indexes, Raup-Crick coefficient at genus level.

677 Supplementary material

678 Supplementary references used to create the database of warm-water benthic foraminifera

679 distribution.

680

681 References

682 Abilkhasimov, K.B., 2016. The character of formation of natural reservoirs in the Paleozoic deposits in

683 Precaspian depression and evaluation of the oil-gas pespectives. Academy of Natural History Publishing

684 House, Moscow.

685 Afanasenkov, A.P., Skvortsov, M.B., Nikishin, A.M., Murzin, S.M., Polyakov, A.A., 2008. Geological evolution

686 and petroleum systems in the north Caspian region. Moscow University Geology Bulletin = Vestnik

687 Moskovskogo Universiteta. Geologiya 63, 131–139. 10.3103/S0145875208030010.

688 Akhmetshina, L.Z., Kukhtinov, D.A., Kukhtinova, L.V., 2013. Atlas of paleoontological remains of the Permian

689 deposit of northern and eastern segments of Precaspian Basin (Kazakhstan part): 242 с., 61 палеонт.

690 табл., 39 ил., Alma-Ata, Kazakhstan.

691 Akramkhodjaev, A.M., Avazkhodjaeva, K.K., Labutina, L.I., 1979. Lithology, environments and oil-gas bearing

692 of pre-Jurassic deposits of Ustyurt. Fan Publishing House, Tashkent, Uzbekistan.

693 Akramkhodjaev, A.M., Yuldashe, Z.Y., Avazkhodjaeva, K.K., Labutina, L.I., 1981. Key and parametric wells of

694 Ustyurt. Fan Publishing House, Tashkent, Uzbekistan.

695 Anonymous, 1994. Excursion Guidebook. Marine Permian in Guizhou and Guangxi. International

696 symposium on Permian stratigraphy, environments & resources with International meetings of Pangea

697 project GSGP, IGCP 306 & 359. Guiyang, China.

698 Bakirov, K.K., Kukhtiniv, D.A., Chimbulatov, M.A., Yakovlev, A.B., 1978. Litological composition and

699 stratigraphy of clastic subsalt sediments of Kenkiyak oil-field. News of Academy of Science of the

700 Kazakhstan S.S.R., Geological Series, 43-49 (In Russian).

701 Bartoli, G., Hoenisch, B., Zeebe, R.E., 2011. Atmospheric CO (sub 2) decline during the Pliocene

702 intensification of Northern Hemisphere glaciations. Paleoceanography 26, Citation PA4213.

703 10.1029/2010PA002055.

704 Bartoli, G., Sarnthein, M., Weinelt, M., Erlenkeuser, H., Garbe‐Schönberg, D., Lea, D., 2005. Final closure of

705 Panama and the onset of Northern Hemisphere glaciation. Earth and Planetary Science Letters 237, 33–

706 44. 10.1016/j.epsl.2005.06.020.

707 Beauchamp, B., Henderson, C.M., Grasby, S.E., Gates, L.T., Beatty, T.W., Utting, J., James, N.P., 2009. Late

708 Permian sedimentation in the Sverdrup Basin, Canadian Arctic; the Lindstrom and Black Stripe

709 Formations. Bulletin of Canadian Petroleum Geology 57, 167–191. 10.2113/gscpgbull.57.2.167.

710 Bensh, F.R., 1962. Pozdnekamennougol'nye i rannepermskie fuzulinidy Severnoi Fergany. Late

711 Carboniferous and early Permian fusulinids of Northern Fergana. In: Verkhov, V.I. (Ed.), Stratigraphy and

712 paleontology of Uzbekistan and surrounding areas. Book 1. Akademy of Sciences of Uzbekistan,

713 Tashkent, 186.

714 Bezhaev, M.M., 1968. The trace of late Paleozoic volcanic activity in the Ural. Doklady Akademii Nauk SSSR

715 179, 661.

716 Birgenheier, L.P., Fielding, C.R., Rygel, M.C., Frank, T.D., Roberts, J., 2009. Evidence for dynamic climate

717 change on sub-10 (super 6) -year scales from the late Paleozoic glacial record, Tamworth Belt, New

718 South Wales, Australia. Journal of Sedimentary Research 79, 56–82.

719 Blakey, R.C., 2013. Using paleogeographic maps to portray Phanerozoic geologic and paleotectonic history

720 of western North America. AAPG Bulletin 97, 146.

721 Boardman, Darwin R., II, Wardlaw, B.R., Nestell, M.K. (Eds.), 2009. Stratigraphy and Conodont

722 Biostratigraphy of the Uppermost Carboniferous and Lower Permian from the North American

723 Midcontinent. Kansas Geological Survey.

724 Brown, D., Alvarez Marron, J., Perez Estaun, A., Gorozhanina, Y., Baryshev, V., Puchkov, V., 1997. Geometric

725 and kinematic evolution of the foreland thrust and fold belt in the Southern Urals. Tectonics 16, 551–

726 562.

727 Cavazza, W., Roure, F., Spakman, W., Stampfli, G.M., Ziegler, P.A., 2004. The TRANSMED Atlas The

728 Mediterranean Region from Crust to Mantle;. Springer, Berlin Heidelberg.

729 Chen, B., Joachimski, M.M., Wang, X., Shen, S., Qi, Y., Qie, W., 2016. Ice volume and paleoclimate history of

730 the Late Paleozoic ice age from conodont apatite oxygen isotopes from Naqing (Guizhou, China).

731 Palaeogeography, Palaeoclimatology, Palaeoecology 448, 151–161. 10.1016/j.palaeo.2016.01.002.

732 Chernykh, V.V., 2006. Lower Permian conodonts in the Ural. Institute of Geology and Geochemistry, Uralian

733 Branch of the Russian Academy of Sciences, Ekaterinburg.

734 Chumakov, N.M. and Semikhatov, M.A., 2004. Climate and climatic zoning during the Permian and Early

735 Triassic. Trudy - Rossiyskaya Akademiya Nauk, Geologicheskiy Institut 550, 230–256.

736 Chumakov, N.M. and Zharkov, M.A., 2002. Climates during Permian-Triassic changes in the biosphere;

737 Paper 1, Early Permian climate. Stratigrafiya, Geologicheskaya Korrelyatsiya 10, 62–81.

738 Chuvashov, B.I., Chernykh, V.V., Bogoslovskaya, M.F., 2002. Biostratigraficheskaya kharakteristika

739 stratotipov yarusov nizhney permi. Biostratigraphic characteristic of Lower Permian stage stratotypes of

740 Permian. Stratigrafiya, Geologicheskaya Korrelyatsiya 10, 3–19.

741 Chuvashov, V.I., 1998. Dynamics of the evolution of the Uralian Foredeep. Geotectonics 32, 186–200.

742 Cocks, L.R.M. and Torsvik, T.H., 2007. Siberia, the wandering northern terrane, and its changing geography

743 through the Palaeozoic. Earth-Science Reviews 82, 29–74. 10.1016/j.earscirev.2007.02.001.

744 Collins, L.S., Budd, A.F., Coates, A.G., 1996. Earliest evolution associated with closure of the Tropical

745 American Seaway. Proceedings of the National Academy of Sciences of the United States of America 93,

746 6069–6072.

747 Crowell, J.C., 1999. Pre-Mesozoic ice ages; their bearing on understanding the climate system.

748 Davydov, V.I., 1986. Upper Carboniferous and Asselian fusulinids of the Southern Ural. In: Chuvashov, E.Ya.

749 Leven and V.I. Davydov (Ed.), Carboniferous--Permian Boundary beds of the Ural, Pre-Ural and Central

750 Asia. Nauka Publishing House, Moscow, 77.

751 Davydov, V.I., 1992. Subdivision and correlation of Upper Carboniferous and Lower Permian deposits in

752 Donets Basin according to fusulinid data. Soviet Geology, 53-61 (In Russian).

753 Davydov, V.I., 1995. The Carboniferous/Permian Boundary in South China. Permophiles, 9–11.

754 Davydov, V.I., 1997. Middle/Upper Carboniferous Boundary: the problem of definition and correlation. In:

755 M. Podemski, S. Dybova-Jachowicz, J. Jureczka and R. Wagner (Ed.), Proceeding of the XIII International

756 Congress on the Carboniferous and Permian, Warszawa, 113–122.

757 Davydov, V.I., 2014. Warm water benthic foraminifera document the Pennsylvanian–Permian warming and

758 cooling events — The record from the Western Pangea tropical shelves. Palaeogeography,

759 Palaeocimatology and Palaeogeography 414, 284–295.

760 Davydov, V.I. and Arefifard, S., 2013. Middle Permian (Guadalupian) fusulinid taxonomy and biostratigraphy

761 of the mid-latitude Dalan Basin, Zagros, Iran and their applications in paleoclimate dynamics and

762 paleogeography. Geoarabia 18, 17–62.

763 Davydov, V.I. and Cozar, P., 2018, in press. The formation of the Alleghenian Isthmus triggered the

764 Bashkirian glaciation: Constraints from warm-water benthic foraminifera. Palaeogeography,

765 Palaeoclimatology, Palaeoecology, 1–15. 10.1016/j.palaeo.2017.08.012.

766 Davydov, V.I., Korn, D., Schmitz, M.D., 2012. The Carboniferous Period. In: Gradstein, F.M., Ogg, J.G.,

767 Schmitz, M.D., Ogg, G. (Eds.), The Geologic Time Scale 2012. Elsevier, Amsterdam, 603–651.

768 Davydov, V.I., Nilsson, I., Stemmerik, L., 2001. Fusulinid zonation of the Upper Carboniferous Kap Jungersen

769 and Foldedal Formations, southern Amdrup Land, eastern north Greenland. Bulletin of the Geological

770 Society of Denmark 48, 31–77.

771 Davydov, V.I., Schiappa, T.A., Snyder, W.S., 2003. Testing the Sequence Stratigraphy Model: Response of

772 Fusulinacean Fauna to Sea Level Fluctuations (examples from Pennsylvanian and Cisuralian of Pre-

773 Caspian-southern Urals Region. In: Olson, H. C., and R. M. Leckie (Ed.), Micropaleontologic Proxies for

774 Sea-Level Changes and Stratigraphic Discontinuities. SEPM Special Publication. SEPM (Society for

775 Sedimentary Geology), SEPM, 359–375.

776 Davydov, V.I., Snyder, W.S., Spinosa, C., 1997a. Fusulinacean biostratigraphy and sequence stratigraphy of

777 the upper Paleozoic of the Southern Urals. Special Publications - Cushman Foundation for Foraminiferal

778 Research 36, 27–30.

779 Davydov, V.I., Snyder, W.S., Spinosa, C., 1997b. Permian foraminiferal biostratigraphy and sequence

780 stratigraphy of Nevada. Special Publications - Cushman Foundation for Foraminiferal Research 36, 31–

781 34.

782 Domeier, M. and Torsvik, T.H., 2014. Plate tectonics in the late Paleozoic. Geoscience Frontiers 5, 303–350.

783 10.1016/j.gsf.2014.01.002.

784 Dronov, V.I., Leven, E.Y., Pronina, G.P., 1997. Upper Permian deposits of the middle Sarykol Range in the

785 central Pamirs. Transactions (Doklady) of the Russian Academy of Sciences. Earth Science Sections 357,

786 1158–1160.

787 Ehrenberg, S.N., Pickard, N.A.H., Svana, T.A., Nilsson, I., Davydov, V.I., 2000. Sequence stratigraphy of the

788 inner Finnmark carbonate platform, Barents Sea: correlation between well 7128/6-1 and the shallow IKU

789 wells. Norsk Geologisk Tidsskrift 80, 129–162.

790 Elsworth, G., Galbraith, E., Halverson, G., Yang, S., 2017. Enhanced weathering and CO2 drawdown caused

791 by latest Eocene strengthening of the Atlantic meridional overturning circulation 10, 213–216.

792 10.1038/ngeo2888.

793 Ettensohn, F.R., Rast, N., Brett, C.E. (Eds.), 2002. Ancient Seismites. The Geological Society of America,

794 Boulder, Colorado.

795 Fedorenko, O.A. and Miletenko, N.V., 2002. Atlas of the Lithology-Paleogeographic, Structural, Palinspastic

796 and Geoenvironmental maps of Central Eurasia. Scientific Research Institute of Natural Resources

797 YUGGEO, Almaaty, Republic of Kazakhstan.

798 Festa, A., Ogata, K., Pini, G.A., Dilek, Y., Alonso, J.L., 2016. Origin and significance of olistostromes in the

799 evolution of orogenic belts: A global synthesis;. Gondwana Research 39, 180–203.

800 Fielding, C.R., Frank, T.D., Birgenheier, L.P., Rygel, M.C., Jones, A.T., Roberts, J., 2008. Stratigraphic imprint

801 of the late Palaeozoic ice age in eastern Australia; a record of alternating glacial and nonglacial climate

802 regime. Journal of the Geological Society of London 165, 129–140.

803 Filimonova, T.V., Gorozhanina, E.N., Isakova, T.N., Gorozhanin, V.M., 2015. Cisuralian Series of the Permian

804 System of the Southeastern SolIletsk Swell: Biostratigraphy and Lithology. Stratigrafiya, Geologicheskaya

805 Korrelyatsiya 23, 131–154.

806 Foster, G.L., Royer, D.L., Lunt, D.J., 2017. Future climate forcing potentially without precedent in the last

807 420 million years. Nature Communication 8, 1–8. 10.1038/ncomms14845.

808 Furnish, W.M. and Glenister, B.F., 1971. Permian ammonoid zonation. Bulletin of Canadian Petroleum

809 Geology 19, 328–329.

810 Gavrilov, B.P., 2011. Geodynamic model of the evolution of northern Ustyurt and surrounding regions of

811 Turan Plate in regards of Paleozoic oil-gas prospectives. Geology, geophysics and oil-gas fileds

812 production.

813 Golonka, J., 2007. Phanerozoic paleoenvironment and paleolithofacies maps; Mesozoic. Geologia (Krakow)

814 33, 211–264.

815 Golonka, J., 2011. Phanerozoic palaeoenvironment and palaeolithofacies maps of the Arctic region: Arctic

816 Petroleum Geology. Spencer, A. M., Embry, A. F., Gautier, D. L., Stoupakova, A. V. & Sørensen, K. (eds.).

817 The Geological Society of London, London.

818 Golonka, J. and Kiessling, W., 2002. Phanerozoic time scale and definition of time slices. Special Publication

819 - Society for Sedimentary Geology 72, 11–20.

820 Gorsky, V.P. and Kalmykova, M.A. (Eds.), 1986. Atlas of characteristic assemblages of Permian fauna and

821 flora of the Ural and Russian Platform. Nedra Publishing House, Leningrad.

822 Groves, J.R. and Wang, Y., 2009. Foraminiferal diversification during the late Paleozoic ice age. Paleobiology

823 35, 367–392.

824 Gusev, A.K., Bogatyrev, V.V., Igonin, V.M., Solodukho, M.G., 1968. Stratigraphy of Upper Paleozoic deposits

825 of Aktobe Preural. Kazan University, 219 p. (In Russian), Kazan.

826 Haig, D.W., McCartain, E., Mory, A.J., Borges, G., Davydov, V.I., Dixon, M., Ernst, A., Groflin, S., Hakansson,

827 E., Keep, M., dos Santos, Z., Shi, G.R., Soares, J., 2014. Postglacial Early Permian (late Sakmarian-early

828 Artinskian) shallow-marine carbonate deposition along a 2000 km transect from Timor to West

829 Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 409, 180–204.

830 10.1016/j.palaeo.2014.05.009.

831 Hammer, Ø. and Harper, D.A.T., 2006. Paleontological Data Analysis. Blackwell Publishing, Oxford.

832 Hammer, Ø., Harper, D.A.T., Ryan, P. D. . . , vol. , issue , : pp., 2001. PAST: Paleontological Statistics Software

833 Package for Education and Data Analysis. Palaeontologia Electronica 4, 9.

834 Haq, B.U. and Schutter, S.R., 2008. A chronology of Paleozoic sea-level changes. Science 322, 64–68.

835 Haug, G.H. and Tiedemann, R., 1998. Effect of the formation of the Isthmus of Panama on Atlantic Ocean

836 thermohaline circulation. Nature (London) 393, 673–676. 10.1038/31447.

837 Isbell, J.L., Henry, L.C., Gulbranson, E.L., Limarino, C.O., Fraiser, M.L., Koch, Z.J., Ciccioli, P.L., Dineen, A.A.,

838 2012. Glacial paradoxes during the late Paleozoic ice age; evaluating the equilibrium line altitude as a

839 control on glaciation. Gondwana Research 22, 1–19. 10.1016/j.gr.2011.11.005.

840 Izotova, M.N., 1985. Fusulinid zones of Lower Permian of northern and eastern slopes of Precaspian

841 depression. Soviet Geology, 76.

842 Izotova, M.N. and Vevel, Y.A., 1998. New species of fusulinoids from the eastern Caspian Sea region.

843 Paleontological Journal 32, 332–335.

844 Jablonski, D., Roy, K., Valentine, J.W., 2006. Out of the Tropics: Evolutionary Dynamics of the Latitudinal

845 Diversity Gradient. Science 314, 102–106.

846 Jin, Y.G., Glenister, B.F., Furnish, W.M., Kotlyar, G., Wardlaw, B.R., Kozur, H., Ross, C., Spinosa, C., 1994.

847 Revised operational scheme of Permian chronostratigraphy. Permophiles 25, 12–15.

848 Kaz'min, V.G. and and Natapov, L.M., 1998. The paleogeographic Atlas of Northern Eurasia. Institute of

849 Tectonics of Lithospheric Plates, Moscow, Moscow.

850 Kaz'min, V.G., Bush, A.V., Lobkovsky, L.I., 2008. Role of Transverse Strike-Slip Faults in the Structure of the

851 Karpinsky Ridge and Their Kinematics. Geotectonics 42, 176–185.

852 Keller, B.M., 1949. Flysh Paleozoic formation in Zalair Synclynorium in Southern Ural and and similar

853 deposits in surrounding areas. Transactions of Geological Institute of Academy of Sciences of USSR 104,

854 1-165 (In Russian).

855 Khain, V.E. and Popkov, V.E. (Eds.), 2009. Tectonics of southern periphery of East-European Platform.:

856 Кубан. гос. ун-т. 2009.213 с. Kubanian Federal University, Krasnodar.

857 Khvorova, I.V., 1961. Flysch and lower molasse formations of the south Ural. Transactions of Geological

858 Institute, Academy of Sciences of the USSR 37, 1–351.

859 Korenevsky, S.M., Bobrov, V.P., Suprunyuk, K.S., Khruschev, D.P., 1968. Halogen formations of North-West

860 Donets Basin and Dniepr-Donets trough. Nedra, Moscow.

861 Kozur, H. and LeMone, D.V., 1995. The Shalem Colony Section of the Abo and Upper Hueco Formation of

862 the Robledo Mountains, Dona Ana County, New Mexico: stratigraphy and new conodont-based age

863 determinations. Museum of Natural History and Science Bulletin 6, 39–55.

864 Lawver, L.A., Gahagan, L.M., Norton, I., 2011. Palaeogeographic and tectonic evolution of the Arctic region

865 during the Palaeozoic. Memoirs of the Geological Society of London 35, 61–77. 10.1144/M35.5.

866 Letavin A.I., Petrenko P.A., Lopatin A.F., Sterlenko Yu.A., Varjagov S.A., Voblikov B.G., 1994. To the problem

867 of prognostic estimation of hydrocarbons in Paleozoic deposits of the Peredovoi Ridge of Northern

868 Caucasus and Fore-Caucasian. Geologiya Nefti i Gaza, 10–14 (In Russian).

869 Leven, E.Y., 2004. Fuzulinidy i permskaya shkala Tetisa. Fusulinids and Permian scale of Tethys. Stratigrafiya,

870 Geologicheskaya Korrelyatsiya 12, 33–47.

871 Leven, E.Y. and Gorgij, M.N., 2011. The Kalaktash and Halvan assemblages of Permian fusulinids in the

872 Padeh and Sang-Variz sections (Halvan Mountains, Yazd Province, central Iran). Stratigrafiya

873 Geologicheskaya Korrelyatsiya 19, 20–39.

874 Leven, E.Y. and Scherbovich, S.F., 1978. Fuzulinidy i stratigrafiya assel'skogo yarusa Darvaza. Fusulinids and

875 stratigraphy of the Asselian Stage of the Darvaz Range. Nauka, Moscow.

876 Leven, E.Y. and Shcherbovich, S.F., 1980. New fusulinid species from the Sakmarian of the Darvaz.

877 Paleontological Journal 14, 18–29.

878 Lowry, D.P., Poulsen, C.J., Horton, D.E., Torsvik, T.H., Pollard, D., 2014. Thresholds for Paleozoic ice sheet

879 initiation. Geology (Boulder) 42, 627–630. 10.1130/G35615.1.

880 Mamet, B., 1977. Foraminiferal zonation of the Lower Carboniferous; methods and stratigraphic

881 implications. Dowden, Hutchinson, & Ross, Stroudsburg, Pa.

882 Maslov, V.V., Goryunova, L.F., Gibshman, N.B., 2016. Evaluation of prospecitves of the oil-gas production in

883 the Upper Paleozoic of Ustyurt on the bases of biostratigraphic analyses. Neftyegaz Territury, 57-63 (In

884 Russian).

885 McCalpin, J.P. (Ed.), 2009. Paleoseismology, Second edition;. Elsevier, Burlington, MA.

886 Metcalfe, I., Crowley, J.L., Nicoll, R.S., Schmitz, M., 2015. High-precision U-Pb CA-TIMS calibration of Middle

887 Permian to Lower Triassic sequences, mass extinction and extreme climate-change in eastern Australian

888 Gondwana. Gondwana Research 28, 61–81.

889 Mizens, G.A., 1997. Upper Paleozoic flysh of western Ural. Institute of Geology and Geochemistry, Uralian

890 Branch, Russian Academy of Sciences, Sverdlovsk.

891 Mizens, G.A., 2002. Sedimentological bassins and geodynamic environments in the late Devonian-early

892 Permian of south of the Ural. Institute of Geology and Geochemistry, Uralian Branch, Russian Academy

893 of Sciences, 190 p. (In Russian), Sverdlovsk.

894 Montanez, I.P. and Poulsen, C.J., 2013. The late Paleozoic ice age; an evolving paradigm. Annual Review of

895 Earth and Planetary Sciences 41, 629–656. 10.1146/annurev.earth.031208.100118.

896 Mory, A.J., Redfern, J., Martin, J.R., Fielding, C.R., Frank, T.D., Isbell, J.L., 2008. A review of Permian-

897 Carboniferous glacial deposits in Western Australia. Special Paper - Geological Society of America 441,

898 29–40. 10.1130/2008.2441(02).

899 Nikishin, A.M., Ziegler, P.A., Bolotov, S.N., Fokin, P.A., 2012. Late Palaeozoic to Cenozoic evolution of the

900 Black Sea-southern Eastern Europe region; a view from the Russian Platform. Turkish Journal of Earth

901 Sciences 21, 571–634. 10.3906/yer-1005-22.

902 Nikishin, A.M., Ziegler, P.A., Stephenson, R.A., Cloetingh, S. A. P. L., Furne, A.V., Fokin, P.A., Ershov, A.V.,

903 Bolotov, S.N., Korotaev, M.V., Alekseev, A.S., Gorbachev, V.I., Shipilov, E.V., Lankreijer, A., Bembinova,

904 E.Y., Shalimov, I.V., 1996. Late Precambrian to Triassic history of the East European Craton; dynamics of

905 sedimentary basin evolution. Tectonophysics 268, 23–63.

906 Nikolaev, A.I., 2011. Fusulinoids of Moscovian Stage in the Precaspian bassin. Transactions of VNIGRI, 168.

907 Nilsson, I. and Davydov, V.I., 1992. Middle Carboniferous-Lower Permian fusulinid stratigraphy in central

908 Spitsbergen, Arctic Norway. In: Mork, A. (Ed.), IKU Report 23.1356.00/07/92, Trondheil, 182.

909 Nilsson, I. and Davydov, V.I., 1997. Fusulinid biostratigraphy in Upper Carboniferous (Gzhelian) and Lower

910 Permian (Asselian-Sakmarian) succession in Spitsbergen, Arctic Norway. Permophiles, 18–27.

911 Pronin, A.P., Kuanyshev, F.L., Salykhova, A., Mil'kina, N.V., 2011. New data on the Paleozoic deposits in the

912 area of joining of Precaspian depression and Turan Plate (Kaspian Sea subsurface). Oil and Gas Geology,

913 21-25 (In Russian).

914 Pronin, A.P., Turkov, O.S., Kalmuratova, S.A., Mil'kina, N.V., 1997. New data regarding the origin of the

915 Paleozoic deposits of Buzachi peninsula. Geology of Kazakstan, 43-52 (In Russian);

916 Pronina, G.P., 1999. Korrelyatsiya verkhnepermskikh otlozheniy Boreal’noy oblasti po melkim

917 foraminiferam [Correlation of the Upper Permian deposits of the Boreal Realm based on small

918 foraminifers]. Proceeding of International Symposium “Upper Permian Stratotypes of the Volga Region”,

919 182-191 (In Russian).

920 Puchkov, V.N., 1997. Tectonics of the Ural; modern concepts. Geotectonics 31, 294–312.

921 Puchkov, V.N., 2000. Paleogeodynamics of Southern and Central Ural.: Пучков В.Н. Палеогеодинамика

922 Южного и Среднего Урала. – Уфа: Даурия, 2000. – 146 с. Dauria, Ufa, Russian.

923 Puchkov, V.N., 2009. The evolution of the Uralian orogen. Special Publications Geological Society 327, 161–

924 195.

925 Puchkov, V.N., 2010. The geology of the Ural and Pre-Ural. DesignPoligraphService, Ufa, Russian.

926 Rahmstorf, S., 2002. Ocean circulation and climate during the past 120,000 years. Nature (London) 419,

927 207–214. 10.1038/nature01090.

928 Rauser-Chernousova, D.M., 1965. Foraminiferi stratotipicheskogo razreza sakmarskogo yarusa (r. Sakmara,

929 Yuzhnyj Ural). Foraminifers from the Sakmarian type section (Sakmara River, S. Ural). Nauka, Moscow.

930 Rind, D., 1998. Latitudinal temperature gradients and climate change Rind, D. Journal of Geophysical

931 Research 103, 5943–5971.

932 Rogova, S.P. and and Pugacheva, R.K., 1983. Oil-gas perspectives of lower Permian deposits of Precaspian

933 Basin: ВНИГНИ, вып. 248, 1983. С. 111-121. Geology and oil-gas perspectives of slope zones of

934 Precaspian Basin. 248, 111-121 (In Russian).

935 Ross, C.A., 1967a. Development of fusulinid (Foraminiferida) faunal realms. Journal of Paleontology 41,

936 1341–1354.

937 Ross, C.A., 1967b. Late Paleozoic Fusulinacea from northern Yukon Territory. Journal of Paleontology 41,

938 709–725.

939 Ross, C.A., 1995. Permian fusulinaceans. In: Scholle, P.A., Peryt, T.M., Ulmer-Scholle, D.S. (Eds.), Permian of

940 Northern Pangea. Vol. 1: Paleogeography, Paleoclimate, Stratigraphy. Springer-Verlag, Berlin, 167–185.

941 Ross, C.A. and Ross, J.R.P., 1987a. Biostratigraphic zonation of late Paleozoic depositional sequences.

942 Special Publications - Cushman Foundation for Foraminiferal Research 24, 151–168.

943 Ross, C.A. and Ross, J.R.P., 1987b. Late Paleozoic sea levels and depositional sequences. Special Publications

944 - Cushman Foundation for Foraminiferal Research 24, 137–149.

945 Ross, C.A. and Ross, J.R.P., 1988. Late Paleozoic transgressive-regressive deposition. In: Wilgus C.K, Hastings

946 B. S. Posamentier H. Van Wagoner J. Ross C. A. Kendall C. G. St C. (Ed.), Sea-Level Changes—An

947 Integrated Approach, 227–247.

948 Rui Lin, Nassichuk, W.W., Thorsteinsson, R., 1994. The Lower Permian Fusulinacean Sphaeroschwagerina in

949 the Sverdrup Basin, Canadian Arctic Archipelago. Pangea: Global Environments and Resources, 891–905.

950 Rygel, M.C., Fielding, C.R., Frank, T.D., Birgenheier, L.P., 2008. The magnitude of late Paleozoic

951 glacioeustatic fluctuations; a synthesis. Journal of Sedimentary Research 78, 500–511.

952 Saltzman, M.R., 2003. Late Paleozoic ice age; oceanic gateway or pCO (sub 2) ? Geology 31, 151–154.

953 Scherbovich, S.F., 1969. Late Gzhelian and Asselian fusulinids in Precaspian Syneclise. Transactions of

954 Geological Institute, Academy of Sciences of the USSR 176, 1-104 (In Russian).

955 Schmitz, M.D. and Davydov, V.I., 2012. Quantitative radiometric and biostratigraphic calibration of the

956 Pennsylvanian - Early Permian (Cisuralian) time scale, and pan-Euramerican chronostratigraphic

957 correlation. Geological Society of America Bulletin 124, 549–577.

958 Scotese, C.R., 2015. Atlas of Plate Tectonic Reconstructions. Technical report.

959 https://www.researchgate.net/profile/Christopher_Scotese3.

960 Shi, G.R. and Waterhouse, J.B., 2010. Late Palaeozic global changes affecting high-latitude environments

961 and biotas: an introduction. Palaeogeography, Palaeoclimatology, Palaeoecology 298, 1–16.

962 Smith, L.B. and Read, J.F., 2000. Rapid onset of late Paleozoic glaciation on Gondwana; evidence from

963 Upper Mississippian strata of the Midcontinent, United States. Geology (Boulder) 28, 279–282.

964 10.1130/0091-7613(2000)028<0279:ROOLPG>2.3.CO;2.

965 Snyder, W.S., Spinosa, C., Davydov, V.I., Belasky, P., 1994. Petroleum geology of the southern Pre-Uralian

966 Foredeep with reference to the northeastern Pre-Caspian Basin. International Geology Review 36, 452–

967 472.

968 Stampfli, G.M., Hochard, C., Verard, C., Wilhem, C., vonRaumer, J., 2013. The formation of Pangea.

969 Tectonophysics 593, 1–19. 10.1016/j.tecto.2013.02.037.

970 Sukhov, E.E., 2003. Permian smaler foraminifera of Biarmian paleogeographic province. Kazan Federal

971 University, Kazan.

972 Tierney, K.E., Saltzman, M.R., Henderson, C.M., Davydov, V.I., Yan, J., Munnecke, A., Cramer, B.D., 2008.

973 Toward High Resolution Isotope Stratigraphy: Permian 87Sr/86Sr and δ13C Stratigraphy from

974 Limestones In China and Nevada. Abstracts with Programs - Geological Society of America 40, 402–403.

975 Ueno, K., Arita, M., Meno, S., Sardsud, A., Saesaengseerung, D., 2015. An Early Permian fusuline fauna from

976 southernmost peninsular Thailand; discovery of Early Permian warming spikes in the peri-Gondwanan

977 Sibumasu Block. Journal of Asian Earth Sciences 104, 185–196. 10.1016/j.jseaes.2014.10.030.

978 Uzakov, K., 1996. Lithological and biostratigraphic characteristics of pre-Jurrasic deposits of Eastern Ustyurt

979 (Karakalpastan). Uzbekiston Geologiya Zhurnali = Uzbekskiy Geologicheskiy Zhurnal, 52-67 (In Russian).

980 Volodin, D.F. (Ed.), 1993. Stratigraphical charts of Precambrian and Panerozoic of Ukraine.: Гос. ком.

981 Украйни по гєологии и использованию недр,. Geoprognoz, Kiev.

982 Volozh, Y.A., Bykadorov, V.A., Antipov, M.P., Parasyna, V.C., Rybal'chenko, V.V., Ogorodnikov, I.V., 2011.

983 Paleozoic and Triassic deposits of Ustyurt (Seismography, Paleogeography and oil-gaz bearing). Bulletine

984 of Moscow Society of Natural studies 86, 47-66 (In Russian).

985 Wardlaw, B.R. and Davydov, V.I., 2000. Preliminary Placement of the International Lower Permian Working

986 Standard to the Glass Mountains, Texas. Permophiles, 11–14.

987 Zeng, J., Cao, C., Davydov, V.I., Shen, S., Babinski, M., Trindade, R.I.F., Tohver, E., Collins, A.S., 2012. Carbon

988 isotope chemostratigraphy and implications of palaeoclimatic changes during the Cisuralian (Early

989 Permian) in the Southern Urals, Russia. Gondwana Research 21, 601–610. 10.1016/j.gr.2011.06.002.

990 Ziegler, P.A., 1989. Evolution of Laurussia: a study in late Palaeozoic plate tectonics. Kluwer Academic

991 Publishers, Dordrecht.

992 Zolotova, V.P. and Baryshnikov, V.V., 1978. Fusulinids of Kungurian Stage of Kamsk Preural.

993 Paleontologichesky Zhurnal, 22.

994 Zonenshain, L.P., Kuzmin, M.I., Natapov, L.M., 1990. Geology of the USSR; a plate-tectonic synthesis.

995 Geodynamics Series 21, 242.

996

997

310-295 Ma Uralian Foredeep

11 ? 10 12 8 7 9 5 6 4 3

2

1 Center of originations

A

295-290 Ma Uralian Foredeep

? 12 11 10 9 8 5 6-7 4 2

1 Center of originations

B

Analyzed locations: 1, Carnic Alps; 2, Taurids, Turkey; 3, Donets Basin; 4, Darvaz, C. Asia; 5, South Tian-Shan, Central Asia; 6-7, northern and eastern Precaspian; 8, Southern Urals; 9, Russian Platform; 10, Central Urals; 11, Timan-Pechora Basin Fauna distribution Tethyan affinity Boreal affinity

WWBF Realms: Tethyan Boreal North American

Fig. 1 Davydov Cluster analyses, species Raup-Crick coefficient Late Asselian Early Sakmarian, Tastubian

Early-middle Asselian 1 8 9 4 1 5 2 6-7 10 1 1

10 9 1 6-7 8 3 4 5 2 1 100 1 9 8 3 2 4 5 1

10 1 6-7 0.96 1.0 20 100 0.96 90 0.84 0.9 0.88 0.8 0.72 0.80 0.7 100 0.60 0.72 0.6 0.48 100 0.64 Similarity Similarity

Similarity 0.5 0.36 0.56 0.4 0.24 0.48 0.3 0.12 0.40 0.2 100 100 100 0.32 0.00 Cophenetic correlation 0.86 Cophenetic correlation 0.73 Cophenetic correlation 0.95

Late Sakmarian, Sterlitamakian Early Artinskian Late Artinskian Cophenetic correlation 0.83 1 8 9 2 4 1 1 6-7 10 1 8 9 4 5 2 1 1 6-7 1 10 8 9 5 4 1 1 10 6-7 100 100 0.96 30 0.96 predominantly 0.96 100 100 0.84 Tethyan fauna 0.84 0.84 100 0.72 0.72 0.72 0.60 predominantly 0.60 0.60 Boreal fauna 0.48 0.48 Similarity

Similarity 0.48 0.36 8 the regions Similarity with the fusulinids 0.36 100 50 0.36 0.24 of Tethyan 0.24 affinity; since 0.24 0.12 90 0.12 100 Sakmarian the 0.12 0.00 fusulinids of these 100 100 0.00 Cophenetic correlation 0.998 regions belong 0.00 Cophenetic correlation 0.96 Cophenetic correlation 0.96 to Boreal affinity 1, Carnic Alps; 2, Taurids, Turkey; 3, Donets Basin; 4, Darvaz; 5, South Tian-Shan’, Central Asia; 6-7, northern and eastern Precaspian; 8, southern Urals; 9, Russian Platform; 10, Central Urals; 11, Timan-Pechora

Fig. 2 Davydov Cluster analyses (genus), Raup- Crick coefficient Early Sakmarian Early-middle Asselian Late Asselian 1 9 8 4 1 5 2 6-7 10 1 1 9 8 3 4 5 1 2 1 10 1 6-7

9 10 1 5 6-7 8 3 4 1 2 1.0 10 30 60 50 60 0.96 0.9 80 80 0.95 80 50 70 0.90 0.90 0.8 0.84 0.85 0.7 0.78 0.80 0.6 0.72 0.75 90 Similarity

Similarity 0.66 0.5 0.70 Similarity 0.60 0.4 0.65 0.54 0.60 0.3 0.48 100 100 100 0.55 Cophenetic correlation 0.71 0.2 0.42 Cophenetic correlation 0.83 Cophenetic correlation 0.90

Late Sakmarian, Sterlitamakian Early Artinskian Late Artinskian

1 predominantly 1 1 8 9 5 4 1 6-7 8 9 10 1 2 4 1 5 6-7 8 9 10 1 4 2 1 6-7 10 1 70 1.0 1.0 100 Tethyan fauna 40 20 0.96 50 90 80 0.9 0.9 0.88 70 0.8 0.8 0.80 predominantly 0.7 0.7 0.72 Boreal fauna 0.6 0.6 80 0.64 60 8 the regions 0.5 0.5 0.56 Similarity with the fusulinids Similarity 0.4 Similarity 0.4 of Tethyan 0.48 affinity; since 0.3 0.3 0.40 Sakmarian the 0.2 0.32 0.2 fusulinids of these 100 100 100 0.1 regions belong 0.24 0.1 Cophenetic correlation 0.81 Cophenetic correlation 0.95 Cophenetic correlation 0.82 to Boreal affinity 1, Carnic Alps; 2, Taurids, Turkey; 3, Donets Basin; 4, Darvaz; 5, South Tian-Shan’, Central Asia; 6-7, northern and eastern Precaspian; 8, southern Urals; 9, Russian Platform; 10, Central Urals; 11, Timan-Pechora

Fig. 3 Davydov NMMS_species, Raup-Crick coefficient

Early-middle Asselian Late Asselian Early Sakmarian, Tastubian

2 11 11 0.24 0.24 0.32 0.18 5 2 0.18 11 0.24 0.12 0.12 1 0.16 0.06 1 0.06 5 10 4 10 0.08 0.00 3 0.00 5 6-7 4 6-7 9 0.00 4 2 Coordinate 2 -0.06 10 -0.06 Coordinate 2 Coordinate 2 6-7 -0.12 -0.12 -0.08 1 9 -0.18 9 -0.18 -0.16 Precaspian Isthmus -0.24 8 -0.24 3 8 8 -0.24 -0.30 -0.30 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 0.4 0.5 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24 0.32 -0.40 -0.32 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24 0.32 Coordinate 1 Coordinate 1 Coordinate 1

Late Sakmarian, Sterlitamakian Early Artinskian Late Artinskian predominantly 5 0.36 2 Tethyan fauna 11 0.32 0.12 4 0.30 Precaspian Isthmus 0.24 0.06 10 0.24 9 predominantly 0.16 0.00 11 8 Boreal fauna 0.18 Precaspian Isthmus -0.06 0.12 10 0.08 6-7 Precaspian Isthmus 11 -0.12 0.06 6-7 10 0.00 8 the regions Coordinate 2 Coordinate 2 Coordinate 2 4 -0.18 0.00 5 8 9 -0.08 1 with the fusulinids 4 -0.24 -0.06 of Tethyan 1 -0.16 6-7 9 2 -0.30 affinity; since -0.12 -0.24 Sakmarian the 8 -0.36 1 -0.18 fusulinids of these -0.48 -0.40 -0.32 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24 -0.40 -0.32 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24 -0.48 -0.40 -0.32 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24 regions belong Coordinate 1 Coordinate 1 Coordinate 1 to Boreal affinity 1, Carnic Alps; 2, Taurids, Turkey; 3, Donets Basin; 4, Darvaz; 5, South Tian-Shan’, Central Asia; 6-7, northern and eastern Precaspian; 8, southern Urals; 9, Russian Platform; 10, Central Urals; 11, Timan-Pechora

Fig. 4 Davydov 1.0

0.9

0.8 Arctic vs Tethys 0.7 species 0.6 genera Arctic vs Urals-Precaspian 0.5 species

Similarity 0.4 collision event 0.3

0.2 WWBF extinction in Boreal Realm

0.1

0 Late Earl.-Midl. Late Early Late Early Late Early Gzhel. Asselian Sakmarian Artinskian Kung.

300 295 290 285 Age (Ma) International Geologic Time Scale Fig. 5 Davydov 38 42 48 54 60 66 72 78 84 90 96 100 28 TIC RC OA KOK 28 LE PA KHM 24 N U TPB F PBA 24 R I M TIM A E U T 20 F

20 M Russian S G A U U Z KCB N F 16 F L C Platform B K KUA N 16 MSB

M 12 P PCB C 12 UST SHB VOR 8 AMD R PA K 8 MNB K DDB C 4 UKM 4 GCM Moscovian-Asselian 0 0 1000 0 42 48 54 60 66 72 78 84 90 96

36 42 48 54 60 66 72 78 84 90 96 100 28 PA Sakmarian WAR NRM 0 1000 28 BDM JGB 24

JNF High mountains 24 KHM Mountains 20 U N F U and plateau B F R I M 20 TIM A CSB Lowland KCB T 16 KUA Lacustrine- E alluvial land U 16 F IN MSB H High salinity lagoone S 12 U STF F R A 12 P Fluvial fan conglomerates T S PCB U AMD Shallow shelf 8 CKK 8 Outer shelf PA - PaleoArctic S KP KAR 4 Bathial basins Volcanic arcs DDB GCM 4 Continental slope Intermauntain depressions UKM and foredeeps 0 ain Ocean (abyssal) basins om Oceanic currents n d 0 thya Te JGB- Junggar-Balkhash Basin NUF - North Uralian Foldbelt JNF- Junggar Forebelt PAR - Parapamiz Zone AMD - Amu-Darya Block KAR - Karabagaz Massif PBA - Pribalkhash Zone BDM - Baidara Massif KCB - Karachytyr Basin PCB - Precaspian Basin SHB - South Hissar Basin CKK - Central Karakum KHM - Khanty-Mansy Block STF - South Tian-Shan foldbelt CNA - Central Kunlun Arc KOK - Kokshetau Massif SUF - South Uralian Foldbelt CPM - Central Pamirs KPS - Karpinsky Swell CSB - Chu-Sarysu Basin KUA - Kurama Arc TIM - Timan Block DDB - Dnepr-Donets Basin MGZ - Magnitogorsk Zone TPB - Timan-Pechora Basin UFB - Uralian Foreland EUF - East-Uralian Foldbelt MNB - Mangyshlak Basin UKM - Ukrainian Shield GCM - Great Caucasus Massif MSB - Moscow Basin HIN - Hindukush Block UST - Ustyurt NKL - North Kunlun VOR - Voronezh Shield 42 48 54 60 66 72 78 84 90 96

Fig. 6 Davydov Relative species diversity Biogenic apatite geochemical record US Midcontinent Fusulinid FAD in Fusulinid FOD Atmospheric Australian Regional 18 S in West Tethys δ O (‰V-SWOM) eustatic sea-level CO (p.p.m.) Glaciation

E Tethyan and Boreal in N. America 2 I Stage magnitude (m) R Stage 10 30 50 70 90 110 Events

realms Regional Stage

E 24 23 22 21 200 0 S Fusullinid Zones 0 277.0 Yangchienia Irenian 3000 2000 1000 40 Southern Urals record Solikamian Robustoschwagerina 280 Fillipovian Kungurian 39 Yangchienia P2B N Saranian

38 Leonardian A

I Biwaella, Toriyamaia 37

L 285.0 36 Sarginian A 35 Chalaroschwagerina R Artinskian Irginian 34 Chalaroschwagerina

U P2A

Burtsevian 33 Toriyamaia S

290 I 290.0 Lenoxian

C Sterlitamakian 32 Sakmarian Robustoschwagerina Urals-Tethys Tastubian 31 ? P1 30 Eoparafusulina Eoparafusulina gateway closure 295.0 Shikhanian 29 Pseudoschwagerina Asselian Uskalykian 28

Pseudoschwagerina Nealian 250 species 298.9 Sjuranian 27 Melekhovian 26 Schellwienia 300 Noginian 25 Schellwienia

Gzhelian Pavlovoposad. 24 N

Rusavkian 23 Biwaella irgilian V A 22

I 303.7 Dorogovilovian 20-21 Triticites (s.l.) .

N Kasimovian Khamovnich. 18-19 Msr A Krevyakian 17 Eowaeringella Eowaeringella V 307.0 Peskovian 16 C4 L Triticites (s.l.)

Myachkovian 15 Y

S Podolskian 14 Bedeeina Wedekindellina 310 Wedekindellina Desmoin. C3

N Moscovian Kashirian 13

N Bedeeina 12 E Tsninian 11 P 314.7 Vereian Profusulinella

Melekesian 10 Atokan Cheremshanian 9 Pseudostaffella Bashkirian Prikamian 8 Profusulinella Monotaxinoides transitorius 320 Severokeltmian 7 Krasnopolyan. 6 Pseudostaffella Voznesenian 5 Millerella Millerella Morrowan Rheic gateway closure C2 323.2 Zapaltjubian

N 4 A

I Protvian Monotaxinoides P

P 3

I Serpukhovian Steshevian transitorius S

S Tarussian

I 2

S C1

S Venevian 1

330 I 330.0 Chesterian (part) Chesterian Ma M

The occurrence of the taxon 15 20 25 30 Climatic optimum Climatic cooling Migration event in each province (ºC) Ocean surface temperature Seismites, olistostromes

Davydov Fig. 7 Seismites at Kondurovsky Section (photo courtesy of W. S. Snyder)

Kasimovian olistolith at Zianchurino village (photo courtesy of G. Mizens)

Fig. 8 Davydov Early‐Middle Asselian Late Asselian_species Region 123456‐7891011 Region 123456‐7891011 1 1 0,9965 1,0000 1,0000 0,9960 0,8955 0,8075 0,2155 0,1425 0,0000 1 1 1,0000 0,8785 1,0000 1,0000 0,9745 0,6095 0,3365 0,5130 0,0465 2 0,9965 1 0,9990 0,9995 1,0000 1,0000 0,8005 0,6090 0,2700 0,1275 2 1,0000 1 0,9430 0,9940 1,0000 0,9835 0,4240 0,4800 0,3460 0,0435 3 1,0000 0,9990 1 1,0000 0,9910 1,0000 0,9980 0,0440 0,0635 0,0000 3 0,8785 0,9430 1 0,9905 0,9210 0,9980 0,9565 0,9230 0,6510 0,2295 4 1,0000 0,9995 1,0000 1 1,0000 1,0000 0,9610 0,0010 0,0005 0,0000 4 1,0000 0,9940 0,9905 1 1,0000 0,9560 1,0000 0,0000 0,0000 0,0000 5 0,9960 1,0000 0,9910 1,0000 1 1,0000 0,9500 0,0265 0,0000 0,0000 5 1,0000 1,0000 0,9210 1,0000 1 0,9950 0,9995 0,0005 0,0100 0,0000 6‐7 0,8955 1,0000 1,0000 1,0000 1,0000 1 1,0000 0,9480 0,8855 0,0000 6‐7 0,9745 0,9835 0,9980 0,9560 0,9950 1 1,0000 0,9995 0,9850 0,8245 8 0,8075 0,8005 0,9980 0,9610 0,9500 1,0000 1 0,7040 0,0030 0,0000 8 0,6095 0,4240 0,9565 1,0000 0,9995 1,0000 1 0,9505 0,0145 0,0000 9 0,2155 0,6090 0,0440 0,0010 0,0265 0,9480 0,7040 1 1,0000 0,3565 9 0,3365 0,4800 0,9230 0,0000 0,0005 0,9995 0,9505 1 1,0000 0,9910 10 0,1425 0,2700 0,0635 0,0005 0,0000 0,8855 0,0030 1,0000 1 0,9935 10 0,5130 0,3460 0,6510 0,0000 0,0100 0,9850 0,0145 1,0000 1 1,0000 11 0,0000 0,1275 0,0000 0,0000 0,0000 0,0000 0,0000 0,3565 0,9935 1 11 0,0465 0,0435 0,2295 0,0000 0,0000 0,8245 0,0000 0,9910 1,0000 1

Early Sakmarian_species Late Sakmarian_species Region 12456‐7891011 Region 1456‐7891011 1 1 0.9995 1 1 0.078 0000 1110.874 0.002 0 0 0 0.001 2 0.9995 1 0.9625 0.981 0.0085 0000 411100000 4 1 0.9625 1 1 0.0225 0000 50.874 1 1 0.0165 0 0 0.001 0.0065 5 1 0.981 1 1 0.109 0 0 0.0005 0 6‐7 0.002 0 0.0165 1 1 1 0.8635 0.4005 6‐7 0.078 0.0085 0.0225 0.109 1 1 1 0.9925 0.6355 80001110.9995 0.0085 800001110.9715 0 900011110.0645 9000011110.0025 10 0 0 0.001 0.8635 0.9995 1 1 0.823 10 0 0 0 0.0005 0.9925 0.9715 1 1 0.9975 11 0.001 0 0.0065 0.4005 0.0085 0.0645 0.823 1 1100000.6355 0 0.0025 0.9975 1

Early Artinskian_species Late Artinskian_species Region 12456‐7891011 Region 1246‐7891011 1 1 0.99 0.964 0.9615 00000 110.0745 0.1055 0.0195 0 0 0 0.031 2 0.99 1 0.6485 0.47 0.1845 0.02 0.037 0.0125 0.05 2 0.0745 1 0.23 0.0315 0 0 0 0.049 4 0.964 0.6485 1 0.9555 00000 40.1055 0.23 100000.006 5 0.9615 0.47 0.9555 1 0.3485 0.108 0.184 0.1075 0.2095 6‐7 0.0195 0.0315 0 1 1 1 0.984 0.82 6‐7 0 0.1845 0 0.3485 11110.6145 800011110.979 8 0 0.02 0 0.108 11110.7325 900011110.982 9 0 0.037 0 0.184 11110.8405 10 0 0 0 0.984 1 1 1 0.9945 10 0 0.0125 0 0.1075 11111 110.031 0.049 0.006 0.82 0.979 0.982 0.9945 1 11 0 0.05 0 0.2095 0.6145 0.7325 0.8405 1 1

Regions with Tethyan dominant WWBF Regions with Boreal dominant WWBF Regions with WWBF of Tethyan affinity that in Sakmarian become of Boreal affinity

For the region names see Fig. 1. Early‐Middle Asselian_genus Late Asselian_genus Region 123456‐7891011 Region 123456‐7891011 1111110.9955 0.917 0.8815 0.7735 0.022 1 1 1 0,907 1 1 0,9965 0,327 0,303 0,3575 0,013 21111110.9665 0.9805 0.8275 0.5755 2 1 1 0,8855 1 1 0,966 0,034 0,2365 0,0715 0,004 311111110.2015 0.163 0 3 0,907 0,8855 1 1 0,932 0,999 0,635 0,5495 0,0845 0,0005 41111110.9785 0.0135 0.0015 0 4 111110,9805 0,387 0 0,0005 0 51111110.983 0.325 0.0075 0 5 1 1 0,932 1 1 1 0,4905 0,012 0,02 0 6‐7 0.9955 1111110.9905 0.948 0 6‐7 0,9965 0,966 0,999 0,9805 1 1 1 0,9925 0,924 0,5185 8 0.917 0.9665 1 0.9785 0.983 1 1 0.816 0.0005 0 8 0,327 0,034 0,635 0,387 0,4905 1 1 1 0,6255 0 9 0.8815 0.9805 0.2015 0.0135 0.325 0.9905 0.816 1 1 0.777 9 0,303 0,2365 0,5495 0 0,012 0,9925 1 1 1 0,846 10 0.7735 0.8275 0.163 0.0015 0.0075 0.948 0.0005 1 1 1 10 0,3575 0,0715 0,0845 0,0005 0,02 0,924 0,6255 1 1 1 11 0.022 0.5755 000000.777 1 1 11 0,013 0,004 0,0005 0 0 0,5185 0 0,846 1 1

Early Sakmarian_genus Late Sakmarian_genus Region 12456‐7891011 Region 1 4 5 6‐7 8 9 10 11 1 11110,1475 0000 1 110,9995 0,007 0000 2 1 1 1 0,997 0,012 0000 4 1110,001 0000 4 11110,064 0000 50,9995 1 1 0,081 0 0 0,0025 0,013 5 1 0,997 1 1 0,557 0,0055 0 0,007 0 6‐7 0,007 0,001 0,081 1 1 1 0,9035 0,3075 6‐7 0,1475 0,012 0,064 0,557 11110,9215 8 00011110,022 8 0 0 0 0,0055 1 1 1 0,9585 0 9 00011110,205 9 000011110,0355 10 0 0 0,0025 0,9035 1 1 1 0,9975 10 0 0 0 0,007 1 0,9585 1 1 1 11 0 0 0,013 0,3075 0,022 0,205 0,9975 1 1100000,9215 0 0,0355 1 1

Early Artinskian_genus Late Artinskian_genus Region 12456‐78 91011 Region 1 2 4 6‐7 8 9 10 11 1 1 1 0,9995 0,945 00000 1 10,434 0,2795 0,005 0 0 0 0,0035 2 1 1 0,9985 0,414 0,029 0 0,0005 0 0,001 2 0,434 1 0,724 0,0435 0 0 0 0,0465 4 0,9995 0,9985 1 0,9085 00000 40,2795 0,724 100000,006 5 0,945 0,414 0,9085 1 0,365 0,134 0,1945 0,122 0,231 6‐7 0,005 0,0435 01110,9975 0,9485 6‐7 0 0,029 0 0,365 11110,876 8 00011110,9975 8 0 0 0 0,134 11110,858 9 00011110,9975 9 0 0,0005 0 0,1945 11110,96 10 0 0 0 0,9975 1111 10 0 0 0 0,122 11111 110,0035 0,0465 0,006 0,9485 0,9975 0,9975 1 1 11 0 0,001 0 0,231 0,876 0,858 0,96 1 1

Regions with Tethyan dominant WWBF Regions with Boreal dominant WWBF Regions with WWBF of Tethyan affinity that in Sakmarian become of Boreal affinity 1 Database References 2 3 Akhmetshina, L.Z., Kukhtinov, D.A., and Kukhtinova, L.V., 2013, Atlas of paleoontological remains of the 4 Permian deposit of northern and eastern segments of Precaspian Basin (Kazakhstan part): 242 с., 61 5 палеонт. табл., 39 ил.: Alma-Ata, Kazakhstan. 6 Akramkhodjaev, A.M., Avazkhodjaeva, K.K., and Labutina, L.I., 1979, Lithology, environments and oil-gas 7 bearing of pre-Jurassic deposits of Ustyurt.: Tashkent, Uzbekistan, Fan Publishing House. 8 Akramkhodjaev, A.M., Yuldashe, Z.Y., Avazkhodjaeva, K.K., and Labutina, L.I., 1981, Key and parametric 9 wells of Ustyurt.: Tashkent, Uzbekistan, Fan Publishing House. 10 Alekseev, A.S., Goreva, N.V., Isakova, T.N., and Kossovaya, O.L., 2009, The stratotype of the Gzhelian 11 Stage (Upper Carboniferous) in Moscow Basin, Russia, in Puchkov, V.N., ed., Carboniferous type 12 sections in Russia, potential and global stratotypes: Ufa, Institute of geology of the Ufa scientific 13 Center, p. 165. 14 Alekseev, A.S., Goreva, N.V., Isakova, T.N., and Makhlina, M.K., 2004, Biostratigraphy of the 15 Carboniferous in the Moscow Syneclise, Russia: Newsletter on Carboniferous Stratigraphy, v. 22, p. 16 28–35. 17 Alekseev, A.S., Goreva, N.V., Kulagina E.I., Kossovaya, O.L., Isakova, T.N., and Reimers, A.N., 2002, Upper 18 Carboniferous of South Urals (Bashkiria, Russia). Guidebook of the field excursion., 56 p. 19 Alekseeva, I.A., Glushenko, N.V., Ivanov, V.K., Inosova, K.I., Kalmykova, M.A., Kashik, D.S., Kozitskaya, 20 R.I., Kruzina, A.K., Movshovich, Y.V., Redichkin, N.A., and Schwartzman, E.G., 1983, The Key-well of 21 the Carboniferous-Permian boundary beds in the south of East-European Platform (Gzhelian and 22 Asselian Stages, Skosyrskaya Well No 4199): Leningrad, Nauka Publishing House; 135 p. (In Russian), 23 Volume 12. 24 Alksne, A.E., 1976, New species of genus Daixina from the Upper Carboniferous depositis of Bashkiria, 25 Russia.: Paleontologichesky Zhurnal, no. 2, p. 29. 26 Baryshnikov, V.V., Zolotova, V.P., and Kosheleva, V.F., 1982, New foraminifera species of Artinskian 27 Stage of Permian Preurals. Preprint.: Sverdlovsk (In Russian). 28 Belyaev, g.M., and Rauser-Chernousova, D.M.R.-C., 1938, On some fusulinids of the Schwagerina horizon 29 (the group of Pseudofusulina uralica Krotow): Acad. Sci. URSS, Inst. Geol., Tr., v. 7, p. 169–196. 30 Bensh, F.R., Zonal'naya stratigrafiya i korrelyatsiya verkhnego karbona i nizhney permi Sredney Azii. 31 Zonal stratigraphy and correlation of the Upper Carboniferous and Lower Permian in Central Asia. 32 Bensh, F.R., 1962, Pozdnekamennougol'nye i rannepermskie fuzulinidy Severnoi Fergany. Late 33 Carboniferous and early Permian fusulinids of Northern Fergana., in Verkhov, V.I., ed., Stratigraphy 34 and paleontology of Uzbekistan and surrounding areas. Book 1: Tashkent, Akademy of Sciences of 35 Uzbekistan, p. 186. 36 Bensh, F.R., editor, 1965, Stratigrafiya i foraminifery kamennougol'nykh otlozheniy yugo-zapadnykh 37 otrogov i yuzhnogo sklona Gissarskogo khrebta. Carboniferous stratigraphy and foraminifera of the 38 southwestern spurs and southern slope of the Hissar Range: Tashkent, Fan. 39 Bensh, F.R., 1968, Stratigrafiya morskikh fatsiy verkhnego karbona Tyan'-Shanya. Stratigraphy of marine 40 facies in the upper Carboniferous of the Tien Shan: Mosk. Obshchest. Ispyt. Prir., Byull., Otd. Geol., v. 41 43, no. 3, p. 59–66. 42 Bensh, F.R., 1969, Stratigrafiya i foraminifery kamennougol'nykh otlozheniy yugo-zapadnykh otrogov i 43 yuzhnogo sklona Gissarskogo khrebta. Carboniferous stratigraphy and foraminifera of the 44 southwestern spurs and southern slope of the Hissar Range: Tashkent, Fan Publishing House, 45 (None). 46 Bensh, F.R., 1972, Stratigrafiya i fuzulinidy verkhnego paleozoya Yuzhnoy Fergany. Stratigraphy and 47 fusulinids of the upper Paleozoic of southern Fergana: Tashkent, Fan, Rauser-Chernousova, D. M., 1- 48 149, 31 paleontol. plates. 49 Bensh, F.R., 1978, Raschleneniye verkhnego karbona i granitsa karbona i permi po materialam Sredney 50 Azii. Subdividing the Upper Carboniferous and the Carboniferous-Permian boundary based on 51 materials from Central Asia: Trudy - Mezhvedomstvennyy Stratigraficheskiy Komitet SSSR, v. 6, p. 52 247–250. 53 Bensh, F.R., 1982, Fuzulinidovye zony i yarusnoe raschlenenie verkhbego karbona i nizhnei permi 54 Srednei Azii. Fusulinid Zonation stadial subdivisions of Upper Carboniferous and Lower Permian of 55 Central Asia: Tashkent, Fan, (None), 1-272 (IN Russian). 56 Bensh, F.R., 1987, Reviziya sistematiki psevdofuzulinid, roda Pseudofusulina Dunbar et Skinner, 1931 i 57 blizkikh rodov. Taxonomic revision of pseudofusulinids, Pseudofusulina Dunbar and Skinner, 1931 58 and similar taxa: Voprosy Mikropaleontologii, v. 29, p. 20. 59 Bensh, F.R., Rumiantseva, Z.S., and Sergunkova, O.I., 1996, The former USSR; Tianshan and Pamirs 60 [modified]: Publication - International Union of Geological Sciences, v. 33, p. 122–153. 61 Bogush, O.I., 1963, Foraminifers and stratigraphy of middle and upper Carboniferous the Eastern part pf 62 Alay Ridge: Novosibirsk, Nauka [In Russian]. 63 Bogush, O.I., and Yuferev, O.V., 1962, Foraminifery i stratigrafiya kamennougolnykh otlozhenii Karatau i 64 Talasskogo Alatau, 234 p. 65 Brazhnikova, N.Y.'y., Vakarchuk, G.I., Vdovenko, M.V., Vinnichenko, L.V., Karpova, M.A., Kolomiyets, Y.I., 66 Potiyevskaya, P.D., Rostovtseva, L.F., and Shevchenko, G.D., 1967, Microfaunal marker horizons in 67 the Carboniferous and Permian deposits of the Dnieper-Donets basin, 194 p. 68 Bykadorov, V.A., Belov, A.A., Bensh, F.R., Volozh, Y.A., Volchegurskiy, L.F., Korzhayev, V.P., Koshkin, V.P., 69 Pilifosov, V.M., Puchkov, V.N., Smirnov, A.V., Smirnov, L.V., Surkov, V.S., Fedorenko, O.A., 70 Shokal'skiy, S.P., Yan, Y., Yatskevich, S.V., Ali-Zade, A., Brodskiy, A.Y., Bryzgalov, S.L., Dzhanseitov, 71 N.S., Getman, O.F., Zamarenov, A.K., Karapetov, S.S., Kim, Y.I., Krasnoborodkin, V.K., Mavyyev, N.C., 72 Myasnikov, A.K., Nikitin, Y.I., Poyezzhayev, I.P., Rustamov, M.I., Sapozhnikov, R.B., Sergeyeva, L.V., 73 Skrinnik, L.I., Smirnova, L.G., Tevelev, A.V., Titova, A.P., Tokarev, V.N., Turkov, O.S., Filin, V.I., 74 Filippova, I.B., and Miletenko, N.V., Early Permian--Rannyaya perm'. 75 Chang, L.S., 1963, Upper Carboniferous fusulinids from Kelpin and its neighbourhood, Hsin-Kiang. Part. 76 1: Acta Pelaeontologica Sinica., v. 11, no. 1, p. 36. 77 Chuvashov, B.I., Fuzulinidy zony Parafusulina solidissima na Urale. The fusulinid zones of Parafusulina 78 solidissima in the Urals. 79 Chuvashov, B.I., Permian deposits of the Urals and Preduralye. 80 Chuvashov, B.I., 1972, Verkhniy karbon zapadnogo sklona Srednego Urala. Upper Carboniferous of the 81 western slope of the Central Urals: Sovetskaya Geologiya, v. 11, p. 106–119. 82 Chuvashov, B.I., 1997, Kungurskiy yarus permskoy sistemy (problemy vydeleniya i korrelyatsii). 83 Kungurian stage of the Permian System; problems of recognition and correlation: Stratigrafiya, 84 Geologicheskaya Korrelyatsiya, v. 5, no. 3, p. 10–28. 85 Chuvashov, B.I., editor, 2002, Guidebook of geological excursions on the Carboniferous of the Urals. Part 86 1. S. Urals excursion.: Ekaterinburg, Institute of Geology and Geochemistry of UB RAS. 87 Chuvashov, B.I., Alksne, A.E., and Polozova, A.N., Zonal'naya stratigrafiya artinskogo yarusa Zapadnogo 88 Urala i Predural'ya (po fuzulinidam). The regional stratigraphy of the Artinskian Stage in the western 89 Urals and the Uralian Foreland, according to fusulinids. 90 Chuvashov, B.I., and Chernykh, V.V., 2000, The Kungurian Stage in the general stratigraphic scale of the 91 Permian System: Doklady Earth Sciences, 375A, p. 1345–1349. 92 Chuvashov, B.I., Chernykh, V.V., and Bogoslovskaya, M.F., 2002, Biostratigraficheskaya kharakteristika 93 stratotipov yarusov nizhney permi. Biostratigraphic characteristic of Lower Permian stage 94 stratotypes of Permian: Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 10, no. 4, p. 3–19. 95 Chuvashov, B.I., Chernykh, V.V., Leven, E.Y., Davydov, V.I., Bowring, S.A., Ramezani, J., Gelnister, B.F., 96 Henderson, C.M., Schiappa, T.A., Northrup, C.J., Snyder, W.S., Spinosa, C., and Wardlaw, B.R., 2002, 97 Proposal for the Base of the Sakmarian Stage: GSSP in the Kondurovsky Section, southern Urals, 98 Russia: Permophiles, v. 41, p. 4–13.

99 Chuvashov, B.I., Djupina, G.V., Mizens, G.A., and Chernykh, V.V., 1990, The Key-sections of Upper 100 Carboniferous and Lower Permian in the western slope of Urals and Preurals.: Ekaterinburg, Institute 101 of Geology and geochemistry of Uralian Branch of Russian Academy of Sciences, (None), 368 p. 102 Chuvashov, B.I., and Dyupina, G.V., 1971, Verkhniy paleozoy rayona poselka Biserti. The upper Paleozoic 103 of the Bisert region: Akademiya Nauk SSSR, Ural'skiy Filial, Institut Geologii i Geokhimii, Trudy, v. 88, 104 p. 3–24. 105 Chuvashov, B.I., and Dyupina, G.V., 1978, Faunisticheskiye kompleksy i problemy korrelyatsii razno 106 fatsial'nykh otlozheniy (na primere assel'skogo yarusa zapadnogo sklona Srednogo Urala). Faunal 107 complexes and problems of the correlation of diverse facies deposits; the Asselian Stage of the 108 western slopes of the Central Urals as an example: Trudy Instituta Geologii i Geofiziki, v. 386, p. 124– 109 146. 110 Chuvashov, B.I., and Ivanova, R.M., Moskovskiye i verkhnekamennougol'nyye otlozheniya v razreze "Uly- 111 Taldyk" (Vostochnyye Mugodzhary). Moscovian and Upper Carboniferous deposits of the "Uly- 112 Taldyk" Section, eastern Mugodzhar Hills. 113 Chuvashov, B.I., and Ivanova, R.M., Sredniy karbon rek Kunary i Iseti. Middle Carboniferous of the 114 Kunara and Iset' rivers. 115 Chuvashov, B.I., and Ivanova, R.M., 1980, Middle Carboniferous of the Kunara and Iset' rivers: 116 Carboniferous stratigraphy, Fusulinidae, and miospores of the Urals. 117 Chuvashov, B.I., and Ivanova, R.M., 1980, Moscovian and Upper Carboniferous deposits of the "Uly- 118 Taldyk" Section, eastern Mugodzhar Hills: Carboniferous stratigraphy, Fusulinidae, and miospores of 119 the Urals. 120 Chuvashov, B.I., Ivanova, R.M., and Kolchina, A.N., 1979, Verkhniy paleozoy basseyna r. Sinary. The 121 upper Paleozoic basin of the Sinara River: Trudy Instituta Geologii i Geokhimii, no, no. 26, p. 95–125. 122 Chuvashov, B.I., Ivanova, R.M., and Kolchina, A.N., editors, 1984, Verkhniy paleozoy vostochnogo sklona 123 Urala; stratigrafiya i geologicheskaya istoriya. Upper Paleozoic deposits of the eastern slope of the 124 Urals; stratigraphy and geologic history: Sverdlovsk, UO AN of USSR, Papulov, G.N., 230 p. 125 Chuvashov, B.I., Leven, E.Ya.and Davydov, V.I., editor, 1986, Carboniferous/Permian Boundary beds of 126 the Urals, Pre-Urals area and Central Asia.: Moscow, Nauka Publishing House. 127 Davydov, V.I., 1981, New species of middle Carboniferous Palaeorechelina.: Paleontological Journal, no. 128 3, 120-124 (In Russian). 129 Davydov, V.I., 1984, Zonal'nye i yarusnye podrazdeleiya verkhnego karbona yu-go-vostochnogo Darvaza. 130 Fusulinid zonal subdivisions of the Upper Carboniferous of the southwestern Darvas: Bulletin of 131 Moscow Society of Natural Studies, Geological Series, v. 59, no. 3, p. 41. 132 Davydov, V.I., 1986, Fusulinids of Carboniferous-Permian boundary beds of Darvas, in B.I. Chuvashov, 133 E.Ya. Leven, and V.I. Davydov, ed., Carboniferous/Permian Boundary beds of the Urals, Pre-Urals 134 area and Central Asia.: Moscow, Nauka Publishing House, p. 103. 135 Davydov, V.I., 1986, Precise definition of the Carboniferous-Permian boundary in Donets Basin and the 136 Northern Caucasus based on paleomagnetic criteria: Soviet Geology, v. 12, p. 74–76. 137 Davydov, V.I., 1986, Upper Carboniferous and Asselian fusulinids of the Southern Urals, in Chuvashov, 138 E.Ya. Leven and V.I. Davydov, ed., Carboniferous--Permian Boundary beds of the Urals, Pre-Urals and 139 Central Asia: Moscow, Nauka Publishing House, p. 77. 140 Davydov, V.I., 1987, Fusulinid zone DAIXINA BOSBYTAUENSIS - D. ROBUSTA in South Ferghana.: Doklady 141 Akademii Nauk SSSR, v. 292, no. 1, 160-164 (In Russian). 142 Davydov, V.I., 1988, Archaediscids in the Upper Carboniferous and Lower Permian: Revue de 143 Paleobiologie, spec. v. 2, p. 39–46. 144 Davydov, V.I., 1988, The origin and evolution some "pseudofusulinids": Paleontologichesky Zhurnal, no. 145 3, p. 10.

146 Davydov, V.I., 1990, Zonal fusilinid subdivisions of Gzhelian in Donets Basin and Pre-Donets Trough, in 147 Problems of Modern Micropaleontology: Leningrad, Nauka. Proceedings of the All-Union 148 Paleontological Society, p. 52–69. 149 Davydov, V.I., 1991, Vozrast verkhney chasti formatsii Belcher Chenell v stratotipe po fuzulinidam. The 150 age of the upper Belcher Channel Formation in the type-section on the ground of fusulinids, in 151 Bondarev, V.I., [No last name!], eds.: Stratigraphy and Paleontology of Paleozoic of the Arctic. 152 Transactions of SEVMORGEO, Nauchn.-Proizvod. Ob"yedin. Sevmorgeologiya, Leningrad, p. 16. 153 Davydov, V.I., 1992, Subdivision and correlation of Upper Carboniferous and Lower Permian deposits in 154 Donets Basin according to fusulinid data: Soviet Geology, no. 5, p. 53. 155 Davydov, V.I., 1995, The Carboniferous/Permian Boundary in South China: Permophiles, no. 26, p. 9–11. 156 Davydov, V.I., 1997, Fusulinid biostratigraphy of the Upper Paleozoic of the Kolguev Island and Franz 157 Josef Land Archipelago: Biostratigraphy of the oil-bearing basins., v. 1, 40-59 (In Russian). 158 Davydov, V.I., 1998, Fusulinid biostratigraphic dating of the Upper Paleozoic successions in wells 7128/4- 159 1, 7228/9-1 and 7124/3-1, Finnmark East Licence area, Barents Sea shelf., in Unpublished Permian 160 Research Institute report, p. 31. 161 Davydov, V.I., 2011, Taxonomy, nomenclature and evolution of the early schubertellids (Fusulinida, 162 Foraminifera) Acta Palaeontologica Polonica., v. 56, no. 1, p. 181–194. 163 Davydov, V.I., Anisimov, R.M., and Nilsson, I., 1999, Upper Paleozoic Sequence stratigraphy of the Arctic 164 Region implied from Graphic Correlation: Spitsbergen and the Barents Sea shelf, in Yin, Hongfu & 165 Tong, Jinnan (eds.), ed., Pangea and the Paleozoic-Mesozoic transition.: Hubei, China University 166 Geosci. Press, p. 94–97. 167 Davydov, V.I., and Arefifard, S., 2007, Permian Fusulinid Fauna of Gondwanan Affinity from Kalmard 168 Region, East-Central Iran and its Significance for the Tectonics and Paleogeography: Paleontologia 169 electronica, http://palaeo-electronica.org/2007_2/00124/index.html, v. 10, p. 40. 170 Davydov, V.I., Barskov, I.S., Bogoslovkaya, M.F., Popov, A.V., Akhmetshina, L.Z., and Kozitskaya, R.I., 171 1992, The Carboniferous-Permian boundary in the former USSR and its correlation: International 172 Geology Review, v. 34, no. 9, p. 889–906. 173 Davydov, V.I., Barskov, I.S., Bogoslovskaya, M.F., and Leven, E.Y., 1994, Granitsa karbona i permi v 174 stratotipicheskikh razrezakh Yuzhnogo Urala i yeye korrelyatsiya. Carboniferous-Permian boundary 175 in stratotype sections of the Southern Urals and its correlation: Stratigrafiya, Geologicheskaya 176 Korrelyatsiya, v. 2, no. 3, p. 32–45. 177 Davydov, V.I., Barskov, I.S., Bogoslovskaya, M.F., Leven, E.Y.A., Popov, A.V., Akhmetshina, L.Z., and 178 Kozitskaya, R.I., 1992, The Carboniferous-Permian boundary in the former USSR and its correlation: 179 International Geology Review, v. 34, no. 9, p. 889–906. 180 Davydov, V.I., Bogoslovskaya, M.F., Chernykh, V.V., Chuvashov, B.I., Leonova, T.B., Schiappa, T.A., 181 Spinosa, C., and Wardlaw, B.R., Lower Permian Cisuralian series; biostratigraphic characteristic in the 182 type area and surrounding regions. 183 Davydov, V.I., Chernykh, V.V., Chuvashov, B.I., D, S.M., and Snyder, W.S., 2008, Faunal assemblage and 184 correlation of Kasimovian-Gzhelian transition at Usolka section, southern Urals, Russia (a potential 185 candidate for GSSP to define base of Gzhelian Stage): Stratigraphy, v. 5, no. 2, p. 113–135. 186 Davydov, V.I., Crowley, J.L., and Schmitz, M.D., 2010, High-precision U-Pb zircon age calibration of the 187 global Carboniferous time scale and Milankovitch band cyclicity in the Donets Basin, eastern 188 Ukraine: Geochemistry, Geophysics, Geosystems: G3, v. 11, no. 1, p. 1–22, doi: 189 10.1029/2009GC002736. 190 Davydov, V.I., Haig, D.W., and Mccartain, E., 2013, A latest Carboniferous warming spike recorded by a 191 fusulinid-rich bioherm in Timor Leste: Implications for East Gondwana deglaciation: 192 Palaeogeography, Palaeocimatology and Palaeogeography, v. 376, p. 22–38, doi: 193 10.1016/j.palaeo.2013.01.022.

194 Davydov, V.I., and Henderson, C.M., 2007, The Cisuralian GSSP Field Workshop: Permophiles, v. 49, p. 4– 195 6. 196 Davydov, V.I., and Kozur, H., 1997, Position of the Carboniferous/Permian Boundary in the Carnic Alps 197 compared with the stratotype region, in M. Podemski, S. Dybova-Jachowicz, J. Jureczka and R. 198 Wagner (eds.), ed., Proceeding of the XIII International Congress on the Carboniferous and Permian: 199 Warszawa, p. 123–130. 200 Davydov, V.I., and Krainer, K., 1999, Fusulinid assemblages and Facies of the Bombaso Fm. andbasal 201 Meledis Fm. (Moscovian-Kasimovian) in the Central Carnic Alps (Austria/Italy): Facies, v. 40, p. 157– 202 196. 203 Davydov, V.I., Krainer, K., and Chernykh, V.V., 2013, Fusulinid biostratigraphy of the Lower Permian 204 Zweikofel Formation (Rattendorf Group; Carnic Alps, Austria) and Lower Permian Tethyan 205 chronostratigraphy: Geological Journal, v. 48, p. 57–100. 206 Davydov, V.I., Krainer, K., and Fritz, A., 1997, Upper Carboniferous and Lower Permian 207 continental/marine correlation based on Carnic Alps and Donets basin sections. 208 Davydov, V.I., and Leven, E.J., 2003, Correlation of Upper Carboniferous (Pennsylvanian) and Lower 209 Permian (Cisuralian) marine deposits of the Peri-Tethys: Palaeogeography, Palaeoclimatology, 210 Palaeoecology, v. 196, 1-2, p. 39–57. 211 Davydov, V.I., and Leven, E.Y., 1999, Fusulinid biostratigraphy at the Asselian-Sakmarian transition in 212 Kondurovsky Section: Permophiles, v. 34, p. 23. 213 Davydov, V.I., and Nilsson, I., 1999, Fusulinids in the Middle Upper Carboniferous Boundary Beds on 214 Spitsbergen, Arctic Norway: Palaeontologia Electronica, http://paleo-electronica.org, v. 2,, no. 1, p. 215 45. 216 Davydov, V.I., Nilsson, I., and Stemmerik, L., 2001, Fusulinid zonation of the Upper Carboniferous Kap 217 Jungersen and Foldedal Formations, southern Amdrup Land, eastern north Greenland: Bulletin of 218 the Geological Society of Denmark, v. 48, no. 1, p. 31–77. 219 Davydov, V.I., and Popov, A.V., 1986, Upper Carboniferous and Lower Permian sections of the southern 220 Urals, in Chuvashov, B.I., Leven, E.Ya.and Davydov, V.I., ed., Carboniferous/Permian Boundary beds 221 of the Urals, Pre-Urals area and Central Asia.: Moscow, Nauka Publishing House, p. 29. 222 Davydov, V.I., Schiappa, T.A., and Snyder, W.S., 2003, Testing the Sequence Stratigraphy Model: 223 Response of Fusulinacean Fauna to Sea Level Fluctuations (examples from Pennsylvanian and 224 Cisuralian of Pre-Caspian-southern Urals Region, in Olson, H. C., and R. M. Leckie, ed., 225 Micropaleontologic Proxies for Sea-Level Changes and Stratigraphic Discontinuities. SEPM Special 226 Publication: SEPM, SEPM (Society for Sedimentary Geology), Volume 75, p. 359–375. 227 Davydov, V.I., Snyder, W.S., and Spinosa, C., 1997, Fusulinacean biostratigraphy and sequence 228 stratigraphy of the upper Paleozoic of the Southern Urals: Special Publications - Cushman 229 Foundation for Foraminiferal Research, v. 36, p. 27–30. 230 Dutkevich, G.A., 1934, Some new species of Fusulinidae from the upper and middle Carboniferous of 231 Verkhne-Chussovskye Gorodki on the Chussovaya river (western slope of the middle Ural), v. 36, p. 232 58–90. 233 Dzhenchuraeva, A.V., 1993, Stratigraphy and foraminifera of the Upper Paleozoic at the axial part of the 234 Turkestan-Alai.: Bishkek, Ilim Publishing House. 235 Dzhenchuraeva, A.V., and Getman, O.F., 2007, Stratigraphy and foraminifers from the uppermost 236 Carboniferous (Kasimovian-Gzhelian) of the Jamantoo and Baibichetoo Ranges (middle Tien-Shan, 237 Kyrgyzstan): Journal of Foraminiferal Research, v. 37, no. 1, p. 46–68, doi: 10.2113/gsjfr.37.1.46. 238 Dzhenchuraeva, A.V., and Getman, O.F., 2010, Stratigraphy and lithology of Upper Paleozoic of Middle 239 Tian-Shan': Bishkek, Ministry of Natural Resources of Kirgiz Republic. 240 Dzhenchuraeva, A.V., Neevin, A.V., Masumov, R.A., Getman, O.F., and Nogaeva, L.P., 2013, Atlas of 241 facies and fossil remains of Paleozoic of Central Tian-Shan'.: Bishkek, Kyrgiz-Slavyansk University, 403 242 p. (In Russian).

243 Dzhenchuraeva, A.V., and Okuyucu, C., 2007, Fusulinid Foraminifera of the Bashkirian-Moscovian 244 boundary in the eastern Taurides, southern Turkey: Journal of Micropalaeontology, 26, Part 1, p. 73– 245 85, doi: 10.1144/0262-821X05-006. 246 Ehrenberg, S.N., Nielsen, E.B., Svana, T.A., and Stemmerik, L., 1998, Depositional evolution of the 247 Finnmark carbonate platform, Barents Sea; results from wells 7128/6-1 and 7128/4-1: Norsk 248 Geologisk Tidsskrift, v. 78, no. 3, p. 185–224. 249 Ehrenberg, S.N., Pickard, N.A.H., Svana, T.A., Nilsson, I., and Davydov, V.I., 2000, Sequence stratigraphy 250 of the inner Finnmark carbonate platform, Barents Sea: correlation between well 7128/6-1 and the 251 shallow IKU wells: Norsk Geologisk Tidsskrift, v. 80, p. 129–162. 252 Ekhlakov, Y.A., and Zolotova, V.P., 1986, Carboniferous-Permian boundary beds on Kos'va and 253 Berezovaya Rivers, in Chuvashov, B.I., Leven, E. Y. and Davydov, V.I., ed., Carboniferous--Permian 254 Boundary beds of the Urals, Pre-Urals and Central Asia. Nauka Publishing House, Moscow. (In 255 Russian): Moscow, Nauka, 12-18 (In Russian). 256 Filimonova, T.V., Gorozhanina, E.N., Isakova, T.N., and Gorozhanin, V.M., 2015, Cisuralian Series of the 257 Permian System of the Southeastern SolIletsk Swell: Biostratigraphy and Lithology: Stratigrafiya, 258 Geologicheskaya Korrelyatsiya, v. 23, no. 2, p. 131–154. 259 Fluegel, E., Fohrer, B., Forke, H., Krainer, K., and Samankassou, E., 1997, Excursion B1; Cyclic sediments 260 and algal mounds in the upper Paleozoic of the Carnic Alps: Gaea Heidelbergensis, v. 4, p. 79–100. 261 Fluegel, E., and Kahler, F., 1974, Fazies-Interpretation der unterpermischen Sedimente in den Karnischen 262 Alpen. Facies interpretation in the Lower Permian sediments of the Carnic Alps: Carinthia II, v. 164, 263 p. 43–62. 264 Forke, H., and Kahler, F., 1997, Franz Kahler (1900-1995); Bemerkungen und Wuensche mit einem 265 optimistischen Ausblick; Letzte Gedanken; schriftlich niedergelegt im Juli 1995. Franz Kahler (1900- 266 1995); observations and desires with an optimistic outlook; final thoughts written in July 1995: 267 Carinthia II, v. 107, no. 1, p. 25–28. 268 Forke, H.C., 1995, Biostratigraphie (Fusuliniden; Conodonten) und Mikrofazies im Unterperm (Sakmar) 269 der Karnischen Alpen (Nassfeldgebiet, Oesterreich). Biostratigraphy (fusulinids, conodonts) and 270 microfacies in the Lower Permian (Sakmarian) of the Carnic Alps, Nassfeld area, Austria: Jahrbuch 271 der Geologischen Bundesanstalt Wien, v. 138, no. 2, p. 207–297. 272 Forke, H.C., 2001, Integrated paleontological studies of the fusulinacea/conodont faunas and 273 biostratigraphy of Upper Carboniferous /Lower Permian deposits from the Southern Alps (Carnic 274 Alps, Karavanke Mts., Austria,/Italy/ Slovenia) and its correlation with Russian type sections 275 (Moscow Basin, Southern Urals).: Erlangen, University Erlangen-Nurnberg, Unpiublished PhD Thesis. 276 Forke, H.C., 2002, Biostratigraphic subdivision and correlation of uppermost Carboniferous/Lower 277 Permian sediments in the southern Alps; fusulinoidean and conodont faunas from the Carnic Alps 278 (Austria/Italy), Karavanke Mountains (Slovenia), and Southern Urals (Russia): Facies, v. 47, p. 201– 279 275. 280 Forke, H.C., Kahler, F., and Krainer, K., 1998, Sedimentology, microfacies and stratigraphic distribution of 281 foraminifers of the Lower "Pseudoschwagerina" Limestone (Rattendorf Group, Late Carboniferous), 282 Carnic Alps (Austria/Italy): Senckenbergiana Lethaea, v. 78, 1-2, p. 1–39. 283 Forke, H.C., and Samankassou, E., 2000, Biostratigraphical correlation of late Carboniferous (Kasimovian) 284 section in the Carnic Alps (Austria/Italy); integrated paleontological data, facies, and discussion: 285 Facies, v. 42, p. 177–210. 286 Forke, H.C., Schonlaub, H.-P., and Samankassou, E., 2006, Late Paleozoic of the Carnic Alps 287 (Austria/Italy). Field Guide-book of the SCCS Task Group to establish GSSP's close to the 288 Moscovian/Kasimovian and Kasimovian/Gzhelian boundaries. July 31-August 1, 2006: Berichte der 289 Geologischen Bundesanstalt, no. 70, p. 57.

290 Goreva, N.V., Alekseev, A.S., Isakova, T.N., and Kossovaya, O.L., 2007, Afanasievo section - neostratotype 291 of Kasimovian Stage (Upper Pennsylvanian Series), Moscow Basin, central Russia: Newsletter on 292 Carboniferous Stratigraphy, v. 25, p. 8–14. 293 Goreva, N.V., Alekseev, A.S., Isakova, T.N., and Kossovaya, O.L., 2009, Biostratigraphical analysis of the 294 Moscovian-Kasimovian transition at the neostratotype of Kasimovian Stage (Afanasievo section, 295 Moscow Basin, Russia): Palaeoworld, v. 18, p. 102. 296 Goreva, N.V., Alekseyev, A.S., Isakova, T.N., and Kossovaya, O.L., 2009, "Podtrititsitovyye" sloi Donskoy 297 Luki i nizhnyaya granitsa kasimovskogo yarusa. "Sub-Triticites" beds in the Don River basin and the 298 Kasimovian's lower boundary: Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 17, no. 1, p. 49–69. 299 Gorgij, M.N., and Leven, E., 2013, The first findings of fusulinids in the sections of the Sabzevar tectonic 300 block (Iran): Stratigraphy and Geological Correlation, v. 21, no. 1, p. 8–17. 301 Gorsky, V.P., and Kalmykova, M.A., editors, 1986, Atlas of characteristic assemblages of Permian fauna 302 and flora of the Urals and Russian Platform.: Leningrad, Nedra Publishing House, Transactions of 303 VSEGEI, new series, 331. 304 Grozdilova, L.P., 1937, Fusulinids from the vicinity of the Simsk works in the southern Urals: Russia, Tp. 305 [Petroleum Geol.-Prosp. Inst., Tr.] s. A, v. 106, p. 24–41. 306 Grozdilova, L.P., 1938, Fuzulinidy neftenosnykh izvestnyakov Ishimbayevskogo rayona: [Petroleum Geol.- 307 Prosp. Inst., Tr.] s. A, v. 101, p. 90–141. 308 Grozdilova, L.P., 1966, Foraminifera of the upper Carboniferous of the northern Timan range: Trudy 309 Vsesoyuznogo Neftyanogo Nauchno-Issledovatel'skogo Geologorazvedochnogo Instituta, v. 250, p. 310 254–331. 311 Grozdilova, L.P., 1966, Foraminifery verkhnego karbona severnogo Timana. Foraminifera of the upper 312 Carboniferous of the northern Timan range, in Trudy Vsesoyuznogo Neftyanogo Nauchno- 313 Issledovatel'skogo Geologorazvedochnogo Instituta: Leningrad, Nedra, p. 254–331. 314 Grozdilova, L.P., Lebedeva, I.S., Lipina, O.A., Malakhova, N.P., Mikhaylova, Z.P., Chermnykh, V.A., 315 Postoyalko, M.V., Simonova, Z.G., Sinitsyna, Z.A., and Shcherbakova, M.V., 1975, Foraminifera: Trudy 316 Vsesoyuznogo Neftyanogo Nauchno-Issledovatel'skogo Geologorazvedochnogo Instituta, no. 383, p. 317 27–64. 318 Grozdilova, L.P., and Lebedeva, N.S., 1960, Foraminifery kamennougolnykh otlozhenii zapadnogo sklona 319 Urala i Timana; atlas naibolee kharakternykh vidov: Trudy - Vsesouznyy Naucno-Issledovatelskiy 320 Geologorazvedocnyy Neftanoy Institut, Geohimiceskiy Sbornik, v. 150, p. 264. 321 Grozdilova, L.P., and Lebedeva, N.S., 1961, Lower Permian foraminifers of North Timan, in anonimmous, 322 ed., Transactions of VNIGRI, p. 161. 323 Grozdilova, L.P., and Lebedeva, N.S., 1965, Paleontologicheskiy Zhurnal: Paleontologicheskiy Zhurnal, v. 324 1, p. 146. 325 Grozdilova, L.P., Lebedeva, N.S., Yermakova, K.A., Chudinova, I.I., Simakova, M.A., Degtyarev, D.D., 326 Rakshin, P.P., Muromtseva, V.A., Tkacheva, I.D., Kruchinina, O.N., Lapina, N.N., and Stanichnikova, 327 M.S., 1978, Paleontological characteristics: Leningrad, USSR (SUN), Izd. Nedra, Leningrad, (None). 328 Grozdilova, L.P., Simakova, M.A., Kruchinina, O.N., and Sosipatrova, G.P., 1980, The paleobiogeography 329 of the Early Permian northern Timan Basin and its possible relationship to the water areas of Novaya 330 Zemlya and Spitsbergen: Sverdlovsk, USSR (SUN), Akad. Nauk SSSR, Ural'sk. Nauchn. Tsentr, 331 Sverdlovsk, The biostratigraphy of the Artinskian and Kungurian stages in the Urals. 332 Gusev, A.K., Bogatyrev, V.V., Igonin, V.M., and Solodukho, M.G., 1968, Stratigraphy of Upper Paleozoic 333 deposits of Aktobe Preurals.: Kazan, Kazan University, 219 p. (In Russian). 334 Guseva, Y.A., Grozdilova, L.P., and Gorskiy, V.P., 1968, Biostratigraphic basis for the boundary between 335 the Artinskian and Kungurian suites of the Urals: Doklady Akademii Nauk SSSR, v. 182, no. 4, p. 893– 336 895. 337 Guseva, Y.A., Grozdilova, L.P., and Gorskiy, V.P., 1968, Biostratigraphic substantiation of the Artinskian- 338 Kungurian boundary in the Urals: Acad. Sci. USSR, Dokl., Earth Sci. Sect., v. 182, p. 73–74.

339 Il'in, V.D., Gubareva, V.S., Zamilatskaya, T.K., and Klenina, L.N., 1987, Subsalt sedimentary assemblage of 340 Karachaganak oil-field: Stratigraphy and Paleontology of Paleozoic of Precaspian Basin, 5-25 (In 341 Russian). 342 Isakova, T.N., Alekseev, A.S., Goreva, N.V., and Kossovaya, O.L., 2005, Lower Kasimovian (Pennsylvanian) 343 of Donskaya Luka (southern Russia): Newsletter on Carboniferous Stratigraphy, v. 23, p. 44–47. 344 Isakova, T.N., and Nazarov, V.B., 1986, The Late Carboniferous-Early Permian stratigraphy and 345 microfauna of South Urals.: Moscow, Nauka [In Russian]. 346 Izotova, M.N., 1985, Fusulinid zones of Lower Permian of northern and eastern slopes of Precaspian 347 depression.: Soviet Geology, no. 9, p. 76. 348 Izotova, M.N., and Goryacheva, L.P., 1980, Early Permian fusulinids of northern margin of Precaspian 349 Basin;: Microfauna i biostratigraphiya neftegazonosnykh raionov SSSP, v. 1, 61-70 (In Russian). 350 Izotova, M.N., and Vevel, Y.A., 1998, New species of fusulinoids from the eastern Caspian Sea region: 351 Paleontological Journal, v. 32, no. 4, p. 332–335. 352 Kahler, F., 1939, Verbreitung und Lebensdauer der Fusuliniden- Gattungen Pseudoschwagerina und 353 Paraschwagerina und deren Bedeutung fuer die Grenze Karbon/Perm: Senckenbergiana, v. 21, 3-4, 354 p. 169–215. 355 Kahler, F., 1943, Der Trogkofelkalk; ein vergessener Kaerntner Marmor: Carinthia II, pp. 356 Kahler, F., 1953, Der Bau der Karawanken und des Klagenfurter Beckens: Carinthia II, v. 78. 357 Kahler, F., 1955, Entwicklungsraeume und Wanderwege der Fusuliniden am eurasiatischen Kontinent: 358 Geologie, v. 2, p. 178–188. 359 Kahler, F., 1970, Marines Unterperm in der Basis der Koschuta (suedliche Karawankenkette, Kaernten). 360 The marine lower Permian at the base of the Koschuta, southern Karawanken Range, Carinthia: 361 Anzeiger der Oesterreichischen Akademie der Wissenschaften, Mathematisch- 362 Naturwissenschaftliche Klasse, v. 106, 1-14, p. 12–13. 363 Kahler, F., 1972, Das Perm der Karnischen Alpen. The Permian of the Carnic Alps: Verhandlungen der 364 Geologischen Bundesanstalt (Wien), v. 1, p. 139–141. 365 Kahler, F., 1973, Beitraege zur Kenntnis der Fusuliniden der Ostalpen; die Gattung Quasifusulina in den 366 Karnischen Alpen. Contributions to knowledge of the fusulinids of the eastern Alps; the genus 367 Quasifusulina in the Carnic Alps: Palaeontographica. Abteilung A: Palaeozoologie-Stratigraphie, 141, 368 Part 5-6, p. 154–173. 369 Kahler, F., 1974, Iranische Fusuliniden. Iranian fusulinids: Jahrbuch der Geologischen Bundesanstalt 370 Wien, v. 117, p. 75–107. 371 Kahler, F., 1984, Ein Vergleich der Fusuliniden-Fauna des Oberkarbon and Unterperm der Ostalpen mit 372 dem Dongebiet (UdSSR). A comparison of the fusulinid fauna of the Upper Carboniferous and Lower 373 Permian of the Eastern Alps with the Don region, USSR: Mitteilungen der Oesterreichischen 374 Geologischen Gesellschaft, v. 77, p. 247–261. 375 Kahler, F., 1987, Fusuliniden-Faunen auf Chios, Kalymnos und Kos in der Aegaeis: Mitteilungen der 376 Oesterreichischen Geologischen Gesellschaft, v. 80, p. 287–323. 377 Kahler, F., 1991, Beziehungen der Fusuliniden der Karnischen Alpen zur Palaeotethys. Relationships of 378 fusulinids of the Carithian Alps to the Paleo-Tethys: Mitteilungen der Oesterreichischen 379 Geologischen Gesellschaft, v. 84, p. 309–326. 380 Kahler, F., and Kahler, G., 1937, Beitraege zur Kenntnis der Fusuliniden der Ostalpen; Die 381 Pseudoschwagerinen der Grenzlandbaenke und des oberen Schwagerinenkalkes: Palaeontographica. 382 Abteilung A: Palaeozoologie-Stratigraphie, v. 87, p. 1–44. 383 Kahler, F., and Kahler, G., 1938, Beobachtungen an Fusuliniden der karnischen Alpen: Zentr. Miner. Abt. 384 B, v. 4, p. 101–115. 385 Kahler, F., and Kahler, G., 1940, Fusuliniden aus dem Tienschan, v. 2.

386 Kahler, F., and Kahler, G., 1941, Beitraege zur Kenntnis der Fusuliniden der Ostalpen; die Gattungen 387 Pseudoschwagerina und ihre Vertreter im unteren Schwagerinenkalk und im Trogkofelkalk: 388 Palaeontographica. Abteilung A: Palaeozoologie-Stratigraphie, v. 92, p. 59–98. 389 Kahler, F., and Kahler, G., 1969, Einige permische Fusuliniden aus dem Irak. Permian fusulinids from Iraq: 390 Neues Jahrbuch fuer Geologie und Palaeontologie. Monatshefte, v. 4, p. 232–241. 391 Kahler, F., and Kahler, G., 1980, Fusuliniden aus den Kalken der Trogkofel-Schichten der Karnischen 392 Alpen. Fusulinids from limestones of the Trogkofel Formation in the Carnic Alps: Carinthia II, 393 Sonderheft, no. 36, p. 183–254. 394 Kahler, F., and Krainer, K., 1993, The Schulterkofel section in the Carnic Alps, Austria; implications for the 395 Carboniferous-Permian boundary: Facies, v. 28, 1-2, p. 257–276. 396 Kahler, F. & Kahler, G., 1938, Beobachtungen an Fusuliniden der Karnischen Alpen: Zentr. Bl. Min. Geol. 397 Paläont., Abt. B,, no. 4, p. 101–115. 398 Kalmykova, M.A., 1967, Permian fusulinids of Darvaz, in Likharev, B.K., ed., Biostratigraphic volume. 399 Transactions of All-Union Geological Research Institute (VSEGEI), Volume 2, p. 116. 400 Karapetov, S.S., and Leven, E.Y., 1973, Verkhnepaleozoyskiye otlozheniya Tsentral'nogo Afganistana 401 (basseyn r. Gil'mend). Upper Paleozoic of central Afghanistan, basin of the Helmand River: Byulleten' 402 Moskovskogo Obshchestva Ispytateley Prirody, Otdel Geologicheskiy, v. 48, no. 1, p. 30–40. 403 Kashik, D.S., Alekseeva, I.A., and Polozova, A.N., 1969, The stratigraphy of lower Permian deposits of 404 north of the Russian Platform: Report of USSR, v. 187, no. 2, p. 399. 405 Khodjanyazova, R., Davydov, V., Montaez, I., and Schmitz, M., 2014, Climate- and eustasy-driven cyclicity 406 in Pennsylvanian fusulinid assemblages, Donets Basin (Ukraine): Palaeogeography, Palaeocimatology 407 and Palaeogeography, v. 396, p. 41–61. 408 Khodjanyazova, R.R., and Davydov, V.I., 2013, Late Moscovian fusulinids from the "N" Formation (Donets 409 Basin, Ukraine): Journal of Paleontology, v. 87, no. 1, p. 44–68, doi: 10.1666/11-132R1.1. 410 Khvorova, I.V., 1961, Flysch and lower molasse formations of the south Urals: Transactions of Geological 411 Institute, Academy of Sciences of the USSR, v. 37, p. 1–351. 412 Kim, A.I., Salimova, F.A., Kim, I.A., and Meshchankina, N.A., editors, 2007, Palaeontological Atlas of 413 Phanerozoic faunas and floras of Uzbekistan. Republic of Uzbekistan State Committee on Geology 414 and Mineral Resources.: PALAEOZOIC (CAMBRIAN, ORDOVICIAN, SILURIAN, DEVONIAN, 415 CARBONIFEROUS, PERMIAN): Tashkent, REPUBLIC OF UZBEKISTAN STATE COMMITTEE ON GEOLOGY 416 AND MINERAL RESOURCES, Volume 1. 417 Kireeva, G.D., Scherbovich, S.F., Dobrokhotova, S.V., Ketat, O.B., Malkovsky, F.S., Semina, S.A., Chernova, 418 I.A., and Yagofarova, F.Z., 1971, Schwagerina vulgaris and Schwagerina fusiformis zone of the 419 Asselian Stage of the Russian Platform and the western slope of the southern Urals: Questions of 420 micropaleonotlogy, no. 14, p. 70. 421 Kobayashi, F., and Altiner, D., 2008, Fusulinoidean faunas from the Upper Carboniferous and Lower 422 Permian platform limestone in the Hadim area, central Taurides, Turkey: Rivista Italiana di 423 Paleontologia e Stratigrafia, v. 114, no. 2, p. 191–232. 424 Kobayashi, F., and Altiner, D., 2008, Late Carboniferous and Early Permian fusulinoideans in the central 425 Taurides, Turkey; biostratigraphy, faunal composition and comparison: Journal of Foraminiferal 426 Research, v. 38, no. 1, p. 59–73. 427 Konovalova, M.V., 1991, Stratigraphy and fusulinids of Upper Carboniferous and Lower Permian of 428 Timan-Pechora oil- and gasbearing province: Moscow, Nedra Publishing House, Ukhta Geological- 429 Exploration Expedition, 201 p. (In Russian). 430 Korolyuk, I.K., Kirillova, I.A., Melamud, Y.L., and Rauser-Chernousova, D.M., 1970, Nizhnepermskiy 431 biogermnyy massiv Shakhtau (Bashkiriya). The lower Permian bioherm at Shakhtau, Bashkiria: 432 Byulleten' Moskovskogo Obshchestva Ispytateley Prirody, Otdel Geologicheskiy, v. 45, no. 4, p. 46– 433 59.

434 Korzhenevsky, I.D., 1940, Nekotorye vidy fusulinid iz nizhnepermskikh izvestnyakov Ishimbaevskikh and 435 Sterlitamakskikh gor-odinochek. On some new fusulinid species from the lower Permian limestones 436 of Ishimbajevo and from monad necks of Sterlitamak (western slope of south Urals), in Transactions 437 of Geological Institute Academy of Sciences of U.S.S.R., Geological series, no 2,, p. 1. 438 Krainer, K., and Davydov, V., 1998, Facies and biostratigraphy of the Late Carboniferous/Early Permian 439 sedimentary sequence in the Carnic Alps (Austria/Italy): Geodiversitas, v. 20, no. 4, p. 643–662. 440 Lee, J.S., 1927, Fusulinidae of North China.: Paleontologica Sinica. Series B, v. 4, no. 1, p. 172. 441 Leven, E., 1993, Early Permian fusulinids from the central Pamir: Rivista Italiana di Paleontologia e 442 Stratigrafia, v. 99, no. 2, p. 151–198. 443 Leven, E.J., 1971, Les gisements permiens et les fusulinides de l'Afghanistan du Nord. The Permian beds 444 and fusulinids of northern Afghanistan: Notes et Memoires sur le Moyen-Orient, 12, Part 1, p. 1–35. 445 Leven, E.J., 1995, Lower Permian fusulinids from the vicinity of Ankara (Turkey): Rivista Italiana di 446 Paleontologia e Stratigrafia, v. 101, no. 3, p. 235–248. 447 Leven, E.J., 1997, Permian stratigraphy and Fusulinida of Afghanistan with their paleogeographic and 448 paleotectonic implications, Special Paper - Geological Society of America, Volume 316, 134 p. 449 Leven, E.J., 1998, Permian fusulinids assemblages and stratigraphy of the Transcaucasia: Rivista Italiana 450 di Paleontologia e Stratigrafia, v. 104, no. 3, p. 299–328. 451 Leven, E.J., 2003, The Permian stratigraphy and fusulinids of the Tethys: Rivista Italiana di Paleontologia 452 e Stratigrafia, v. 109, no. 2, p. 267–280. 453 Leven, E.J., and Davydov, V.I., 2001, Stratigraphy and fusulinids of the Kasimovian and lower Gzhelian 454 (Upper Carboniferous) in the southwestern Darvaz (Pamir): Rivista Italiana di Paleontologia e 455 Stratigrafia, v. 107, no. 1, p. 3–45. 456 Leven, E.J., and Gorgij, M.N., 2002, Upper Carboniferous-Permian stratigraphy and fusulinids from the 457 Anarak region, central Iran: Russian Journal of Earth Sciences, v. 8, no. 2. 458 Leven, E.J., and Gorgij, M.N., 2011, First record of Gzhelian and Asselian Fusulinids from the Vazhnan 459 Formation (Sanandaj-Sirjan zone of Iran): Stratigraphy and Geological Correlation, v. 19, no. 5, p. 460 486–501, doi: 10.1134/S0869593811050066. 461 Leven, E.J., and Okay, A.I., 1996, Foraminifera from the exotic Permo-Carboniferous limestone blocks in 462 Karakaya Complex, northwestern Turkey: Rivista Italiana di Paleontologia e Stratigrafia, v. 102, no. 2, 463 p. 139–174. 464 Leven, E.J., and Taheri, A., 2003, Carboniferous-Permian stratigraphy and fusulinids of east Iran; 465 Gzhelian and Asselian deposits of the Ozbak-Kuh region: Rivista Italiana di Paleontologia e 466 Stratigrafia, v. 109, no. 3, p. 399–415. 467 Leven, E.Y., 1967, Stratigrafiya i fuzulinidy permskikh otlozheniy Pamira; stratigraphy and fusulinides of 468 the Pamirs Permian deposits: Trudy - Geologicheskiy Institut (Moskva), v. 167, p. 215. 469 Leven, E.Y., 1974, Biostratigrafiya verkhnego paleozoya Yugo-Zapadnogo Darvaza. Biostratigraphy of the 470 upper Paleozoic in southwestern Darvaz: Izvestiya Akademii Nauk SSSR. Seriya Geologicheskaya, v. 3, 471 p. 55–62. 472 Leven, E.Y., 1979, Bolorskiy yarus permi; obosnovaniye, kharakteristika, korrelyatsiya. The Bolorian 473 Stage of the Permian: designation, characteristics and correlation: Izvestiya Akademii Nauk SSSR. 474 Seriya Geologicheskaya, v. 1979, no. 1, p. 53–65. 475 Leven, E.Y., 1980, The Bolorian Stage of the Permian System; establishment, characteristics, and 476 correlation: International Geology Review, v. 22, no. 5, p. 587–597. 477 Leven, E.Y., 1980, Yakhtashskiy yarus permi; obosnovaniye, kharakteristika, korrelyatsiya. Yakhtashskiy 478 Stage of the Permian designation, characteristics, correlation: Izvestiya Akademii Nauk SSSR. Seriya 479 Geologicheskaya, v. 1980, no. 8, p. 50. 480 Leven, E.Y., 1981, Verkhniy paleozoy basseynov rek Charymdary, Gundary i Zidadary (Yugo-Vostochnyy 481 Darvaz). The upper Paleozoic of the Charymdara, Gandara, and Zidadara basins; southwestern

482 Darvaz: Byulleten' Moskovskogo Obshchestva Ispytateley Prirody, Otdel Geologicheskiy, v. 56, no. 4, 483 p. 40–52. 484 Leven, E.Y., 1982, The Permian Yakhtashian Stage; its basis, characteristics, and correlation: 485 International Geology Review, v. 24, no. 8, p. 945–954. 486 Leven, E.Y., editor, 2009, Upper Carboniferous and Permian of the western Tethys: fusulinids, 487 stratigraphy, paleogeography: Moscow, Transactions of Geological Institute of Russian Academy of 488 Sciences, Volume 590, 1-237 (In Russian). 489 Leven, E.Y., Boyko, M.S., Reuymers, A.N., Leonova, T.B., and Bogoslovskaya, M.F., 2002, Lower Permian 490 near Verkhneozernoye village; Southern Urals: Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 10, 491 no. 5, p. 44–58. 492 Leven, E.Y., and Davydov, V.I., 1979, Novyye dannyye po stratigrafii permskikh krasnotsvetnykh tolshch 493 yugo-zapadnogo Darvaza. New stratigraphic data on Permian red beds of southwestern Darvaz: 494 Izvestiya Vysshikh Uchebnykh Zavedeniy. Geologiya i Razvedka, v. 1979, no. 8, p. 13–20. 495 Leven, E.Y., and Davydov, V.I., 1991, O granitsakh i ob"yeme nizhney zony assel'skogo yarusa. The 496 boundaries and volume of the lower zone of the Asselian stage: Izvestiya Akademii Nauk SSSR. Seriya 497 Geologicheskaya, v. 12, p. 61–73. 498 Leven, E.Y., Davydov, V.I., and Gorgij, M N, 2006, Pennsylvanian stratigraphy and fusulinids of central 499 and eastern Iran: Palaeontologia Electronica, v. 8, no. 1, p. 36. 500 Leven, E.Y., and Dmitriyev, V.Y., 1974, Stratotype of the Darvaz Stage of the Permian: Doklady. Earth 501 Science Sections, v. 215, 1-6, p. 22–23. 502 Leven, E.Y., and Gordzh, M.N., 2011, First occurrence of Gzhelian and Asselian fusulinids in Vajnan 503 Formation; Sanandaj-Sirjan Zone, Iran: Stratigrafiya Geologicheskaya Korrelyatsiya, v. 19, no. 5, p. 504 16–31. 505 Leven, E.Y., and Gorgij, M.N., 2006, Gzhelian Fusulinids First Discovered in Central Iran: Stratigraphy, 506 Geological Correlations, v. 14, no. 1, p. 19–29. 507 Leven, E.Y., and Gorgij, M.N., 2006, Pervyye nakhodki gzhel'skikh fuzulinid v Tsentral'nom Irane. First 508 discovery of Gzhelian fusulinids in central Iran: Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 14, 509 no. 1, p. 22–32. 510 Leven, E.Y., and Gorgij, M.N., 2008, Bolorian and Kubergandian stages of the Permian in the Sanandaj- 511 Sirjan zone, Iran: Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 16, no. 5, p. 3–14. 512 Leven, E.Y., and Gorgij, M.N., 2011, The Kalaktash and Halvan assemblages of Permian fusulinids in the 513 Padeh and Sang-Variz sections (Halvan Mountains, Yazd Province, central Iran): Stratigrafiya 514 Geologicheskaya Korrelyatsiya, v. 19, no. 2, p. 20–39. 515 Leven, E.Y., Leonova, T.B., and Dmitriyev, V.Y., 1992, Perm Darvaz-Zaalaiskoi zony Pamira (fusulinidy, 516 ammonoidei, stratigraphiya). The Permian of Darvas-Zaalai zone of Pamirs (fusulinids, ammonoids, 517 stratigraphy), Transactions of Paleontological Institute of Russian Academy of Sciences, 203 p. 518 Leven, E.Y., and Ozkan, R., 2004, Novyye nakhodki permskikh fuzulinid v Turtsii i nekotoryye voprosy ikh 519 biogeografii. New occurrences of Permian Fusulinidae in Turkey and aspects of biogeography: 520 Stratigrafiya, Geologicheskaya Korrelyatsiya, v. 12, no. 4, p. 20–31. 521 Leven, E.Y., and Scherbovich, S.F., 1978, Fuzulinidy i stratigrafiya assel'skogo yarusa Darvaza. Fusulinids 522 and stratigraphy of the Asselian Stage of the Darvaz Range: Moscow, Nauka, Moscow Society of 523 Natural Studies, Geological. Series, 1-162 (In Russian). 524 Leven, E.Y., and Shcherbovich, S.F., 1980, Kompleks fuzulinid sakmarskogo yarusa Darvaza. A fusulinid 525 assemblage of the Darvaz Sakmarian Stage: Voprosy Mikropaleontologii, no, v. 23, p. 71–85. 526 Leven, E.Y., and Shcherbovich, S.F., 1980, New fusulinid species from the Sakmarian of the Darvaz: 527 Paleontological Journal, v. 14, no. 3, p. 18–29. 528 Leven, E.Y., and Vaziri, M.H., 2004, Carboniferous-Permian stratigraphy and fusulinids of eastern Iran; 529 the Permian in the Bag-e-Vang section (Shirgesht area): Rivista Italiana di Paleontologia e 530 Stratigrafia, v. 110, no. 2, p. 441–465.

531 Makhlina, M.K., Alekseyev, A.S., Goreva, N.V., Goryunova, R.V., Isakova, T.N., Kossovaya, O.L., Lazarev, 532 S.S., Lebedev, O.A., and Shkolin, A.A., 2001, Sredniy karbon Moskovskoy sineklizy (yuzhnaya chast’); 533 v 2 tomakh; Tom 2, Middle Carboniferous of southern Moscow Syneclise; Volume 2, Paleontological 534 Characteristics: Moscow, Rossiyskaya Akademiya Nauk, Paleontologicheskiy Institut [In Russian], 535 (None). 536 Makhlina, M.K., and Isakova, T.N., 1997, Melekhovian Horizon: A New Stratigraphic Unit of the Gzhelian 537 Stage, Upper Carboniferous (East European Platform): Stratigraphy, Geological Correlations, v. 5, no. 538 5, p. 458. 539 Malakhova, N.P., and Chuvashov, B.I., 1970, Verkhnepaleozoyskiye terrigennyye otlozheniya Urala. 540 Upper Paleozoic terrigenous deposits of the Urals: Akademiya Nauk SSSR, Ural'skiy Filial, Institut 541 Geologii i Geokhimii, Trudy, v. 88, p. 13. 542 Maslov, V.V., Goryunova, L.F., and Gibshman, N.B., 2016, Evaluation of prospecitves of the oil-gas 543 production in the Upper Paleozoic of Ustyurt on the bases of biostratigraphic analyses.: Neftyegaz 544 Territury, no. 11, 57-63 (In Russian). 545 Masumov, A.S., Borisov, O.M., and Bensh, F.R., 1978, Verkhniy paleozoy Sredinnogo i Yuzhnogo Tyan'- 546 Shanya. The upper Paleozoic of the central and southern Tien Shan: Tashkent, Uzbekistan, Fan 547 Publishing House. 548 Mikhailova, Z.P., 1974, Fusulinids of Upper Carboniferous of Pechoras Preurals: Leningrad, Nauka, Inst. 549 Geol. KOMI dep. of Academy of Science of the. U.S.S.R, 116 (In Russian). 550 Mikhailova, Z.P., 1997, Fuzulinidy verkhnego karbona i nizhney permi Bol'shezemel'skoy tundry. 551 Fusulinids from Upper Carboniferous and Lower Permian of Bolshezemelskaya Tundra: Trudy - 552 Institut Geologii, v. 91, p. 14–29. 553 Mikhailova, Z.P., and Chermnykh, V.A., 1997, Fusulinoida zones of the Upper Carboniferous and Lower 554 Permian in the Northern Urals, in Ross, C.A., Ross, J.R.P., et al., eds., Special Publications - Cushman 555 Foundation for Foraminiferal Research, p. 99–103. 556 Miklukho-Maclay, A.D., 1963, Upper Paleozoic of Central Asia: Leningrad, Leningrad State University. 557 Miklukho-Maklay, A.D., 1949, Verkhnepaleozojskie fuzulinidy Srednej Azii (Fergana, Darvaz i Pamir), in 558 Monografiya LGU: Leningrad, p. 127. 559 Morozova, A.P., Zolotova, V.P., Konovalova, M.B., and Ogneva, I.I., 1980, New species of 560 Pseudofusulinids of Sakmarian and Artinskian Stages of northern part of Preurals Trough and 561 surrounding areas, in Rauser-Chernousova, D.M., and Chuvashov, B.I., ed., Biostratigraphy of 562 Artinskian and Kungurian Stages of the Urals.: Ekaterinburg, Academy of Sciences of USSR, Uralian 563 Branch., p. 11. 564 Murav'yev, I.S., Grigor'yeva, A.D., Gizatulin, Z.Z., Yermoshkin, N.V., Zhulitova, V.Y., Igonin, V.M., Isakova, 565 T.N., Kossovaya, O.L., and Morozova, I.P., 1984, The "Yablonevyy Ovrag" section (Samara Luka) is the 566 hypostratotype of the Gzhelian Stage and the probable stratotype of Carboniferous-Permian 567 boundary: Trudy - Mezhvedomstvennyy Stratigraficheskiy Komitet SSSR, v. 13, p. 26–42. 568 Nikolaev, A.I., 2011, Fusulinoids of Moscovian Stage in the Precaspian bassin.: Transactions of VNIGRI, p. 569 168. 570 Nilsson, I., and Davydov, V.I., 1997, Fusulinid biostratigraphy in Upper Carboniferous (Gzhelian) and 571 Lower Permian (Asselian-Sakmarian) succession in Spitsbergen, Arctic Norway: Permophiles, no. 30, 572 p. 18–27. 573 Nilsson, I., Hakansson, E., Madsen, L., Schack Pedersen, S.A., and Stemmerik, L., 1991, Stratigraphic 574 significance of new fusulinid samples from the upper Palaeozoic Mallemuk Mountain Group, North 575 Greenland: Rapport - Gronlands Geologiske Undersogelse (1964), v. 150, p. 29–32. 576 Novak, M., 2007, Depositional environment of Upper Carboniferous – Lower Permian beds in the 577 Karavanke Mountains (Southern Alps, Slovenia): Geologija, v. 50, no. 2, p. 247–268.

578 Okuyucu, C., 2007, New Middle Permian foraminifers (Chitralinidae) from the Karakaya Complex, in 579 northwestern Turkey: Comptes rendus Palevol, v. 6, no. 5, p. 311–319, doi: 580 10.1016/j.crpv.2007.05.004. 581 Okuyucu, C., 2008, Biostratigraphy and systematics of late Asselian-early Sakmarian (Early Permian) 582 fusulinids (Foraminifera) from southern Turkey: Geological Magazine, v. 145, no. 3, p. 413–434, doi: 583 10.1017/S0016756808004482. 584 Okuyucu, C., 2013, Fusulinid zonation of the late Moscovian-early Sakmarian sequences from the 585 Taurides, southern Turkey: Neues Jahrbuch fuer Geologie und Palaeontologie. Abhandlungen, v. 586 268, no. 3, p. 237–258, doi: 10.1127/0077-7749/2013/0328. 587 Orlov-Labkovsky, O., and Bensh, F.R., 2015, Atlas of Foraminifera of the Carboniferous and Permian 588 (Cisuralian) of Uzbekistan and adjacent regions, Tien Shan.: Sofia, Bulgaria, Pensoft Publishers. 589 Pnev, V.P., Grozdilova, L.P., Izotova, M.N., Kruchinina, O.N., Mikhaylova, Y.N., and Simakova, M.A., 1971, 590 The Belaya Gora (Tastuba) Formation of the Sakmarian age in the western Urals: Zapiski 591 Leningradskogo, Ordena Lenina, Ordena Oktyabr'skoy Revolyutsii i Ordena Trudovogo Krasnogo 592 Znameni Gornogo Instituta im G.V. Plekhanova, v. 59, no. 2, p. 128–143. 593 Pnev, V.P., Grozdilova, L.P., Simakova, M.A., Apukhtina, M.A., Kruchinina, O.A., and Mikhaylova, Y.N., 594 1967, The Filinok (Novo-Kurkino) horizon of the Assel suite in the western Urals: Leningrad, Gorn. 595 Inst., Zap., v. 53, no. 2, p. 45–50. 596 Pnev, V.P., and Polozova, A.N., 1981, The data on the stratigraphy of Artinskian Stage of Middle 597 Southern Urals;: Записки Ленинградского ордена Ленина, ордена Октябрьской Революции и 598 ордена Трудового Красного Знамени горного института им. Г. В. Плеханова, т. LXXXV, 1981 г.: 599 Notes of St. Petersburg Mining Institute, v. 85, 42-49 (In Russian). 600 Polozova, A.N., 1979, New species of fusulinid from the Carboniferous-Permian boundary beds in the 601 Orenburg and Aktubinsk Preurals, in Wagner, R.H., Higgins, A.C., and Meyen, S.V., ed., The 602 Carboniferous of the U.S.S.R; its stratigraphic significance and outstanding problems of world-wide 603 correlation. Yorkshire Geological Society Occasional Publication, p. 192–195. 604 Ponomareva, G.Y., Kossovaya, O.L., and Khopta, I.S., 2015, Middle Urals. Carboniferous and Permian 605 marine and continental successions Field Trip Guidebook of 13 ICCP Perm city: "Aster" Printing 606 house 112: Perm, Russia, "Aster" Printing house 112 p. 607 Pronin, A.P., Kuanyshev, F.L., Salykhova, A., and Mil'kina, N.V., 2011, New data on the Paleozoic deposits 608 in the area of joining of Precaspian depression and Turan Plate (Kaspian Sea subsurface).: Oil and 609 Gas Geology, no. 4, 21-25 (In Russian). 610 Pronin, A.P., Turkov, O.S., Kalmuratova, S.A., and Mil'kina, N.V., 1997, New data regarding the origin of 611 the Paleozoic deposits of Buzachi peninsula.: Geology of Kazakstan, no. 4, 43-52 (In Russian); 612 Rauser-Chernousova, D.M., Granitsa karbona i permi. The Carboniferous-Permian boundary. 613 Rauser-Chernousova, D.M., O nekotorykh kriteriyakh paleobiogeograficheskogo rayonirovaniya (na 614 primere izucheniya assel'skikh i sakmarskikh foraminifer). Some criteria for paleobiogeographic 615 zoning, exemplified by studies of Asselian and Sakmarian foraminifera. 616 Rauser-Chernousova, D.M., 1938, The upper Palaeozoic Foraminifera of the Samara bend and the trans- 617 Volga region: Acad. Sci. URSS, Inst. Geol., Tr., v. 7, p. 69–167. 618 Rauser-Chernousova, D.M., 1940, Stratigraphy of the upper Carboniferous and Artinskian [Permian] 619 stage on the western slope of the Urals and materials concerning the fauna of fusulinids: Cep., 620 [Acad. Sci. USSR, Inst. Geol. Sci., Tr., Geol. Ser., v. 2, no. 7, p. 37–101. 621 Rauser-Chernousova, D.M., editor, 1949, Foraminifers of Upper Carboniferous and Artinskian deposits of 622 Bashkirian Preurals. 623 Rauser-Chernousova, D.M., 1949, Stratigrafiya verkhnekamennougol’nykh i artinskikh otlozhenyi 624 Bashkirskovo Priuralya (Stratigraphy of Upper Carboniferous and Artinskian deposits of Bashkirian 625 Pre-Ural Mts.): Proceedings of Geological Institute of Academy of Sciences of USSR, Series 626 Geological, v. 105, no. 35, p. 3.

627 Rauser-Chernousova, D.M., 1950, Fatsii verkhnekamennougolnykh i artinskikh otlozhenii Sterlitamaksko- 628 Ishimbaiskogo Priuralya (na osnove izucheniya fuzulinid): Trudy - Geologicheskiy Institut, Akademiya 629 Nauk SSSR, v. 119, no. 43, p. 1–108. 630 Rauser-Chernousova, D.M., 1961, Reviziya shvagerin s blizkimi rodami i granitsa karbona i permi. The 631 revision of Schwagerina and related genera and the Carboniferous-Permian boundary.: Voprosy 632 micropaleontologii, no. 4, p. 3. 633 Rauser-Chernousova, D.M., 1965, Foraminiferi stratotipicheskogo razreza sakmarskogo yarusa (r. 634 Sakmara, Yuzhnyj Ural). Foraminifers from the Sakmarian type section (Sakmara River, S. Urals).: 635 Moscow, Nauka, Transactions of Geologcial Institute of Academy of Sciences of SSSR. 636 Rauser-Chernousova, D.M., 1973, Paleobiogeografiya assel'skikh i sakmarskikh morey po fuzulinidam v 637 aspekte zonal'nykh podrazdeleny. Paleobiogeography of the Asselian and Sakmarian seas based on 638 fusulinids in the light of zonal subdivisions: Voprosy Mikropaleontologii, v. 16, p. 36–61. 639 Rauser-Chernousova, D.M., Balahova, N.N., Chernova, E.I., Dalmatskaya, I.I., and Reitlinger, E.A., 640 Stratigrafiya srednekamennougolnykh otlozhenii tsentralnoi i vostochnoi chastei Russkoi platformy 641 (na osnove izucheniya foraminifer); 1, Moskovskaya sinekliza (Regionalnaya stratigrafiya SSSR, tom 642 2). 643 Rauser-Chernousova, D.M., Belyaev, g.M., and Reitlinger, E.A.R., 1940, On Carboniferous Foraminifera of 644 the Samara Bend [Volga river region], v. 88. 645 Rauser-Chernousova, D.M., Chernova, E.I., Gryzleva, N.D., Kireeva, G.D., Leontovich, G.E., and Safonova, 646 T.P., Srednekamennougolnye fuzulinidy Russkoi platformy i sopredelnykh oblastei; spravochnik, 647 opredelitel. 648 Rauser-Chernousova, D.M., Chernysheva, N.E., Glebovskaya, E.M., Grozdilova, L.P., Lipina, O.A., 649 Suleimanov, I.S., and Vissarionova, A.Y., 1948, Stratigrafiya i foraminifery nizhnego karbona Russkoi 650 platformy i Priuralya: Trudy - Geologicheskiy Institut, Akademiya Nauk SSSR, v. 62, p. 19. 651 Rauser-Chernousova, D.M., Grozdilova, L.P., Pnev, V.P., Sultanayev, A.A., and Alksneh, A.E., 1981, 652 Stratotip burtsevskogo goriyouta artinskogo yarusa (Yuzhnyy Ural). Stratotype of Burtsevka Horizon 653 of the Artinskian, South Urals: Izvestiya Akademii Nauk SSSR. Seriya Geologicheskaya, v. 1981, no. 9, 654 p. 64–72. 655 Rauser-Chernousova, D.M., Gryzlova, N.D., Kireeva, G.D., Leontovich, G.E., Safonova, T.P., and Chernova, 656 E.I., 1951, Middle Carboniferous fusulinids in the Russian platform and adjacent area. The Guide- 657 book: Moscow, Academy of Sciences of the USSR, (None), 3-380 (In Russian.). 658 Rauser-Chernousova, D.M., Ivanova, E.A., Grozdilova, L.P., Makhlina, M.K., Alkene, A.E., Kireeva, G.D., 659 Konovalova, M.B., Meyen, S.V., Morozova, I.P., Rosovskaya, S.E., Faddeeva, I.Z., Shamov, D.F., 660 Shchegolev, A.K., and Shcherbakova, M.V., 1979, The Upper Carboniferous Series: Occasional 661 Publication Yorkshire Geological Society, no, v. 4, p. 147–174. 662 Rauser-Chernousova, D.M., and Ivanova, Y.A., 1978, Verkhniy karbon. The Upper Carboniferous: Trudy - 663 Mezhvedomstvennyy Stratigraficheskiy Komitet SSSR, v. 6, p. 223–228. 664 Rauser-Chernousova, D.M., Ivanova, Y.A., Korolyuk, I.K., Morozova, I.P., and Fotiyeva, N.N., 1977, K 665 kharakteristike stratotipa sterlitamakskogo gorizonta (nizhnyaya perm', massiv Shakhtau, 666 Bashkiriya). Characteristics of the stratotype of the Sterlitamak Horizon, Lower Permian, Shakhtau 667 Massif, Bashkiria: Byulleten' Moskovskogo Obshchestva Ispytateley Prirody, Otdel Geologicheskiy, v. 668 52, no. 6, p. 24–37. 669 Rauser-Chernousova, D.M., and Izotova, M.N., 1980, K revizii rannepermskikh vidov gruppy 670 Pseudofusulina urdalensis. The revision of the Lower Permian species of the Pseudofusulina 671 urdalensis group: Voprosy Mikropaleontologii, no, v. 23, p. 86–95. 672 Rauser-Chernousova, D.M., Rozovskaya, S.E., and Scherbovich, S.E., editors, 1974, Foraminifers: 673 Yerevan, Academy of Sciences of Armenian SSR, Atlas of Fossils of Armenian SSR, 86-103. 674 Rauser-Chernousova, D.M., and Shchegolev, A.K., 1979, The Carboniferous Permian boundary in the 675 USSR: Occasional Publication Yorkshire Geological Society, no, v. 4, p. 175–191.

676 Rauser-Chernousova, D.M., and Shcherbovich, S.F., 1958, o shvagerinovom gorizonte tsentral'noi chasti 677 Russkoi Platformy. About schwagerins horizons in the Central part of the Russian Platform, in 678 Menner, V.V., ed., Transactions of Geological Institute, Academy of Sciences of the USSR: Moscow, 679 Academy of Sciences of the USSR, Volume 13, p. 3. 680 Rauser-Chernousova D.M. and Scherbovich S.F., editor, 1949, Schwagerins of European part of U.S.S.R. 681 Transactions of Institute of Geological Sciences, No 35. 682 Rauser-Chernousova, D.M., and Chuvashov, B.I., editor, 1980, Biostratigraphy of Artinskian and 683 Kungurian Stages of the Urals.: Ekaterinburg, Academy of Sciences of USSR, Uralian Branch. 684 Rauzer-Chernousova, D.M., Grozdilova, L.P., Pnev, V.P., Sultanayev, A.A., and Alksneh, A.E., 1981, 685 Stratotype of Burtsevka Horizon of the Artinskian, South Urals: Izvestiya Akademii Nauk SSSR. Seriya 686 Geologicheskaya, v. 1981, no. 9, p. 64–72. 687 Remizova, S.T., 1989, Novyye nizhnepermskiye fuzulinellidy Severnogo Timana. New Lower Permian 688 fusulinids of northern Timan: Trudy - Institut Geologii, v. 71, p. 17–25. 689 Remizova, S.T., 1995, Foraminifers and biostratigraphy of Upper Carboniferous of the Northern Timan: 690 Syktyvkar, Transactions of KOMI Scientific Center of Uralian branch of Russian Academy of Sciences. 691 Remizova, S.T., 1997, Fusulinids and stratigraphy of the Sakmarian Stage in the northern Timan: Prace 692 Panstwowego Instytutu Geologicznego (1988), v. 157, Part 1, p. 329–338. 693 Remizova, S.T., 2004, Fusulinoids of Timan: evolution, biostratigraphy and paleobiogeography.: 694 Ekaterinburg, Russian Academy of Sciences, (None). 695 Rozovskaya, S.E., 1940, Contribution to the study of Fusulinidae of the Moscow Basin: Doklady Akademii 696 Nauk SSSR, v. 28, no. 5, p. 477–480. 697 Rozovskaya, S.E., 1949, Stratigraficheskoe raspredelenie fuzulinid v verkhnekamennougolnykh i 698 nizhnepermskikh otlozheniyakh yuzhnogo Urala: Doklady Akademii Nauk SSSR, v. 69, no. 2, p. 249– 699 252. 700 Rozovskaya, S.E., 1950, Rod Triticites, ego razvitie i stratigraficheskoe znachenie. Triticites genus its 701 evolution and stratigraphic significance: Kiev, Trudy Paleontologicheskogo Instituta [In Russian], 80 702 p. 703 Rozovskaya, S.E., 1952, Fuzulinidy verkhnego karbona i nizhnei permi yuzhnogo Urala: Trudy 704 Paleontologicheskogo Instituta, v. 40, p. 5–48. 705 Scherbakova, M.V., 1986, Carboniferous-Permian boundary beds in Ostanets section, on Kos'va River., in 706 Chuvashov, B.I., Leven, E. Y. and Davydov, V.I., ed., Carboniferous--Permian Boundary beds of the 707 Urals, Pre-Urals and Central Asia. Nauka Publishing House, Moscow. (In Russian): Moscow, Nauka, 708 18-21 (In Russian). 709 Scherbovich, S.F., 1969, Late Gzhelian and Asselian fusulinids in Precaspian Syneclise: Transactions of 710 Geological Institute, Academy of Sciences of the USSR, v. 176, 1-104 (In Russian). 711 Schmitz, M.D., and Davydov, V.I., 2012, Quantitative radiometric and biostratigraphic calibration of the 712 Pennsylvanian - Early Permian (Cisuralian) time scale, and pan-Euramerican chronostratigraphic 713 correlation.: Geological Society of America Bulletin, v. 124, p. 549–577. 714 Shamov, D.F., 1940, Geology of Ishimbay oil-field area: Soviet Geology,, no. 11, p. 6. 715 Shamov, D.F., 1958, The group of inflate-fusiform pseudofusulinids from Shwagerina Horizon of Ishimby- 716 Sterlitamak oil-bearing region.: Transactions of Geological Institute of Academy of Sciences of USSR, 717 no. 13, 134-157 (In Russian). 718 Shcherbakova, M.V., Shcherbakov, O.A., Chuvashov, B.I., and Kitayev, P.M., 1979, Kamennougol'nyye 719 otlozheniya v razreze "Orel". Carboniferous deposits in the "Orel" Profile: Trudy Instituta Geologii i 720 Geokhimii, no, no. 26, p. 48–59. 721 Sjomina, S.A., 1961, Stratigrafiya i foraminifery (fuzulinidy) shvagerinovogo gorizonta Oksko-Tsninskogo 722 podnyatiya [Stratigraphy and Foraminifers (Fusulinids) from the Schwagerina-Horizon of the Oka- 723 Tsna Swell].: Moscow, Nauka, Transactions of Geological Institute of Academy of Sciences of USSR.

724 Sofronitskiy, P.A., Zolotova, V.P., and Baryshnikov, V.V., 1974, Guide-book on the Permian deposits 725 along Kos'va, Sylva and Kama.: Perm, Russia 101 p. (In Russian), Perm State University. 726 Stemmerik, L., Hakansson, E., Madsen, L., Nilsson, I., Piasecki, S., Pinard, S., Rasmussen, J.A., and 727 Anonymous, 1996, Stratigraphy and depositional evolution of the upper Palaeozoic sedimentary 728 succession in eastern Peary Land, North Greenland: Bulletin - Gronlands Geologiske Undersogelse, v. 729 171, p. 45–71. 730 Uzakov, K., 1996, Lithological and biostratigraphic characteristics of pre-Jurrasic deposits of Eastern 731 Ustyurt (Karakalpastan).: Uzbekiston Geologiya Zhurnali = Uzbekskiy Geologicheskiy Zhurnal, no. 4, 732 52-67 (In Russian). 733 Vissarionova, A.Y., Kireeva, G.D., Lipina, O.A., Morozova, V.G., Rauser-Chernousova, D.M., Shamov, D.F., 734 Shcherlovich, S.F., and Suleimanov, I.S., 1949, Foraminifery verkhnekamennougolnykh i artinskikh 735 otlozhenii Bashkirskogo Priuralya: Trudy - Geologicheskiy Institut, Akademiya Nauk SSSR, v. 105, p. 736 35. 737 Volozhanina, P.P., 1962, Upper Carboniferous fusulinids of Timano-Pechora Basin.: Voprosy 738 Mikropaleontologii, no. 6, 116-154 (In Russian). 739 Wahlman, G.P., Davydov, V.I., Nilsson, I., and Anonymous, 1995, Fusulinid biostratigraphy of subsurfaces 740 cores from Conoco 7128/6-1 Well, offshore Barents Sea, Arctic Norway, Panstwowy Instytut 741 Geologiczny, 150 p. 742 Zamilatskaya, T.K., Afanasieva, M.S., Efremova, G.D., and Malyukova, T.N., 1987, Stratigraphy and facies 743 of lower Permian deposits of Karachaganak High (Precaspian Basin).: Izvestiya Vysshikh Uchebnykh 744 Zavedeniy. Geologiya i Razvedka, no. 10, 13-26 (In Russian). 745 Zamilatskaya, T.K., and Gorshkova, V.A., 1987, Stratigraphy and correlation of pre-Kungurian deposits of 746 northern margin of Precaspian Basin: Stratigraphy and Paleontology of Paleozoic of Precaspian 747 Basin, 122-131 (In Russian). 748 Zolotova, V.P., and Baryshnikov, V.V., 1978, Fusulinids of Kungurian Stage of Kamsk Preurals.: 749 Paleontologichesky Zhurnal, no. 3, p. 22. 750 Zolotova, V.P., and Baryshnikov, V.V., 1980, Foraminifers of Kungurian Stage in the type area.: 751 Biostratigraphy of Artinskian and Kungurian Stage of the Urals., no. 1, 72-110 (In Russian). 752 Zolotova, V.P., Shcherbakova, M.V., Ekhlakov, Y.A., Alksne, A.E., Polozova, M.V., Konovalova, M.B., and 753 Kosheleva, V.F., 1977, Fusulinids of the Gzhelian-Asselian Boundary Beds of the Urals, Preurals, and 754 Timan: Voprosy Mikropaleontologii, no. 20, p. 93. 755 756 252 sources