Ergasiophytes of old cemeteries of the Right-Bank of Dnipro Grass Steppe District (Ukraine)
DOI:
https://doi.org/10.32999/ksu1990-553X/2025-21-2-6Keywords:
biodiversity, steppe, cultural heritage objects, old cemeteries, alien species, UkraineAbstract
Questions: What is the species richness, diversity of life forms of ergasiophytes of old cemeteries of the Right-Bank of Dnipro Grass Steppe District? How does the species richness and diversity of life forms of ergasiophytes vary depending on their location in an urbanized environment? Locations: Right-Bank of Dnipro Grass Steppe District (Ukraine) Materials and methods: field observations and herbarium collections Nomenclature: A checklist of vascular plants, bryobionts, and lichens of Ukraine for storing vegetation plots in TURBOVEG (Vynokurov et al. 2025) Results: Every year, the number of ergasiophytes is significantly increases at cultural heritage sites and old cemeteries. The results of comprehensive study on ergasiophytes in the alien fraction of the flora of old cemeteries of the Right-Bank of Dnipro Grass Steppe District are presented. Ergasiophytes include 119 species from 101 genera and 48 families of vascular plants, which is 17.3 % of the spontaneous flora of old cemeteries. The group of ergasiophytes is the most numerous group in terms of the degree of naturalization in the alien flora of old cemeter-ies. A high proportion of ergasiophytes is a characteristic feature of the flora of old cemeteries compared to other cultural heritage sites, including kurgans and ancient settlements. The largest representation of ergasiophytes is observed in old cemeteries located in cities (17.2 %), slightly inferior to them in this indicator are rural cemeteries (16.3 %), while cemeteries located far outside of settlements are characterized by a small participation of ergasiophytes (7.3 %). In addition, old cemeteries closed for burial have twice the percentage of ergasiophytes (8.3 %) compared to active cemeteries (16.8 %). In the spectrum of life forms among ergasiophytes, there is a very high proportion of phanerophytes (39.5 %), especially when compared to natural zonal flora. The sponta-neous spread of trees and shrubs from cultivation sites leads to the dis-placement of steppe plants in cemeteries and outside them, so it is neces-sary to control their spread.
References
Arkushyna, H.F. (2003). About rare species of plants in the flora of cemeteries of Kirovograd. Science and education. Materials of III international research and practice conference, Dnepropetrovsk, 19–20 Martch, 2003: 3–4. (in Ukrainian)
Arkushyna, H.F. (2007). Urban flora of Kirovograd. PhD thesis. Yalta: Nikitskiy botanical garden of Ukrainian Agrarian Academy of Sciences. (in Ukrainian)
Besarabchuk, I. & Volhin, S. (2017). Zoning of the city of Lutsk for comparative urban floristic research. Scientific Bulletin of the Lesia Ukrainka East European National University 7 (357): 61–67. (in Ukrainian)
Burda, R.Y. & Humech, B.C. (1988). Anthropogenic transformation of the urban flora of the city of Zhdanov for last century. Plant introduction and acclimatization 10: 9–14. (in Ukrainian)
Burda, R. I., Pashkevych, N. A., Boiko, H. V. & Fitsailo, T. V. (2015). Alien species of protected floras of the Forest-Steppe of Ukraine. Kyiv: Naukova dumka, 117 p. (in Ukrainian)
Boiko M.F. (2001). Fundamentals of scientific research. Biology: Methodical recommendations. Kherson: Ailant, 12 p. (in Ukrainian)
Dayneko, P., Moysiyenko, I., Dembicz, I., Zachwatowicz, M. & Sudnik–Wójcikowska, B. (2020). Ancient settlements in Southern Ukraine: how do local and landscape factors shape vascular plant diversity patternsin the last remnants of grass steppe vegetation? Tuexenia 40: 459–478. https://doi.org/10.14471/2020.40.015
Dayneko, P., Moysiyenko, I., Sudnik-Wójcikowska, B., Dembicz, I., Zachwatowicz, M. & Skobel, N. (2023). Ancient settlements as natural heritage sites: the first occurrence dataset on vascular plant species from ancient settlements in the Lower Dnipro region (Ukraine). Biodiversity Data Journal 11 https://doi.org/10.3897/bdj.11.e99041
Didukh, Ya.P. & Shelyag–Sosonko, Yu.R. (2003). Geobotanical zoning of Ukraine and adjacent territories. Ukrainian Botanical Journal 60 (1): 6–17. (in Ukrainian)
Dřevojan, P., Čeplová, N., Štěpánková, P. & Axmanová, I. (2023). Life form. Available at: http:/www.FloraVeg.eu. [19/02/2024]
Eremenko, Yu.A. (2013). City cemeteries as places of distribution of adventive scrub species. Plants and urbanization. Materials of III international research and practice conference, Dnepropetrovsk, Martch 19– 20, 2013: 12 (in Ukrainian)
Euro+Med (2024). Euro+Med PlantBase – the information resource for Euro–Mediterranean plant diversity. Available at: http://www.europlusmed.org. [19/02/2024]
FloraVeg.EU (2024). FloraVeg.EU – Database of European Vegetation, Habitats and Flora. https://floraveg.euccessed [15/02/2024].
Gerasimyuk, N.V. (2014). Spring flora of cemeteries of Odessa. Biological Bulletein of Bogdan Khmelnitsky Melitopol State Pedagogical University 4 (1): 170–181. (in Ukrainian)
Gladka, L.V. (ed.). (1969). History of cities and villages of the Ukrainian SSR: Odesa region. K.: URE AN URSR, 930 p. (in Ukrainian)
Gubar, L.M. (2006). Urban flora of the eastern part of Maly Polissya. PhD thesis Kyiv: M.H. Kholodny Institute of Botany. (in Ukrainian)
Hamulia, Yu.H. & Zviahyntseva, K.A. (2010). Features of zoning of habitats of natural and anthropogenic vegetation in Kharkov. The Journal of V.N. Karazin Kharkiv National University. Series: biology 11 (905): 43–54. (in Russian)
Karger, D.N., Conrad, O., Böhner, J., Kawohl, T., Kreft, H., Soria–Auza, R.W., Zimmermann, N.E, Linder, H.P. & Kessler, M. (2017). Climatologies at high resolution for the earth’s land surface areas. Sci Data. http://www.10.1038/sdata.2017.122
Kasyanenko, O.Ye. (ed.). (1972). History of cities and villages of the Ukrainian SSR: Kherson region. K.: URE AN URSR, 688 p. (in Ukrainian)
Kornaś, J. (1981). A geographical–historical classification of synantropic plants. Materialy Zakladu Fitosochologii Stosowanej, UW 25: 33–41.
Krytska, L.Y. (1985). Analysis of the flora of steppes and limestone outcrops of the Right–Bank Grassland Steppe. Ukrainian Botanical Journal 42 (2): 1–5. (in Ukrainian)
Krytska, L.Y. (1987). Flora of steppes and limestone outcrops of the Right-Bank Grassland steppe. PhD thesis, 03.00.05 – botany, abstract. Kyiv: M.G. Kholodny Institute of Botany NASU, 16 p. (in Russian)
Kushnyr, O.A. (2004). Dendroflora of State Historical-Memorial Lukyanivskiy Reserve. Scientific Bulletin Reserve Management in Galychyna, Podolia, and Volhynia 14 (8): 293–296. (in Ukrainian)
Kushnyr, O.A (2006). Bioecological features of plants and formation of design shrubs in cemeteries. DSc thesis, 06.03.00 – forestry. Kyiv: National Agrarian University. (in Ukrainian)
Löki, V., Deák, B., Lukács, A.B. & Molnár, V.A. (2019). Biodiversity potential of burial places – a review on the flora and fauna of cemeteries and churchyards. Global Ecology Conservation 18: 1–14.
Maltseva, C.Yu. (2019). Urban flora of the south-western part of the Northern Priazovye (for example, Berdyansk, Primorsk, Henichesk). PhD thesis, Kyiv: M.M. Gryshko National Botanical Garden. (in Ukrainian)
Malyna, V.V. (2009). Stone crosses in Ukraine. XVIII – XX. Mykolayiv: Khudozhniy kram, 864 p. (in Ukrainian)
Marynych, O.M. & Shyshchenko, P.G. (2005). Physical geography of Ukraine. Kyiv: Znannia, 511 p. (in Ukrainian)
Mosyakin S. L. (ed.). (1999). Vascular Plants of Ukraine. A Nomenclatural Checklist. Kyiv: M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 346 p.
Moysiyenko, I.I. (1997). Flora of Kherson cemeteries. “Problems of botany and mycology on the threshold of the third millennium”. Proceedings of the X Congress of the Ukrainian Botanical Society, Kyiv-Poltava, 1997: 39–40. (in Ukrainian)
Moysiyenko, I.I. (1999). Urban flora of Kherson. PhD thesis, 03.00.05 – botany. Yalta: Nikitskyi Botanical Garden of Ukrainian Agrarian Academy of Sciences. (in Ukrainian) Moysiyenko, I.I. (2011). The Flora of the Northern Prychornomoria Region (Structural Analysis, Synantropization, Conservation). Dsc thesis, 03.00.05 – botany. Kyiv: Taras Shevchenko National University of Kyiv. (in Ukrainian)
Moysiyenko, I., Sudnik-Wójcikowska, B., Dembicz, I., Zachwatowicz, M. & Skobel, N. (2023). The first dataset of vascular plant species occurrences on kurgans in Southern Ukraine. Biodiversity Data Journal 10: https://doi.org/10.3897/bdj.10.e96879
Moysiyenko, I.I. & Sudnik–Wójcikowska, B. (2006a). The flora of kurgans in the desert steppe zone of southern Ukraine. Chornomorski Botanical Journal 2 (1): 5–35. https://doi.org/2010.14255/2308–9628/06.21/1
Moysiyenko, I.I. & Sudnik–Wójcikowska, B. (2006b). The flora of kurgans in the steppe zone of Southern Ukraine – phytogeographical and ecological aspects. Polish Botanical Studies 22: 387–398.
Moysiyenko, I.I. & Sudnik-Wójcikowska, B. (2009). Flora of kurgans in the Pontic herb(–rich) grass steppe zone in Ukraine. Chornomorski Botanical Journal 5 (3): 333–369. https://doi.org/10.14255/2308–9628/09.53/3
Moysiyenko, I.I. & Sudnik–Wójcikowska, B. (2010). Kurgans in Ukraine as a refuge of steppe flora. Bulletin of the Eurasian Dry Grassland Group IAVS 6: 6–10.
Moysiyenko, I.I., Skobel, N.O., Sudnik–Wójcikowska, B., Dembicz, I., Zachwatowicz, M., Zakharova, M.Y., Dzerkal, V.M. (2021). Flora of old cemeteries Lower Dnipro as steppe refuge. Chornomorski Botanical Journal 17 (3): 98–106. https://doi.org/10.32999/ %20ksu1990–553X/2021–17–2–1
Moysiyenko, I.I., Zachwatowicz, M., Sudnik–Wójcikowska, B. & Jabłońska, E. (2014). Kurgans help to protect endangered steppe species in the Pontic grass steppe zone, Ukraine. Wulfenia 21: 83–94.
Mucina, L., Bültmann, H., Dierßen, K., Theurillat, J.-P., Raus, T., Čarni, A., Šumberová, K., Willner, W., Dengler, J., Gavilán García, R., Chytrý, M., Hájek, M., Di Pietro, R., Iakushenko, D., Pallas, J., Daniëls, F.J.A., Bergmeier, E., Santos Guerra, A., Ermakov, N., Valachovič, M., Schaminée, J.H.J., Lysenko, T., Didukh, Ya.P., Pignatti, S., Rodwell, J.S., Capelo, J., Weber, H.E., Solomeshch, A., Dimopoulos, P., Aguiar, C., Hennekens, S.M. & Tichý, L. (2016). Vegetation of Europe: Hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science 19, Suppl. 1: 3–264. https://doi.org/10.1111/avsc.12257
Nowińska, R., Czarna, A. & Kozłowska, M. (2020). Cemetery types and the biodiversity of vascular plants – a case study from south-eastern Poland. Urban Forestry & Urban Greening 49: 1–10. https://doi.org/10.1016/j.ufug.2020.126599
Pashchenko, A.Ya. (ed.). (1969). History of cities and villages of the Ukrainian SSR: Dnipropetrovsk region. K.: URE AN URSR, 958 p. (in Ukrainian)
POWO (2024). Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. Available at: http://www.plantsoftheworldonline.org/Retrieved 19 February 2024. [19/02/2024]
Protopopova, V.V. (1991). Synanthropic flora of Ukraine and ways of its development. Kyiv: Naukova dumka, 204 p. (in Russian)
Protopopova, V.V., Shevera, M.V., Mosyakin, S.L., Solomakha, V.A., Solomakha, T.D., Vasilyeva, T.V. & Petryk, S.P. (2009). Invasive species in the flora of the Northern Black Sea region. Kyiv: Phytosociocenter, 56p. (in Ukrainian)
Protopopova, V.V. & Shevera, M.V. (2013). Ergasiophytes – a potential reserve of the adventive fraction of Ukraine's flora. Non-traditional, new and forgotten plant species: scientific and practical aspects of cultivation – 2013: proceedings of the 1st International Scientific Conference, Ukraine, September 10–12, 2013: 99 –101. (in Russian)
Protopopova, V.V. & Shevera, M.V. (2015). Ergasiophytes of the Ukrainian flora. Biodiversity Research and Conservation 35: 31–36.
Protopopova, V.V., Mosjakin, S. L. & Shevera, M.V. (2002). Plant invasions in Ukraine as a threat to biodiversity: the present situation and tasks for the future. Kyiv: M. G. Kholodny Institute of Botany NASU, 29 p. (in Ukrainian)
Pyšek, P., Prach, K., Reymanek, M. & Wade, M. (1995). Plant invasion. General aspect and special problem. Amsterdam: SPB Academic Publishing, 257 p.
Raunkiaer, C. (1934). The life form of plants and statistical plant geography. Oxford: Clarendon Press, 632 p.
Rubtsov, N.Y., Pryvalova, L.A. & Kryukova, Y.V. (1979). Geographical (arealogical) qualification of species of the flora of Crimea. Yalta: State Nikitsky Botanic Garden, 90 p. (in Russian)
Schubert (1865). Maps of Schubert. Available at: http://www.etomesto.ru/shubert/ [19/02/2024]
Shynder, O. (2017). Rare species in spontaneous flora National Botanic Garden of the NAS of Ukraine. In: The gene pool of botanical gardens and arboretums collections is a pledge of stable phytocoenoses in conditions of climate change. Odesa: ONU, 123–126. (in Ukrainian)
Shynder, O. (2019). Spontaneous flora of M.M. Gryshko National botanical garden of the NAS of Ukraine (Kyiv). 2. Methodological problems and criteria for selection of escaped plants in botanical garden conditions. Plant introduction 82: 3–16.
Skobel, N.O. & Moysiyenko, I.I. (2022). Old cemeteries of Kherson city as refuge of steppe flora Chornomorski Botanical Journal 18 (1): 52–70. (in Ukrainian) https://doi.org/10.32999/ksu1990–553X/2022–18–1–3
Skobel, N., Moysiyenko, I., Sudnik-Wójcikowska, B., Dembicz, I., Zachwatowicz, M., Zakharova, M., Marushchak, O. & Dzerkal, V. (2023). Vascular plants of old cemeteries in the Lower Dnipro region (Southern Ukraine). Biodiversity Data Journal 11: e99004. https://doi.org/10.3897/BDJ.11.e99004
Skobel, N.O., Shapoval, V.V. & Moysiyenko, I.I. (2024). Protected habitats of the old cemeteries of the Right-Bank of Dnipro Grass Steppe District. Chornomorski Botanical Journal 20 (4): 458–470. (in Ukrainian) https://doi.org/10.32999/ksu1990–553X/2024–20–4–6
Sudnik-Wójcikowska, B. & Koźniewska, B. (1988). Słownik z zakresu synantropizacji szaty roślinnej. Warszawa: Wydawnictwa Uniwersytetu Warszawskiego, 93 p. (in Polish)
Sudnik-Wójcikowska, B., Moysiyenko, I., Dembicz, I., Galera, H., Rowińska, A. & Zachwatowicz, M. (2012) Kurgans in the ‘Wild Field’ – a Cultural Heritage and Refugium of the Ukrainian steppe. [Kurhany na „Dzikich Polach” – dziedzictwo kultury i ostoja ukraińskiego stepu/Kurgani “Dikogo polia” – kulturna spadshchina i prikhistok ukrainskogo stepu]. Warsaw:Uniwersytet Warszawski: 194.
Sukhanova, O.A. (2010). Dendroflora of Baykovoe Cemetery. Scientific Bulletin of National University of Biological Resources and Nature 152 (1): 180–184.
Ukraine Incognita (2024). Ancient Cemeteries of Ukraine with Stone Cossack and Ancient Ukrainian Crosses. Available at: https://ukrainaincognita.com/mista/karta–starovynnykh–tsvyntariv–z–kam–ianymy–kozatskymy–khrestamy [19/02/2024]
WorldPop (2020). School of Geography and Environmental Science, University of Southampton; Department of Geography and Geosciences, University of Louisville; Departement de Geographie, Universite de Namur) and Center for International Earth Science Information Network (CIESIN), Columbia University (2018). Global High Resolution Population Denominators Project – Funded by The Bill and Melinda Gates Foundation (OPP1134076).
Vasyliev, V.O. (ed.). (1971). History of cities and villages of the Ukrainian SSR: Mykolaiv region. K.: URE AN URSR, 798 p. (in Ukrainian)
Vynokurov, D., Borovyk, D., Chusova, O., Davydova, A., Davydov, D., Danihelka, J., Dembicz, I., Iemelianova, S., Kolomiiets, G., Moysiyenko, I., Shapoval, V., Shynder, O., Skobel, N., Buzhdygan, O. & Kuzemko, A. (2024) Ukrainian Plant Trait Database: UkrTrait v. 1.0. Biodiversity Data Journal 12: e118128. https://doi.org/10.3897/BDJ.12.e118128
Vynokurov, D., Borovyk, D., Darmostuk, V., Davydov, D., Iemelianova, S. & Danihelka, J. (2025). Ukraine_SL: A checklist of vascular plants, bryobionts, and lichens of Ukraine for storing vegetation plots in TURBOVEG (1.0.0) Data set. Zenodo. https://doi.org/10.5281/zenodo.15192162
