Late Pennsylvanian vegetation dynamics of the Donets Basin, Ukraine

Nataliya Boyarina https://orcid.org/0000-0001-5316-4090

Institute of Geological Sciences, NAS of Ukraine (Kyiv, Ukraine)

Cite as

Boyarina, N. 2024. Late Pennsylvanian vegetation dynamics of the Donets Basin, Ukraine. GEO&BIO, 26: 45-78. [English, with Ukrainian summary]

doi: https://doi.org/10.53452/gb2605

pdf: gb2605-boyarina.pdf

Abstract

The dynamics of the Late Pennsylvanian vegetation cover of the Donets Basin is represented by a succession of formations of new plant communities (phytocoenogenesis) of a wetland forest and woodland biome and a seasonally dry woodland biome because of the changing landscape and climate conditions resulting from the glacio-eustatic changes during the Late Paleozoic Ice Age. The Late Pennsylvanian vegetation consisted of plant communities of wetland marattialean fern-dominated forests on coastal lowlands and wetland lycopsid-fern forests on deltaic plains in the Kasimovian as well as wetland marattialean fern-dominated forests with new dominants on coastal lowlands and wetland lycopsid-pteridosperm-calamitalean-fern forests with new dominants on deltaic plains in the early Gzhelian that were formed according to the evolutionary progressive model of phytocoenogenesis under conditions of an expansion of coastal lowlands and deltaic plains inthe long-term period of a relatively stable higher sea level with frequent sea level fluctuations during the late Kasimovian–early-mid-Gzhelian interglacial interval. The subsequent transformation of wetland forests to wetland woodlands on coastal lowlands and deltaic plains according to the substitutionary regressive model of phytocoenogenesis and the formation of the new seasonally dry peltaspermalen pteridosperm-dominated woodland communities in river valleys according to the evolutionary progressive model of phytocoenogenesis took place against the background of a reduction of coastal lowlands and an expansion of river landscapes in the long-term period of a relatively stable sea level drop during the early phase of the late Gzhelian glacial interval. The new plant communities of wetland calamitalean-fern-pteridosperm woodlands on coastal lowlands and seasonally dry fern-pteridosperm woodlands on lagoon coasts in the late Gzhelian were formed according to the migration progressive model of phytocoenogenesis due to a migration of plants from reduced river landscapes to coastal lowlands and lagoon coasts in the period of the further sea level drop with low-amplitude sea level oscillations and an increased drying during the continued late Gzhelian glacial interval.

Key word: Vegetation changes, phytocoenogenesis, glacioeustasy, Kasimovian, Gzhelian, Donets Basin.

Correspondence to

Nataliya Boyarina; Institute of Geological Sciences, NAS of Ukraine; 55-b Oles Honchar Street, Kyiv, 01054 Ukraine; Email: nboyarina@ukr.net

Article info

Submitted: 15.10.2024. Accepted: 30.12.2024.

References

Bashforth, A. R., C. J. Cleal, M. R. Gibling, H. R. Falcon-Lang, R. F. Miller. 2014. Paleoecology of Early Pennsylvanian vegetation on a seasonally dry tropical landscape (Tynemouth Creek Formation, New Brunswick, Canada). Review of Palaeobotany and Palynology, 200: 229–263. https://doi.org/10.1016/j.revpalbo.2013.09.006

Bashforth, A. R., W. A. DiMichele, C. F. Eble, W. J. Nelson. 2016. Dryland vegetation from the Middle Pennsylvanian of Indiana (Illinois Basin): the dryland biome in glacioeustatic, paleobiogeographic, and paleoecologic context. Journal of Paleontology, 90 (5): 785–814. https://doi.org/10.1017/jpa.2016.25

Bashforth, A. R., W. A. DiMichele, C. F. Eble, H. J. Falcon-Lang, C. V. Looy, S. G. Lucas. 2021. The environmental implications of upper Paleozoic plant-fossil assemblages with mixtures of wetland and drought-tolerant taxa in tropical Pangea. Geobios, 68: 1–45. https://doi.org/10.1016/j.geobios.2021.04.002

Borisenko, Yu. A. 1975. Features of the late Carboniferous lithogenesis in the south-western part of the Donbass. Geologiya i geohimiya iskopaemih uglei, 44: 10–16. [In Russian]

Borisenko, Yu. A. 2014. Facies of continental and subcontinental sediments of the upper Carboniferous of the western part of the Donets Coal Basin. Xarkov, 1–163. [In Russian]. https:// www.gigabaza.ru/doc/174649-pall.html

Boyarina, N. I. 2010. Late Gzhelian pteridosperms with callipterid foliage of the Donets Basin, Ukraine. Acta Palaeontologica Polonica, 55 (2): 343–359. https://doi.org/10.4202/app.2009.0020

Boyarina, N. I. 2016. The late Carboniferous palaeophytocoenoses of the Donets Basin (according to the classification of vegetation by the Braun- Blanquet method). Problems of the Phanerozoic geology of Ukraine: Collection of scientific works of the VII All-Ukrainian Scientific Conference. Lviv, 36–39. [In Ukrainian]

Boyarina, N. I. 2017. Palaeophytocoenological studies of the late Carboniferous vegetation of the Donets Basin. Collection of scientific works of the Institute of Geological Sciences of NAS of Ukraine, 10: 4–14. [In Russian] https://doi.org/10.30836/igs.2522-9753.2017.141682

Boyarina, N. I.2022. Late Pennsylvanian vegetation cover changes in the Donets Basin: syndynamic aspect. Visnyk of V.N. Karazin Kharkiv National University, series "Geology. Geography. Ecology", 56: 8–23. https://doi.org/10.26565/2410-7360-2022-56-01

Boyarina, N. I. 2023. The Late Pennsylvanian vegetation of the Donets Basin, Ukraine: Syntaxonomy of plant communities. GEO&BIO, 24: 64–98. https://doi.org/10.15407/gb2406

Braun-Blanquet, J. 1964. Pflanzensociologie. Grundzüge der Vegetationskunde. 3nd. ed. Springer-Verlag, Aufl. Wien, 1–865.

Broutin, J., J. Doubinger, G. Farjanel, P. Freytet, H. Kerp, [et al.]. 1990. Le renouvellement des flores au passage Carbonifére–Permian: approche stratigraphique, biologique, sédimentologique. Comptes Rendus de l'Académie des sciences, 2 (311): 1563–1569.

Cecil, C. B. 2013. An overview and interpretation of autocyclic and allocyclic processes and the accumulation of strata during the Pennsylvanian–Permian transition in the central Appalachian Basin, USA. International Journal of Coal Geology, 119: 21–31. https://doi.org/10.1016/j.coal.2013.07.012

Cecil, C. B., W. A. DiMichele, S .D. Elrick. 2014. Middle and Late Pennsylvanian cyclothems, American Midcontinent: Ice-age environmental changes and terrestrial biotic dynamics. Comptes Rendus. Géoscience, 346 (7–8): 159–168. https://doi.org/10.1016/j.crte.2014.03.008

Cecil, C. B, F. T. Dulong, R. R. West, R. Stamm, B. A. Wardlaw, N. T. Edgar. 2003. Climate controls on the stratigraphy of a Middle Pennsylvanian cyclothem in North America. In: Cecil, C. B., Edgar, N. T. (eds.). Climate controls on stratigraphy. Special Publications,77: 151–182. https://doi.org/10.2110/pec.03.77.0151

Cecil, C. B, R. W. Stanton, S. G. Neuzil, F. T. Dulong, L. F. Ruppert, B. S. Pierce. 1985. Paleoclimate controls on late Paleozoic sedimentation and peat formation in the central Appalachian Basin (U.S.A.). International Journal of Coal Geology, 5: 195–230. https://doi.org/10.1016/0166-5162(85)90014-X

Cleal, C. J. 2007. The Westphalian-Stephanian macrofloral record from the South Wales Coalfield, UK. Geological Magazine 144, 465–486. https://doi.org/10.1017/S0016756807003305

Cleal, C. J., B. A. Thomas. 2005. Palaeozoic tropical rainforests and their effect on global climates: is the past the key to the present? Geobiology, 3: 13–31. https://doi.org/10.1111/j.1472-4669.2005.00043.x

Cleal, C. J., S. Opluštil, B. A. Thomas, Y. Tenchov. 2011. Pennsylvanian vegetation and climate in tropical Variscan Euramerica. Episodes, 34: 3–12. https://doi.org/10.18814/epiiugs/2011/v34i1/002

Cleal, C. J., D. Uhl, B. Cascales-Miñana, B. A. Thomas, A. R. Bashforth, [et al.]. 2012. Plant biodiversity changes in Carboniferous tropical wetlands. Earth-Science Reviews, 114: 124–155. https://doi.org/10.1016/j.earscirev.2012.05.004

Cridland, A. A., J. E. Morris. 1963. Taeniopteris, Walchia, and Dichophyllum in the Pennsylvanian system of Kansas. University of Kansas Science Bulletin, 44: 71–82.

Davydov, V.I., J. L. Crowley, M. D. Schmitz, V. I. Poletaev. 2010. High-precision U–Pb zircon age calibration of the global Carboniferous time scale and Milankovitch band cyclicity in the Donets Basin, eastern Ukraine. Geochemistry, Geophysics, Geosystems, 11 (1): 1–22. https://doi.org/10.1029/2009GC002736

DiMichele, W. A. 2014. Wetland-dryland vegetational dynamics in the Pennsylvanian ice age tropics. International Journal of Plant Sciences, 175: 123–164. https://doi.org/10.1086/675235

DiMichele, W. A., R. B. Aronson. 1992. The Pennsylvanian–Permian vegetational transition: a terrestrial analogue to the onshore–offshore hypothesis. Evolution, 46: 807–824. https://doi.org/10.1111/j.1558-5646.1992.tb02086.x

DiMichele, W. A., T. L. Phillips. 1996. Climate change, plant extinctions, and vegetational recovery during the Middle-Late Pennsylvanian transition: the case of tropical peat-forming environments in North America. In: Hart, M. L. (ed.). Biotic Recovery from Mass Extinctions. Geological Society, London, Special Publication, 102: 201–221. https://doi.org/10.1144/GSL.SP.1996.001.01.14

DiMichele, W. A., H W. Pfefferkorn, R. A. Gastaldo. 2001. Response of Late Carboniferous and Early Permian plant communities to climate change. Annual Review of Earth and Planetary Science, 29: 461–487. https://doi.org/10.1146/annurev.earth.29.1.461

DiMichele, W. A., R. A. Gastaldo, H. W. Pfefferkorn. 2005.Plant biodiversity partitioning in the Late Carboniferous and Early Permian and its implications for ecosystem assembly. Proceedings of the California Academy of Sciences, 56 (1), No 4: 32–49.

DiMichele, W. A., N. J. Tabor, D. S. Chaney, W J. Nelson. 2006. From wetlands to wet spots: environmental tracking and the fate of Carboniferous elements in Early Permian tropical floras. In: Greb, S. F., DiMichele, W. A. (eds.). Wetlands through Time. Geological Society of America Special Paper, 399: 223–248.

DiMichele, W. A., H. Kerp, N. J. Tabor, C. V. Looy. 2008. Revisiting the so-called “Paleophytic–Mesophytic” transition in equatorial Pangea: vegetational integrity and climatic tracking. Palaeogeography, Palaeclimatology, Palaeoecology, 268: 152–163. https://doi.org/10.1016/j.palaeo.2008.06.006

DiMichele, W. A., I. P. Montañez, C. J. Poulsen, N. J. Tabor. 2009. Climate and vegetational regime shifts in the late Paleozoic ice age earth. Geobiology, 7: 200–226. https://doi.org/10.1111/j.1472-4669.2009.00192.x

DiMichele, W. A., B. Cecil, I. P. Montañez, H. J. Falcon-Lang. 2010. Cyclic changes in Pennsylvanian paleoclimate and its effects on floristic dynamics in tropical Pangaea. International Journal Coal Geology, 83: 329–344. https://doi.org/10.1016/j.coal.2010.01.007

DiMichele, W. A., A. R. Bashforth, H. J. Falcon-Lang, S. G. Lucas. 2020. Uplands, lowlands, and climate: Taphonomic megabiases and the apparent rise of a xeromorphic, drought-tolerant flora during the Pennsylvanian-Permian transition. Palaeogeography, Palaeoclimatology, Palaeoecology, 559. Article 109965. Advance online publication. https://doi.org/10.1016/j.palaeo.2020.109965

DiMichele, W. A., C. F. Eble, H. W. Pfefferkorn, S. D. Elrick, W. J. Nelson,

. 2023. Kasimovian floristic change in tropical wetlands and the Middle–Late Pennsylvanian Boundary Event. Geological Society, London, Special Publications, 535 (1): 293–335. https://doi.org/10.1144/SP535-2022-228

DiMichele, W. A., S. G. Lucas, C. F. Eble, H. Kerp, S. J. Reynolds, [et al.]. 2024. A detailed stratigraphic and taphonomic reassessment of the late Paleozoic fossil flora from Promontory Butte, Arizona. Review of Palaeobotany and Palynology, 320: 105004. https://doi.org/10.1016/j.revpalbo.2023.105004

Dimitrova, T. K., C. J. Cleal, B. A. Thomas. 2011. Palynological evidence for Pennsylvanian extra-basinal vegetation in Atlantic Canada. Journal of the Geological Society, 168: 559–569. https://doi.org/10.1144/0016-76492010-028

Elrick, S. D., W. J. Nelson, P. R. Ames, W. A. DiMichele. 2017. Floras characteristic of Late Pennsylvanian peat swamps arose in the late Middle Pennsylvanian. Stratigraphy, 14: 123–141. https://doi.org/10.29041/strat.14.1-4.123-141

Eros, J. M., I. P. Montañez, D. A. Osleger, V. I. Davydov, T. I. Nemyrovska, [et al.]. 2012. Sequence stratigraphy and onlap history of the Donets Basin, Ukraine: Insight into Carboniferous icehouse dynamics. Palaeogeography, Palaeoclimatology, Palaeoecology, 313–314: 1–25. https://doi.org/10.1016/j.palaeo.2011.08.019

Gastaldo, R. A, H. W. Pfefferkorn, W. A. DiMichele. 1995. Taphonomic and sedimentologic characterization of roof-shale floras. Memoir of the Geological Society of America, 185: 341–52. https://doi.org/10.1130/MEM185-p341

Gastaldo, R. A. 1996. Flöznah and flözfern assemblages: potential predictors of Late Carboniferous biome replacement? In: Leary, R. L. (ed.). Patterns in Paleobotany: Proceedings of a Czech–U.S. Carboniferous Paleobotany Workshop. Illinois State Museum Scientific Papers, 26:19–27.

Gorak, S.V., V.I. Poletaev. 1993. Hercynian cycle of tectonic history of Ukraine (Middle Devonian-Permian). In: Tsehelnyuk, P.D. (ed.). Geological history of the territory of Ukraine. Paleozoic. Naukova Dumka,Kyiv, 69–75. [in Russian]

Havlena, V. 1971. Die zeitgleichen Floren des europäischen Oberkarbons und die mesophile Floras des Ostrau-Karwiner Steinkohlenreviers. Review of Palaeobotany and Palynology, 12: 245–270. https://doi.org/10.1016/0034-6667(71)90015-7

Heckel, P. H. 2008. Pennsylvanian cyclothems in Midcontinent North America as farfield effects of waxing and waning of Gondwana ice sheets. In: Fielding, C. R., Frank, T. D., Isbell, J. L. (eds.). Resolving the late Paleozoic Ice Age in Time and Space. Geological Society of America Special Paper, 441: 275–289. https://doi.org/10.1130/2008.2441(19)

Horton, D. E, C. J. Poulsen, D. Pollard. 2010. Influence of high-latitude vegetation feedback on late Paleozoic glacial cycles. Nature Geoscience, 3: 572–577. https://doi.org/10.1038/ngeo922

Isbell, J. L., L. C. Henry, E. L. Gulbranson, C. O. Limarino, M. L. Fraiser, [et al.]. 2012. Glacial paradoxes during the late Paleozoic ice age; evaluating the equilibrium line altitude as a control on glaciation. Gondwana Research, 22: 1–19. https://doi.org/10.1016/j.gr.2011.11.005

Falcon-Lang, H. J. 2003. Response of Late Carboniferous tropical vegetation to transgressive–regressive rhythms at Joggins, Nova Scotia. Journal of the Geological Society, 160: 643–648. https://doi.org/10.1144/0016-764902-114

Falcon-Lang, H. J. 2004. Pennsylvanian tropical rain forests responded to glacialinterglacial rhythms. Geology, 32: 689–692. https://doi.org/10.1130/G20523.1

Falcon-Lang, H. J., W. J. Nelson, S. Elrick, C. V. Looy, P R. Ames, [et al.]. 2009. Incised channel fills containing conifers indicate that seasonally dry vegetation dominated Pennsylvanian tropical lowlands. Geology, 37: 923–926. https://doi.org/10.1130/G30117A.1

Falcon-Lang, H. J., W. J. Nelson, P. H. Heckel, W. A. DiMichele, S. D. Elrick. 2018. New insights on the stepwise collapse of the Carboniferous Coal Forests: evidence from cyclothems and coniferopsid tree-stumps near the Desmoinesian–Missourian boundary in Peoria County, Illinois, USA. Palaeogeography, Palaeoclimatology, Palaeoecology, 490; 375–392. https://doi.org/10.1016/j.palaeo.2017.11.015

Feofilova, A. P. 1966. Transition of coal­bearing deposits to salt­bearing in western part of the Donets Basin. Nauka, Moscow, 1–176. [In Russian]

Fielding, C. R., T. D. Frank, J. L. Isbell. 2008. The late Paleozoic ice age — A review of current understanding and synthesis of global climate patterns. In:Fielding, C. R., Frank, T. D., Isbell, J. L. (eds). Resolving the Late Paleozoic Ice Age in Time and Space. Geological Society of America Special Paper, 441: 343–354. https://doi.org/10.1130/2008.2441(24)

Fisunenko, O. P. 1975. The Donets Basin as a floristic standard of the Carboniferous in the south of the European part of the USSR. In: Timofeev, P. P. (ch. ed.). Stratigraphy of the Carboniferous and geology of coal-bearing formations of the USSR: Proceedings of the VII International Congress on Stratigraphy and Geological of the Carboniferous. Nedra, Moskow, 90–101. [In Russian]

Fisunenko, O. P. 1991. Zonal phytostratigraphical scale of the Lower and Middle Carboniferous of the Donets Basin. Geologichnyj zhurnal, 3 (258): 55–64. [In Russian]

Fisunenko, O. P. 2000. On the problem of the Moscovian Stage. Izdatelstvo LGPU, Lygansk, 1–66. [In Russian]

Looy, C. V., H. Kerp, I. A. P. Duijnstee, W. A. DiMichele. 2014. The Late Paleozoic ecological-evolutionary laboratory, a land-plant fossil record perspective. Sedimentary Record, 12: 4–10. https://doi.org/10. 2110/sedred.2014.4.4

Looy, C. V., R. A. Stevenson, T. B. Van Hoof, L. Mander. 2014. Evidence for coal forest refugia in the seasonally dry Pennsylvanian tropical lowlands of the Illinois Basin, USA. PeerJ, 2: e630. https://doi.org/10.7717/peerj.630 

Lyons, P. C., W. C. Darrah. 1989. Earliest conifers in North America: upland and/or paleoecological indicators? Palaios, 4: 480–486. https://doi.org/10.2307/3514592

Mirkin, B. M., L. G. Naumova. 2012. Current state of basic concepts in plant science. Gilem, Ufa, 1–488. [In Russian]

Montañez, I. P. 2022. Current synthesis of the penultimate icehouse and its imprint on the Upper Devonian through Permian stratigraphic record. In: Lucas, S. G., Schneider, J. W., Wang, X., Nikolaeva, S. (eds.). The Carboniferous Timescale. Geological Society, London, Special Publications, 512: 213–245. https://doi.org/10.1144/SP512-2021-124

Montañez, I. P., C. J. Poulsen. 2013. The late Paleozoic ice age: an evolving paradigm. Annual Review of Earth and Planetary Science: 629–656. https://doi.org/10.1146/annurev.earth.031208.100118

Montañez, I. P., N. J. Tabor, D. Niemeier, W. A. DiMichele, T. D. Frank, [et al.]. 2007. CO2-forced climate instability and linkages to tropical vegetation during Late Paleozoic deglaciation. Science, 315: 87–91. https://doi.org/10.1126/science.1134207

Opluštil, S., C. J. Cleal. 2007. A comparative analysis of some Late Carboniferous basins of Variscan Europe. Geological Magazine, 144: 417–448. https://doi.org/10.1017/S0016756807003330

Opluštil, S., Z. Šimůnek, J. Zajíc, V. Mencl. 2013. Climatic and biotic changes around the Carboniferous/Permian boundary recorded in the continental basins of the Czech Republic. International Journal of Coal Geology, 119: 114–151. https://doi.org/10.1016/j.coal.2013.07.014

Peyser C. E., C. J. Poulsen. 2008. Controls on Permo-Carboniferous precipitation over tropical Pangaea: a GCM sensitivity study. Palaeogeography, Palaeoclimatology, Palaeoecology, 268: 181–192. https://doi.org/10.1016/j.palaeo.2008.03.048

Pfefferkorn, H. W. 1980. A note on the term “upland flora”. Review of Palaeobotany and Palynology, 30: 157–158. https://doi.org/10.1016/0034-6667(80)90011-1

Pfefferkorn, H. W, M. Thomson. 1982. Changes in dominance patterns in Upper Carboniferous plant-fossil assemblages. Geology, 10: 641– 44. https://doi.org/10.1130/0091-7613(1982)10<641:CIDPIU>2.0.CO;2

Pfefferkorn, H. W, R. A. Gastaldo, W A. DiMichele. 2017. Impact of an icehouse climate interval on tropical vegetation and plant evolution. Stratigraphy, 14 (1–4): 365–376. https://doi.org/10.29041/strat.14.1-4.365-376

Phillips, T. L., R. A. Peppers. 1984. Changing patterns of Pennsylvanian coal-swamp vegetation and implications of climatic control on coal occurrence. International Journal of Coal Geology, 3: 205–255. https://doi. org/10.1016/0166-5162(84)90019-3

Poletaev, V. I., M. V. Vdovenko, O. K. Shchogolev, N. I. Boyarina, I. A. Makarov. 2011. Stratotypes of the Carboniferous and Lower Permian regional stratigraphic subdivisions of the Don-Dnieper Depression. Logos,Kyiv, 1–236. [In Ukrainian]

Richey, J. D, I. P. Montañez, Y. Goddéris, C. V. Looy, N. P. Griffis, W. A. DiMichele. 2020. Influence of temporally varying weatherability on CO₂-climate coupling and ecosystem change in the late Paleozoic. Climate of the Past, 16 (5): 1759–1775. https://doi.org/10.5194/cp-16-1759-2020

Richey, J. D., I P. Montañez, J. D. White, W. A. DiMichele, W. Matthaeus, [et al.]. 2021. Modeled physiological mechanisms for observed changes in the late Paleozoic plant fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology, 562: 110056. https://doi.org/10.1016/j.palaeo.2020.110056

Scott, A. C. 1977. A review of the ecology of Upper Carboniferous plant assemblages with new data from Strathclyde. Palaeontology, 20: 447–73.

Shchegolev, A. K. 1964. Differentiation of vegetation in the late Carboniferous of the Westphalian Province. Questions of regularities and forms of development of the organic world: Proceedings of the VII session of the VPO. Nedra, Moscow, 158–170.

Shchegolev, A. K. 1965. Flora at the Carboniferous-Permian boundary in the Donetsk Basin. In: Geology of coal-bearing formations and stratigraphy of the Carboniferous of the USSR. Nauka, Moscow, 234–243. [In Russian]

Shchegolev, A. K.1975. Evolution of the flora and vegetation of the territory of the south of the European part of the USSR from the end of the Middle Carboniferous to the beginning of the Permian. The volume and division of the upper, Stephanian, division of the Carboniferous system. In: Timofeev, P. P. (ch. ed.). Stratigraphy of the Carboniferous and geology of coal-bearing formations of the USSR: Proceedings of the VII International Congress on Stratigraphy and Geological of the Carboniferous. Nedra, Moskow, 101–108. [In Russian]

Shchegolev, A. K. 1991. Lycopsida and Sphenopsida of the late Carboniferous. Naukova dumka, Kiev, 1–128. [in Russian]

Stratigraphyof the Upper Proterozoic and Phanerozoic of Ukraine in two volumes. Vol. 1: Stratigraphy of the Upper Proterozoic, Paleozoic and Mesozoic of Ukraine. 2013. Gozhyk, P. F. (Chief ed.). Logos, Kiev, 1–637. [In Ukrainian]

Sukachev, V. N. 1928. Plant communities (Introduction to phytosociology). Kniga, Moscow-Leningrad, 1–232. [In Russian]

Sukachev, V. N. 1954. Some general theoretical questions of phytocoenology. Botany issues, 1: 291–309. [In Russian]

Tabor, N. J, C. J. Poulsen. 2008. Palaeoclimate across the Late Pennsylvanian–Early Permian tropical palaeolatitudes: a review of climate indicators, their distribution, and relation to palaeophysiographic climate factors. Palaeogeography, Palaeoclimatology, Palaeoecology,268, 293–310. https://doi.org/10.1016/j.palaeo.2008.03.052

Thomas, B. A., C. J. Cleal. 2017. Distinguishing Pennsylvanian-age lowland, extrabasinal and upland vegetation. Palaeobiodiversity and Palaeoenvironments, 97: 273–293. https://doi.org/10.1007/s12549-017-0277-0

Van der Maarel, E. 1988. Vegetation dynamics: patterns in Time and Space. Vegetatio, 77: 7–19. https://doi.org/10.1007/BF00045745

Wagner, R. H., 1997. Floral palaeoecology of the Carboniferous/Permian. In: Aguirre, E., Morales, J., Soria, D. (eds.). Registros Fósiles e Historia de la Tierra. Cursos de Verano de El Escorial, Editorial Complutense, Madrid, 143–172.

Willard, D. A., T. L. Phillips, A. D. Lesnikowska, W. A. DiMichele. 2007. Paleoecology of the Late Pennsylvanian-age Calhoun coal bed and implications for long-term dynamics of wetland ecosystems. International Journal of Coal Geology, 69: 21–54. https://doi.org/10.1016/j.coal.2006.03.011

Zhemchuzhnikov, Y. A., V. S. Yablokov, L. I. Bogolioubova, L. I. Botvinkina, A. P. Feofilova, [et al.]. 1960. Structure and environment of the main coal-bearing suites and coal seams of the middle Carboniferous of the Donets Basin. II. Trudy Geological Institute Akademii Nauk SSSR, 15: 1–347. [In Russian]