Analysis of the Eurasian steppe flora – species inventory, species richness and rates of endemism
UDC 581.93(212.6)(4/5)
Abstract
The Eurasian steppe belt extends from the Danube to the Amur River and represents the largest expanse of steppe vegetation in the world. Despite numerous studies of the flora of individual regions, there is no a reliable estimate of the total number of higher plant species characteristic of this biome as a whole. Filling this gap is the aim of this study. Based on an analysis of modern floristic summaries, a count of species associated with steppe ecosystems was conducted, with an emphasis on the Euro-Siberian and Mongol-Chinese subregions. Special attention is paid to the correct assignment of taxa to the steppe flora, taking into account possible errors in taxonomic status and anthropogenic landscape changes. Estimates of the number of steppe species vary: for northern Kazakhstan – 750–850, for Mongolia – 800–900. Our calculations revealed that the species composition of the Pontic steppe ranges from 200 to 400 species, while for the Kazakh, Mongolian, and Chinese steppe regions, it ranges from 700 to 900 species. The rate of endemism in the ‘lowland’ steppes was extremely low (less than 1 %), which contrasts with the high rates in mountainous provinces such as Anatolia and Altai. This is primarily explained by the history of climate change in the Late Pleistocene, as well as the fact that the cryoarid conditions of glacial periods contributed to the impoverishment of flora and decreased isolation, hindering speciation. The results emphasize the dominant role of paleoclimatic factors in shaping the modern floristic structure of the Eurasian steppes and point to the need for further comprehensive research, particularly in the poorly studied regions of Central Asia.
Downloads
References
Aćić S., Šilc U., Petrović M., Tomović G., Stevanović Z. D. 2015. Classification, ecology and biodiversity of Central Balkan dry grasslands. Tuexenia 35: 329–353.
Akgün F., Kayseri M. S., Akkiraz M. S. 2007. Palaeoclimatic evolution and vegetational changes during the Late Oligocene–Miocene period in the Western and Central Anatolia (Turkey). Palaeogeogr. Palaeoclimatol. Palaeoecol. 253: 56–106. https://doi.org/10.1016/j.palaeo.2007.03.034
Alibekov D., Kubentayev S., Baasanmunkh S., Mukhtubayeva S., Tustubayeva S., Kubentayeva B., Suleimen Y., Choi H. J. 2025. The first complete checklist of vascular plants in the Akmola region, Kazakhstan: A key to understanding biodiversity importance. J. Asia-Pacific Biodiversity 18(3): 618–627. https://doi.org/10.1016/j.japb.2025.03.002
Ambarh D., Zeydanh U. S., Balkız Ö., Aslan S., Karaçetin E., Sözen M., et al. 2016. An overview of biodiversity and conservation status of steppes of the Anatolian biogeographical region. Biodivers. Conserv. 25: 2491–2519.
Baasanmunkh S., Urgamal M., Oyuntsetseg B., Grabovskaya‐Borodina A., Oyundelger K., Tsegmed Z. et al. 2021. Updated checklist of vascular plants endemic to Mongolia. Diversity 13, 12: 619. https://doi.org/10.3390/d13120619
Baasanmunkh S., Urgamal M., Oyunstsetseg B., Sukhorukov A. P., Tsegmed Z., Son D. C., Erst A., Oyundelger K., Kechaykin A. A., Norris J., Kosachev P. A., Ma J. S., Chang K. S., Choi H. J. 2022. Flora of Mongolia: annotated checklist of native vascular plants. Ptytokeys 192: 63–169. https://doi.org/10.3897/phytokeys.192.79702
Baryshnikov G. F., Markova A. K. 1992. Map of Main Mammal Assemblages during maximum cooling of the last glaciation about 24.000 to 12.000 yr B. P. and explanatory notes to the map. In: B. Frenzel, M. Pécsi, A. A. Velichko (eds.). Atlas of Paleoclimates and Paleoenvironments of the Northern Hemisphere, Late Pleistocene – Holocene. Budapest, Stuttgart: Gustav-Fischer-Verlag. Pp. 60–61, 127–129.
Biurrun I., Pielech R., Dembicz I., Gillet F., Kozub Ł., Marcenò C., et al. 2021. Benchmarking plant diversity of Palaearctic grasslands and other open habitats. Journal of Vegetation Science 32(4): e13050. https://doi.org/10.1111/jvs.13050
Borovyk D., Dembicz I., Dengler J., Guarino R., Kuzemko A., Lavrinenko K., et al. 2024. Plant species richness records in Ukrainian steppes. Tuexenia 44: 225–239. https://dx.doi.org/10.14471/2024.44.002
Bruchmann I. 2011. Plant endemism in Europe: spatial distribution and habitat affinities of endemic vascular plants. Dissertation. Flensburg, Germany: University of Flensburg. 258 pp. URL: www.zhb-flensburg.de/dissert/bruchmann
Bruchmann I., Hobohm C. 2014. Factors that create and increase endemism. In: C. Hobom (ed.). Endemism in vascular plants. Plant and Vegetation. Vol. 9. Ch. 3. Springer: Science+Business Media Dordrecht. Pp. 51–68. https://doi.org/10.1007/978-94-007-6913-73
Divíšek J., Hájek M., Jamrichová E., Petr L., Večeřa M., Tichý L., Willner W., Horsák M. 2020. Holocene matters: Landscape history accounts for current species richness of vascular plants in forests and grasslands of eastern Central Europe. J. Biogeogr. 47: 721–735. https://doi.org/10.1111/jbi.13787
Du C., Liao S., Boufford D. E., Ma J. 2020. Twenty years of Chinese vascular plant novelties, 2000 through 2019. Plant Diversity 42, 393–398. https://doi.org/10.1016/j.pld.2020.08.004
Fedorov An. A. (ed.-in chief). (1988–2007). Flora of Russia. The European Part and Bordering Regions. Vol. 1–11. Rotterdam, Brookfield: Balkema A. A.
Flora Altaica. 2005. R. V. Kamelin (ed.). Vol. 1. Barnaul: Azbuka. 340 pр. [In Russian] (Флора Алтая. Т. 1. Под ред. Р. В. Камелина. Барнаул: Азбука, 2005. 340 с.).
Frenzel B. 1992. Vegetation during the maximum cooling of the last glacial. Map and explanatory notes. In: B. Frenzel, M. Pécsi, A. A. Velichko (eds.). Atlas of Paleoclimates and Paleoenvironments of the Northern Hemisphere, Late Pleistocene – Holocene. Budapest, Stuttgart: Gustav-Fischer-Verlag. Pp. 55, 122.
Grichuk V. P. 1992. Vegetation during the last interglacial. In: B. Frenzel, M. Pécsi, A. A. Velichko (eds.). Atlas of Paleoclimates and Paleoenvironments of the Northern Hemisphere, Late Pleistocene – Holocene. Budapest, Stuttgart: Gustav-Fischer-Verlag. Pp. 11, 85.
Грубов В. И. Определитель сосудистых растений Монголии (с атласом). Л.: «Наука», 1982. 441 с.
Губанов И. А. Конспект флоры Внешней Монголии (сосудистые растения). Под ред. Р. В. Камелина. М.: «Валанг», 1996. 136 с.
Hájek M., Hájková P., Roleček J. 2020. A novel dataset of perma nent plots in extremely species-rich temperate grasslands. Folia Geobotanica 55: 257–268. https://doi.org/10.1007/s12224-020-09372-6
Hurka H., Friesen N., Bernhardt K.-G., Neuffer B., Smirnov S. V., Shmakov A. I., Blattner F. R. 2019. The Eurasian steppe belt: status quo, origin and evolutionary history. Turczaninowia 22, 3: 5–71. https://doi.org/10.14258/turczaninowia.22.3.1
Yavuz-Işık N., Toprak V. 2010. Palynostratigraphy and vegetation characteristics of Neogene continental deposits interbedded with the Cappadocia ignimbrites (Central Anatolia, Turkey). Internat. J. Earth Sciences 99: 1887–1897. https://doi.org/10.1007/s00531-009-0486-5
Kamelin R. V. 2005. New flora of Altai (aims and conception of new floristic revision). In: R. V. Kamelin (ed.). Flora Altaica. Vol. 1. Barnaul: Azbuka. Pp. 55–97. [In Russian and Englisch]
Kamp J., Koshkin M. A., Bragina T. M., Katzner T. E., Milner-Gulland E. J., Schreiber D., Sheldon R., Shmalenko A., Smelansky I., Terraube J., Urazaliev R. 2016. Persistent and novel threats to the biodiversity of Kazakhstan’s steppes and semi-deserts. Biodivers. Conserv. 25: 2521–2541. https://doi.org/10.1007/s10531-016-1083-0
Комаров В. Л. (гл. ред.) Флора СССР. Т. 1–30. М.; Л.: Изд-во АН СССР, 1934–1964.
Kuzemko A. A., Steinbauer M. J., Becker T., Didukh Ya. P., Dolnik Ch., Jeschke M., Naqinezhad A., Uğurlu E., Vassilev K., Dengler J. 2016. Patterns and drivers of phytodiversity in steppe grasslands of Central Podolia (Ukraine). Biodivers. Conserv. 25: 2233–2250. https://doi.org/10.1007/s10531-016-1060-7
Королюк А. Ю. Степи Северного Казахстана – синтаксономическая ревизия // Растительность России, 2017. № 30. С. 61–77. https://doi.org/10.31111/vegrus/2017.30.61
Красноборов И. М., Артемов И. А. (отв. ред.). Определитель растений Республики Алтай. Новосибирск: Изд-во СО РАН, 2012. 701 с.
Куприянов А. Н. Конспект флоры Казахского мелкосопочника. Новосибирск: Акад. изд-во «Гео», 2020. 423 с.
Kürschner H., Parolly G. 2012. The Central Anatolian steppes. In: M. J. A. Werger, M. A. van Staalduinen (eds.). Eurasian Steppes. Ecological Problems and Livelihoods in a Changing World, Plant and Vegetation 6. Ch. 4. Springer: Science+Business Media B. V. Pp. 149–171. https://doi.org/10.1007/978-94-007-3886-7_4
Lavrenko E. M. 1969. Über die Lage des Euroasiatischen Steppengebiets in dem System der pflanzengeographischen Gliederung des außertropischen Eurasiens. Vegetatio 19: 11–20.
Лавренко Е. М. Провинциальное разделение При черноморско-Казахстанской подобласти Степной области Евразии // Бот. журн., 1970. Т. 55, № 5. С. 609–625.
Лавренко Е. М. Провинциальное разделение Центральноа зиатской подобласти Степной области Евразии // Бот. журн., 1970. Т. 55, № 12. С. 1734–1741.
Lavrenko E. M., Karamysheva Z. V. 1993. Steppes of the former Soviet Union and Mongolia. In: R. T. Coupland (ed.). Natural Grasslands. Eastern Hemisphere and Resume. Amsterdam: Elsevier. Pp. 3–59.
Лавренко Е. М., Карамышева З. В., Никулина Р. И. Степи Евразии. Л.: Наука, 1991. 146 с.
Li J., He B., Zhou S., Zhang X., Li C., Han G. 2022. Prediction of plant diversity under different stocking rates based on functional traits of constructive species in a desert steppe, northern China. Front. Ecol. Evol. 10: 865703.
Li W., Tojibaev K. S., Hisoriev H., Shomurodov K. F., Luo M., Feng Y., Ma K. 2020. Mapping Asia Plants: Current status of floristic information for Central Asian flora. Global Ecology and Conservation 24: e01220. https://doi.org/10.1016/j.gecco.2020.e01220
Liu D., Liu G., Chen L., Han W., Wang D. 2022. Plant diversity is coupled with soil fungal diversity in a natural temperate steppe of northeastern China. Soil Ecology Letters 4: 454–469. https://doi.org/10.1007/s42832-021-0113-3
Liu G., Xie X., Ye D., Ye X., Tuvshintogtokh I., Mandakh B., Huang Zh., Dong M. 2013. Plant functional diversity and species diversity in the Mongolian Steppe. PLoS ONE 8(10): e77565. https://doi.org/10.1371/journal.pone.0077565
Liu H., Cui H. 2009. Patterns of plant biodiversity in the woodland-steppe ecotone in southeastern Inner Mongolia. Contemporary Problems Ecol. 2: 322–329. https://doi.org/10.1134/S1995425509040043
Luo W., Zhao W., Liu B. 2016. Growth stages affect species richness and vegetation patterns of nebkhas in the desert steppes of China. Catena 137: 126–133. http://dx.doi.org/10.1016/j.catena.2015.09.011
Ma Z. G., Ren X. B. 2007. Drying trend over China from 1951 to 2006. Adv. Clim. Change Res. 3(4): 195–201. [In Chinese]
Malyshev L. I. (ed.-in-chief). 2000–2008. Flora of Siberia. Vol. 1–14. Enfield, NH: Science Publishers.
Merunková K., Preislerová Z., Chytrý M. 2012. White Carpathian grasslands: Can local ecological factors explain their extraordinary species richness? Preslia 84(2): 311–325.
Molnár Á. P., Demeter L., Biró M., Chytrý M., Bartha S., Gantuya B., Molnár Z. 2023. Is there a massive glacial–Holocene flora continuity in Central Europe? Biol. Rev. 98: 2307–2319.
Намзалов Б. Б. Степи Тувы и Юго-Восточного Алтая. Новосибирск: Изд-во «Гео», 2015. 294 с.
Palpurina S., Chytrý M., Tzonev R., Danihelka J., Axmanová I., Merunková K., Karakiev T. 2015. Patterns of fine-scale plant species richness in dry grasslands across the eastern Balkan Peninsula. Acta Oecologica 63: 36–46. https://doi.org/10.1016/j.actao.2015.02.001
Павлов Н. В. Принципы составления «Флоры…», сокращения и обозначения // Флора Казахстана. Т. 1. Алма-Ата: изд-во АН КазССР, 1956. С. 19–54.
Павлов Н. В. (гл. ред.). Флора Казахстана. Т. 1–9. Алма-Ата: изд-во АН КазССР, 1956–1966.
Пешкова Г. А. Флорогенетический анализ степной флоры гор Южной Сибири. Новосибирск: Наука, 2001. 192 c.
Pils G. 2013. Endemism in mainland regions – case studies: Turkey. In: C. Hobohm (ed.). Endemism in vascular plants. Dordrecht: Springer. Pp. 311–321. https://doi.org/10.1007/978-94-007-6913-7
Полуянов А. В. Флора Курской области. Курск: Курск. гос. ун-т, 2005. 264 с.
Polyakova M. A., Dembicz I., Becker T., Becker U., Demina O. N., Ermakov N., Filibeck, G., Guarino R., Janišová M., Jaunatre R., Kozub L., Steinbauer M. J., Suzuki K., Dengler J. 2016. Scale- and taxon-dependent patterns of plant diversity in steppes of Khakassia, South Siberia (Russia). Biodivers. Conserv. 25: 2251–2273. https://doi.org/10.1007/s10531-016-1093-y
Pyak A. I., Shaw S. C., Ebel A. L., Zverev A. A., Hodgson J. G., Wheeler B. D., Gaston K. J. 2008. Endemic plants of the Altai Mountain Country. Old Basing: Wild Guides Ltd. 368 pp.
Qian H., Qian S., Zhang J., Kessler M. 2024. Effects of climate and environmental heterogeneity on the phylogenetic structure of regional angiosperm floras worldwide. Nat. Commun. 15: 1079. https://doi.org/10.1038/s41467-024-45155-9
Rachkovskaya E. I., Bragina T. M. 2012. Steppes of Kazakhstan: Diversity and present state. In: M. J. A. Werger, M. A. van Staalduinen (eds.). Eurasian Steppes. Ecological Problems and Livelihoods in a Changing World. Plant and Vegetation 6. Ch. 3. Springer: Science+Business Media B. V. Pp. 103–148. https://doi.org/10.1007/978-94-007-3886-7_3
Roleček J., Dřevojan P., Hájkova P., Willner W., Janišová M., Lengyel A., Chorney I., Kuzemko A., Goia I., Vassilev K., Hájek M. 2025. Peri-Carpathia forest-steppe grasslands: Distribution, indicator species and extreme species richness. J. Biogeogr. 52(3): 712–721. https://doi.org/10.1111/jbi.15069
Sandel B., Weigelt P., Kreft H., Keppel G., van der Sande M. T., Levin S., et al. 2020. Current climate, isolation and history drive global patterns of tree phylogenetic endemism. Glob. Ecol. Biogeogr. 29: 4–15. https://doi.org/10.1111/geb.13001
Sârbu I., Ştefan N., Oprea A. 2013. Plante Vasculare din România. Bucharest: Victor B Victor. 1320 pp.
Shapoval V. V. 2012. Flora of vascular plants of the Askania Nova steppe reserve. Armyansk: Andreev Publ. House. 195 pp. [In Ukrainian]
Schroeder F.-G. 1998. Lehrbuch der Pflanzengeographie. Wiesbaden: Quelle & Meyer. 457 pp.
Shmalenko A., Smelansky I., Terraube J., Urazaliev R. 2016. Persistent and novel threats to the biodiversity of Kazakhstan’s steppes and semi-deserts. Biodivers. Conserv. 25: 2521–2541. https://doi.org/10.1007/s10531-016-1083-0
Strömberg C. A. E., Werdelin L., Friies E. M., Sarac G. 2007. The spread of grass-dominated habitats in Turkey and surrounding areas during the Cenozoic: Phytolith evidence. Palaeogeogr. Palaeoclimatol. Palaeoecol. 250: 18–49. https://doi.org/10.1016/j.palaeo.2007.02.012
Takhtajan A. 1986. Floristic Regions of the World. Berkeley; Los Angeles, London: University of California Press. 522 pp.
Tsogbadral K. 2021. Biogeographical characteristics of Mongolia. In: B. Yembuu (ed.). Physical Geography of Mongolia. Springer. Pp. 161–176.
Turtureanu P. D., Palpurina S., Becker T., Dolnik C., Ruprecht E., Sutcliffe L. M. E., Szabó A., Dengler J. 2014. Scale- and taxon-dependent biodiversity patterns of dry grassland vegetation in Transylvania. Agric. Ecosyst. Environ. 182: 15–24. http://dx.doi.org/10.1016/j.agee.2013.10.028
Tutin T. G., Heywood V. H., Burges N. A., Moore D. M., Valentine D. W., Walters S. M., Webb D. A. (eds.). 1964–1980. Flora Europaea. Vol. 1–5. Cambridge: Cambridge University Press.
Tuvshintogtokh I. 2014. The steppe vegetation of Mongolia. Ch. Sanchir (ed.). Ulaanbaatar: Bembi San. 610 pp. [In Mongolian with English resume]
Urgamal M., Oyuntsetseg B., Nyambayar D., Dulamsuren C. 2014. Conspectus of the vascular plants of Mongolia. Ulaanbaatar, Mongolia: Admon Printing. 332 pp.
Van der Maarel E., Titlyanova A. 1989. Above-ground and below-ground biomass relations in steppes under different grazing conditions. Oikos 56: 364–370.
Vasilevich V. I. 2009. Dry grass pine forests of Eastern Europe. Bot. Zhurn. 94(11): 1601–1613. [In Russian] (Василевич В. И. Сухотравные сосняки Восточной Европы // Бот. журн., 2009. Т. 94, № 11. С. 1601–1613).
Velichko A. A., Isayeva L. L. 1992. Landscapes during the last glacial maximum. In: B. Frenzel, M. Pécsi, A. A. Velichko (eds.). Atlas of Paleoclimates and Paleoenvironments of the Northern Hemisphere, Late Pleistocene – Holocene. Budapest, Stuttgart: Gustav-Fischer-Verlag. Pp. 59, 125–126.
Vynokurov D., Lysenko T., Dutova Z., Shylnikov D., Doroshina G., Urbanavichene I., Urbanavichus G., Tsepkova N. 2021. The Dry grasslands (Festuco–Frometea) of the North Caucasus: first data on numerical classification and biodiversity patterns. Tuexenia 41: 175–201. https://doi.org/10.14471/2021.41.004
Wang X. X., Dong S. K., Sherman R., Liu Q. R., Liu S. L., Li Y. Y., Wu Y. 2015. A comparison of biodiversity-ecosystem function relationships in alpine grasslands across a degradation gradient on the Qinghai-Tibetan Plateau. Rangeland Journal 37: 45–55.
Wang G. H., Fang J. Y., Guo K., Xie Z. Q., Tang Z. Y., Shen Z. H., Wang R. Q., Wang X. P., Wang D. L. 2020. Contents and protocols for the classification and description of Vegetation Formations, Alliances and Associations of vegetation of China. Chinese Journal of Plant Ecology 44: 128–178. [In Chinese]. https://doi.org/10.17521/cjpe.2019.0272
Wang Z., Meng, P., Wang Z., Lv S., Han G., Hou D., Wang J., Wang H., Zhu A. 2024. Spatial distribution of shrubs and perennial plants under grazing disturbance in the desert steppe of inner mongolia. Glob. Ecol. Conserv. 54: e03193.
Wilson J. B., Peet R. K., Dengler J., Pärtel M. 2012. Plant species rich ness: the world records. Journal of Vegetation Science 23: 796–802. https://doi.org/10.1111/j.1654-1103.2012.01400.x
Wu Z., Raven P. H., Hong D. Y. (eds.). 1994–2013. Flora of China. Vol. 1–25. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press.
Zhao S., Fang J. 2006. Patterns of species richness for vascular plants in China’s nature reserves. Diversity Distrib. 12: 364–372. https://doi.org/10.1111/j.1366-9516.2006.00232.x
Zhu Y., Shan D., Wang B., Shi Z., Yang X., Liu Y. 2019. Floristic features and vegetation classification of the Hulun Buir Steppe in North China: Geography and climate-driven steppe diversification. Global Ecology and Conservation 20: e00741. https://doi.org/10.1016/j.gecco.2019.e00741
Золотухин Н. И., Решетникова Н. М. Флора участка «Острасьевы яры» государственного природного заповедника «Белогорье» // Полевой журнал биолога, 2023. Т. 5, № 3. С. 223–280. https://doi.org/10.52575/2712-9047-2023-5-3-223-280
Turczaninowia is a golden publisher, as we allow self-archiving, but most importantly we are fully transparent about your rights.
Authors may present and discuss their findings ahead of publication: at biological or scientific conferences, on preprint servers, in public databases, and in blogs, wikis, tweets, and other informal communication channels.
Turczaninowia allows authors to deposit manuscripts (currently under review or those for intended submission to Turczaninowia) in non-commercial, pre-print servers such as ArXiv.
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are permitted and encouraged to post their work online (e.g., in institutional repositories or on their website) prior to and during the submission process, as it can lead to productive exchanges, as well as earlier and greater citation of published work (See The Effect of Open Access).
