Terrestrial algae, bryophytes, and lichens of biological soil crusts and corticolous biofilms on the bed of the former Kakhovka Reservoir
DOI:
https://doi.org/10.32999/ksu1990-553X/2026-22-1-3Keywords:
biodiversity, biological soil crusts, takyr, silt, sand, bark of trees, willow forest, war, UkraineAbstract
Question: What is the diversity and role of terrestrial algae, bryophytes and lichens in the initial stages of overgrowth of the former Kakhovka Reservoir bed, in particular, in the formation of soil biocrusts and corticolous biofilms? Location: Kakhovka Reservoir, Dnipropetrovsk, Kherson and Zaporizhzhia regions, Ukraine. Materials and methods: field and laboratory methods of collection and investiga-tion of algae, bryophytes and lichens, direct microscopy and culture methods (Kostikov et al. 2001, Bischoff & Bold 1963, Stanier et al. 1971) Nomenclature: algae and cyanobacteria (Guiry & Guiry 2026), bryophytes (Virchenko & Nyporko 2022), lichens (https://www.indexfungorum.org/) Results: During a three-year study of the exposed bed of the former Kakhovka Res-ervoir (2023-2025), 55 species of terrestrial algae (Cyanobacteria – 17 species, Chlorophyta – 27, Charophyta – 4, Heterokontophyta – 7), 2 species of bryophytes (Marchantia polymorpha and Funaria hygrometrica) and 1 species of lichen (Physcia adscendens) were identified. 50 algal species were found in soil biocrusts. Cyanobacteria and Chlorophyta were the most numerous there, dominating species were Phormidium takyricum, Chlorosarcinopsis cf. aggregata, Stenomitos sp., Microcoleus vaginatus, Klebsormidium cf. flaccidum, etc. On takyr-like (silt) substrates, 30 species of algae were found. Here, in a young willow-poplar forest, the liverwort Marchantia polymorpha was occasionally observed. A greater diversity of algae was found on sands: 36 species. Biocrusts dominated by the moss Funaria hygrometrica were often recorded on sands. 10 species of algae were recorded within the biofilms on willow bark, with green algae prevailing and Chloroidium cf. ellipsoideum as the dominant species. Additionally, the first epiphytic lichen (Physcia adscendens) was recorded at the same substrate. Conclusions: High cyanobacterial diversity of and their dominance in communi-ties are characteristic for soils in the steppe zone of Ukraine. A distinctive feature of biocrusts on silt is the predominance of fine-filamentous cyanobacteria. Biocrusts on sand were characterized by a higher diversity of algae and the occur-rence of aquatic species due to the periodic flooding of these areas. During the three years following the dam destruction, algal diversity in the biocrusts increased considerably. The low species richness in willow bark biofilms and the dominance of green algae are characteristic of aerophytic bark communities in the temperate zone. In the early successional stages of biodiversity recovery, bryophytes and lichens contribute minimally to the development of soil biocrusts and corticolous assemblages of willow trees.
References
Algae of Ukraine: diversity, nomenclature, taxonomy, ecology and geography. Cyanoprocaryota, Euglenophyta, Chrysophyta, Xanthophyta, Raphidophyta, Phaeophyta, Dinophyta, Cryptophyta, Glaucocystophyta, and Rhodophyta (2006). / Ed. by P.M. Tsarenko, S.P. Wasser & E. Nevo. Vol. 1. Liechtenstein: A.R.G. Gantner Verlag, Ruggell, 713 p.
Algae of Ukraine: diversity, nomenclature, taxonomy, ecology and geography. Bacillariophyta (2009). / Ed. by P.M. Tsarenko, S.P. Wasser & E. Nevo. Vol. 2. Liechtenstein: A.R.G. Gantner Verlag, Ruggell, 413 p.
Algae of Ukraine: diversity, nomenclature, taxonomy, ecology and geography. Chlorophyta (2011). / Ed. by P.M. Tsarenko, S.P. Wasser & E. Nevo. Vol. 3. Liechtenstein: A.R.G. Gantner Verlag, Ruggell, 511 p.
Barkmann, J.J. (1958). Phytosociology and ecology of cryptogamic epiphytes. Assen: Van Gorkum, xiii + 628 pp. https://doi.org/10.1017/S0024282971000227
Belnap, J. (2002). Nitrogen fixation in biological soil crusts from southeast Utah, USA. Biology and Fertility of Soils 35: 128–135. https://doi.org/10.1007/s00374-002-0452-x
Belnap, J. (2006). The potential roles of biological soil crusts in dryland hydrologic cycles. Hydrological Processes 20: 3159–3178. https://doi.org/10.1002/hyp.6325
Belnap, J., Büdel, B. & Lange, O.L. (2001). Biological soil crusts: characteristics and distribution. In: Biological soil crusts: structure, function, and management (Belnap, J. & Lange, O.L., editors). Berlin: Springer, 3–30.
Bischoff, H.W. & Bold, H.C. (1963). Phycological studies. IV. Some soil algae from Enchanted Rock and related algal species. University Texas Publications 6318: 1–95.
Bohunicka, M., Pietrasiak, N., Johansen, J.R., Berrendero Gomez, E., Hauer, T., Gaysina, R.A. & Lukesova, A. (2015). Roholtiella, gen. nov. (Nostocales, Cyanobacteria) – a tapering and branching cyanobacteria of the family Nostocaceae. Phytotaxa 197(2): 84–103 https://doi.org/10.11646/phytotaxa.197.2.2
Büdel, B., Dulić, T., Darienko, T., Rybalka, N. & Friedl, T. (2016). Cyanobacteria and algae of biological soil crusts. In Biological Soil Crusts: Structure, Function, and Management (Weber, B., Büdel, B. & Belnap, J., editors). Berlin: Springer, 55–80.
Darienko, T., Gustavs, L., Mudimu, O., Menendes, C., Schumann, R., Karsten, U., Friedl, T. & Pröschold, T. (2010). Chloroidium, a common terrestrial coccoid green alga previously assigned to Chlorella (Trebouxiophyceae, Chlorophyta). European Journal of Phycology, 45: 79–95.
Didukh, Ya.P., Kuzemko, A.A., Khodosovtsev, O.Ye., Chusova, O.O., Borsukevych, L.M., Skobel, N.O., Mikhailyuk, T.I. & Moysiyenko, I.I. (2024). First year of floodplain forest restoration at the bottom of the former Kakhovka reservoir. Chornomorski Botanical Journal 20(3): 305–326. https://doi.org/10.32999/ksu1990-553X/2024-20-3-5
Gaysina, L.A., Bohunická, M., Hazuková, V. & Johansen, J.R. (2018). Biodiversity of terrestrial cyanobacteria of the South Ural region. Cryptogamie Algologie 39(2): 167–198. https://doi.org/10.7872/crya/v39.iss2.2018.167
Grote, E.E., Belnap, J., Housman, D.C., Sparks, J.P. (2010). Carbon exchange in biological soil crusts communities under differential temperatures and soil water contents: implications for global change. Global Change Biology 16: 2763–2774. https://doi.org/10.1111/j.1365-2486.2010.02201.x
Guiry, M.D. & Guiry, G.M. (2026). AlgaeBase. Worldwide electronic publication, Nat. Univ. Ireland, Galway. http://www.algaebase.org
Harper, K.T. & Belnap, J. (2001). The influence of biological soil crusts on mineral uptake by associated vascular plants. Journal of Arid Environments 47: 347–357. https://doi.org/10.1006/jare.2000.0713
Kondratyeva, N.V. (1968). Blue-green algae – Cyanophyta. Guide to the fresh water algae of the Ukrainian SSR. Kyiv, 321 с. (in Ukrainian)
Kostikov, I.Yu., Romanenko, P.O., Demchenko, E.M., Darienko, T.M., Mikhailyuk, T.I., Rybchinskiy, O.V. & Solonenko, A.M. (2001). The soil algae from Ukraine (history and methods of investigation, classification system, floristics). Kyiv: Phytosociocenter, 300 р. (in Ukrainian)
Kuzemko, A.A., Prylutskyi, O.V., Kolomytsev, G.O., Didukh, Ya.P., Moysiyenko, I.I., Borsukevych, L.M., Chusova, O.O., Khodosovtsev, O.Ye. (2025). Initial stages of revegetation at the bottom of the drained Kakhovka Reservoir (Ukraine): synthesis of field surveys and remote sensing. Ukrainian Botanical Journal 82 (5): 488−501. https://doi.org/10.15407/ukrbotj82.05.488
Mikhailyuk, T.І., Vinogradova, O.N., Glaser, K. & Karsten, U. (2016). New taxa for the flora of Ukraine, in the context of modern approaches to taxonomy of Cyanoprokaryota/Cyanobacteria. International Journal on Algae 18(4): 301–320. https://doi.org/10.1615/InterJAlgae.v18.i4.10
Mikhailyuk, T., Glaser, K., Tsarenko, P., Demchenko, E. & Karsten, U. (2019). Composition of biological soil crusts from sand dunes of the Baltic Sea coast, in the context of an integrative approach to the taxonomy of microalgae and cyanobacteria. European Journal of Phycology 54: 263–290.
Mikhailyuk, T.I., Vinogradova, O.M., Demchenko, E.M., Petlovana, V.R., Glaser, K. & Karsten, U. (2025). Terrestrial algae and cyanobacteria of the Holosiiv National Nature Park (Kyiv, Ukraine), with the description of Leptochlorella arboricola sp. nov. (Trebouxiophyceae, Chlorophyta). Ukrainian Botanical Journal 82: 3–30. https://doi.org/10.15407/ukrbotj82.01.003
Munoz-Martin, M.A., Becerra-Absalon, I., Perona, E., Fernandez-Valbuena, L., Garcia-Pichel, F. & Mateo, P. (2019). Cyanobacterial biocrust diversity in Mediterranean ecosystems along a latitudinal and climatic gradient. New Phytologist 221(1): 123–141.
Panou, M. & Gkelis, S. (2022). Unravelling unknown cyanobacteria diversity linked with HCN production. Molecular Phylogenetics and Evolution 166: 107322 https://doi.org/10.1016/j.ympev.2021.107322
Prikhodkova, L.P. (1992). Blue-green algae of soils of Steppe zone of Ukraine. Kyiv: Naukova Dumka. 299 p. (in Russsian)
Schulz, K., Mikhailyuk, T., Dreßler, M., Leinweber, P. & Karsten, U. (2016). Biological soil crusts from coastal dunes at the Baltic Sea: cyanobacterial and algal biodiversity and related soil properties. Microbial Ecology 71: 178–193. https://doi.org/doi:10.1007/s00248-015-0691-7
Stanier, R.Y., Kunisawa, R., Mandel, M. & Cohen-Bazire, G. (1971). Purification and properties of unicellular blue-green algae (order Chroococcales). Bacteriology Reviews, 35: 171–205. https://doi.org/10.1128/br.35.2.171-205.1971
Tsarenko, P.M., Vinogradova, O.M., Burova, O.V., Bryantseva, Yu.V., Borysova, O.V., Lilitska, G.G., Raida, O.V., Beresovska, V.Yu., Kryvosheia-Zakharova, O.M., Sadogurska, S.S. (2024a). Prodromus of spore plants of Ukraine: algae. Book 1. Ed. P.M. Tsarenko. Kyiv: Naukova Dumka, 879 pp. (in Ukrainian) https://doi.org/10.15407/ 978-966-00-1870-9
Tsarenko, P.M., Mikhailyuk, T.I., Burova, O.V., Borysova, O.V., Lilitska, G.G., Demchenko, E.M. (2024b). Prodromus of spore plants of Ukraine: algae. Book 2. Ed. P.M. Tsarenko. Kyiv: Naukova Dumka, 679 pp. (in Ukrainian) https://doi.org/10.15407/978-966-00-1954-6
Van den Acker, J.A.M. & Jungerius, P.D. (1985). The role of algae in the stabilization of coastal dune blowouts. Earth Surface Processes and Landforms 10: 189–192. https://doi.org/doi:10.1002/esp.3290100210
Vinogradova, O.M., Mikhailyuk, T.I., Glaser, K., Holzinger, A. & Karsten, U. (2017). New species of Oculatella (Synechococcales, Cyanobacteria) from terrestrial habitats of Ukraine. Ukrainian Botanical Journal 74(4): 451–465. https://doi.org/doi:10.15407/ukrbotj74.06.509
Vinogradova, O.M., Mikhailyuk, T.I. & Gromakova, A.B. (2019). New and interesting records of cyanobacteria in biological soil crusts from chalk outcrops of Kharkiv Region (Ukraine). VI Int. Conf. «Advances in Modern Phycology» (15–17 May 2019, Kyiv, Ukraine). Abstracts. Kyiv: 122–123.
Virchenko, V.M. & Nyporko, S.O. (2022). Prodromus of Sporen Plants of Ukraine: bryophytes. K.: Naukova Dumka, 176 p. (in Ukrainian)
Wu, Y., Rao, B., Wu, P., Liu, Y., Li, G. & Li, D. (2013). Development of artificially induced biological soil crusts in fields and their effects on top soil. Plant and Soil 370: 115–124. https://doi.org/10.1007/s11104-013-1611-6
