The Role of the Marginal Filter and Climate in Changing the Black Sea Ecosystem

V. V. Melnikov

A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS, Sevastopol, Russia

e-mail: sevlin@ibss-ras.ru

Abstract

The paper presents a review of the current state of knowledge regarding the role of the marginal filter and climate in the ecosystem changes of the Black Sea. The marginal filter (MF) is a key buffer zone at the river–sea interface, where the bulk of terrigenous material delivered from land is deposited and transformed. The MF concept was developed by A.P. Lisitsyn, who demonstrated that the five-stage filter in this contact zone retains 93–95% of suspended particulate matter and about 40% of dissolved substances. The study is the first basin-wide synthesis in which the Lisitsyn concept is applied not for local flux calculations, but as an interpretive tool for understanding the cascade of ecosystem collapses across the entire sea, complementing existing local-scale studies of the Black Sea MF. The functioning of the Black Sea MF as an integrated geoecological buffer system remains poorly understood: this sea is distinguished by its extensive shelf, shallow pycnocline, and hydrogen sulphide zone. The limited volume of oxygenated waters, together with the sea’s dependence on river runoff, renders the MF critically important for the ecosystem. The author hypothesises that the destruction of the biological block of the MF in the 1960s–1980s by intensive bottom trawling, which eradicated the Phyllophora field and benthic filter-feeders, triggered a cascading degradation: a disaster on the shelf (1950s–1980s), a disaster in the open sea (1980–1988), and a collapse of the entire ecosystem (1988–2000). However, the shift to a novel state (>2000) is already linked to climate: warming caused the disappearance of the Cold Intermediate Layer (2019), which had harboured boreal relicts – key forage zooplankton taxa and commercial fish species. Further warming will fundamentally alter the Black Sea ecosystem.

Keywords

Black Sea, marginal filter, deoxygenation, water masses, ecosystem

Acknowledgments

The study was performed under state order on the IBSS RAS topic no. 124030100137-6.

About the authors

Viktor V. Melnikov, Leading Researcher, A. O. Kovalevsky Institute of Biology of the Southern Seas of the Russian Academy of Sciences (299011, Russia, Sevastopol, 2 Nakhimov Ave.), PhD (Biol.), ORCID ID: 0000-0001-6856-0467, ResearcherID: N-9219-2014, Scopus Author ID: 57197396948, sevlin@ibss-ras.ru

For citation

Melnikov, V.V., 2026. The Role of the Marginal Filter and Climate in Changing the Black Sea Ecosystem. Ecological Safety of Coastal and Shelf Zones of Sea, (3), pp. 21–37.

References

  1. Zaitsev, Y.P., 2006. An Introduction on the Black Sea Ecology. Odessa: Even, 224 p. (in Russian).
  2. Gordeev, V.V., 2012. [Geochemistry of the River-Sea System]. Moscow: IP Matushkina I.I., 452 p. (in Russian).
  3. Savenko, V.S., 2006. [The Chemical Composition of Suspended Sediment of the Rivers of the World]. Moscow: GEOS, 173 p. (in Russian).
  4. Lisitsyn, A.P., 1994. A Marginal Filter of the Oceans. Oceanology. 34(5), pp. 735–747 (in Russian).
  5. Miskevich, I.V., Lokhov, A.S. and Chultsova, A.L., 2021. Specificity of Formation of Marginal Filters in Tidal Mountains of Small Rivers in the Winter Period the Example of the River of Solza in the Dvinsky Gulf of the White Sea. Advances in Current Natural Sciences, (7), pp. 54–59. https://doi.org/10.17513/use.37662 (in Russian).
  6. Miskevich, I.V., Kotova, E.I. and Moseev, D.S., 2023. Study of the Marginal Filter of the Mesotidal Estuary of the Indiga River (Barents Sea Basin). Oceanology, 63(3), pp. 432–433. https://doi.org/10.1134/s0001437023030116
  7. Kotova, E.I., Miskevich, I.V., Moseev, D.S. and Chultsova, A.L., 2022. Characteristics of the Marginal Filter in the Indiga Estuary (Barents Sea). _Advances in Current Natural Science_s, (9), pp. 39–44. https://doi.org/10.17513/use.37890
  8. Lukashin, V.N., Kravchishina, M.D., Klyuvitkin, A.A., Novigatsky, A.N. and Politova, N.V., 2019. Geochemistry of Suspended Particulate Matter in the Volga River Marginal Filter. Oceanology, 59(3), pp. 381–391. https://doi.org/10.1134/S0001437019030135
  9. Gordeev, V.V. and Lisitsin, A.P., 2017. [Geochemistry of the Surface Layer of White Sea Bottom Sediments]. In: I. A. Nemirovskaya, V. P. Shevchenko, V. G. Vorontsova, eds., 2017. The White Sea System. Volume IV. The Processes of Sedimentation, Geology and History. Moscow: Scientific World, pp. 571–639 (in Russian).
  10. Nemirovskaya, I.A., ed., 2016. The Caspian Sea System. Moscow: Nauchny Mir, 480 p. (in Russian).
  11. Gordeev, V.V., 2018. Features of River Flow Geochemistry in the Black Sea. In: A. P. Lisitzin, ed., 2018. The Black Sea System. Moscow: Nauchny Mir, pp. 247–286. https://doi.org/10.29006/978-5-91522-473-4.2018.247-286 (in Russian).
  12. Zaitsev, Yu.P., Alexandrov, B.G. and Minicheva, G.G., eds., 2006. North-Western Part of the Black Sea: Biology and Ecology. Kyiv: Naukova Dumka, 702 p. (in Russian).
  13. Egorov, V., Milchakova, N., Bobko, N. and Marchenko, Yu., 2025. Macrophytocenoses as Biogeochemical Barriers to Water Hypereutrophication by Mineral Phosphorus Off the Southwestern Crimea. Marine Biological Journal, 10(2), pp. 3–17.
  14. Berlinsky, N.A., Den’ga, Yu.M., Matveev, A.V., Podust, O.S., Popov, Yu.I. and Tretyak, I.P., 2014. The Influence of Marine Environment Variability to the Dynamic of the Black Sea Phillophora Area (Zernov Field). Odesa National University Herald. Geography and Geology, 19(2), pp. 40–57 (in Russian).
  15. Fashchuk, D., 2018. The Black Sea: Ups-And-Downs in Domestic Fishery. Priroda, (10), pp. 65–79. https://doi.org/10.31857/S0032874X0001453-7 (in Russian).
  16. Zernov, S.A., 1908. Report on a Field Trip to the North-Western part of the Black Sea to Study the Fauna and Collect Specimens for the Zoological Museum of the Imperial Academy of Sciences (Odessa Bay, the Dnieper-Bug Estuary, and the Karkinitsky and Dzharylgatsky Bays). Ezhegodnik Zoologicheskogo Muzeya Imperatorskoy Akademii Nauk, 13. Saint Petersburg: Imperatorskaya Akademiya Nauk, pp. 154–166 (in Russian).
  17. Zernov, S.A., 1913. [Facies of Phyllophora (Algae-Rhodophceae): Phyllophora Field in the North-Western Black Sea]. Ezhegodnik Zoologicheskogo Muzeya Imperatorskoy Akademii Nauk, 14. Saint Petersburg: Imperatorskaya Akademiya Nauk, pp. 181–191 (in Russian).
  18. Zernov, S.A., 1913. [On Studying Life of the Black Sea]. In: IAS, 1913. Zapiski Imperatorskoy Akademii Nauk. Saint Petersburg: Imperatorskaya Akademiya Nauk. Vol. 32, iss. 1, 304 p. (in Russian).
  19. Shchapova, T.F., 1954. [Phyllophora of the Black Sea]. Trudy Instituta Okeanologii, 11, pp. 3–35 (in Russian).
  20. Tolmazin, D., 1985. Changing Coastal Oceanography of the Black Sea. I: Northwestern Shelf. Progress in Oceanography, 15(4), pp. 217–276. https://doi.org/10.1016/0079-6611(85)90038-2
  21. Ivanov, L.I., Beşiktepe, Ş. and Özsoy, E., 1997. The Black Sea Cold Intermediate Layer. In: E. Özsoy and A. Mikaelyan, eds., 1997. Sensivity to Change: Black Sea, Baltic Sea and North Sea. NATO ASI Series; vol. 27. Dordrecht: Kluwer Academic Publishers, pp. 253–264. https://doi.org/10.1007/978-94-011-5758-2_20
  22. Oguz, T. and Besiktepe, S., 1999. Observations on the Rim Current Structure, CIW Formation and Transport in the Western Black Sea. Deep Sea Research Part I: Oceanographic Research Papers, 46(10), pp. 1733–1753. https://doi.org/10.1016/S0967-0637(99)00028-X
  23. Stanev, E.V. and Staneva, J.V., 2001. The Sensitivity of the Heat Exchange at Sea Surface to Meso and Sub-Basin Scale Eddies: Model Study for the Black Sea. Dynamics of Atmospheres and Oceans, 33(3), pp. 163–189. https://doi.org/10.1016/S0377-0265(00)00063-4
  24. Beckers, J.M., Gregoire, M., Nihoul, J.C.J., Stanev, E., Staneva, J. and Lancelot, C., 2002. Modelling the Danube-influenced Northwestern Continental Shelf of the Black Sea. I: Hydrodynamical Processes Simulated by 3-D and Box Models. Estuarine, Coastal and Shelf Science, 54(3), pp. 453–472. https://doi.org/10.1006/ecss.2000.0658
  25. Ovchinnikov, I.M. and Popov, Yu.I., 1984. [On the Formation of the Cold Intermediate Layer in the Black Sea], Doklady AN SSSR, 279(4), pp. 986–989 (in Russian).
  26. Ovchinnikov, I.M. and Popov, Yu.I., 1987. Cold Intermediate Layer Formation in the Black Sea. Okeanologiya, 27(5), pp. 739–746 (in Russian).
  27. Ovchinnikov, I.M. and Popov, Yu.I., 1990. [Features of Formation of the Cold Intermediate Layer in the Black Sea under Severe Winter Conditions]. In: R. R. Belevich, ed., 1990. Proceedings of SOI, 190, pp. 132–151 (in Russian).
  28. Gertman, I.F., 1991. [Thermohaline Structure of the Black Sea]. In: A. I. Simonov and E. N. Altman, eds., 1991. Hydrometeorology and Hydrochemistry of Seas in the USSR. Vol. 4. Black Sea. Issue 1. Hydrometeorological Conditions. Saint Petersburg: Gidrometeoizdat, pp. 146–195 (in Russian).
  29. Titov, V.B., 2003. Interannual Renewal of the Cold Intermediate Layer in the Black Sea over the Last 130 Years. Russian Meteorology and Hydrology, (10), pp. 51–56.
  30. Eremeev, V.N., Ivanov, V.A. and Tuzhilkin, V.S., 1991. [Climate Features of Intra-annual Variations of the Black Sea Hydrophysical Fileds]. Sevastopol. 36 p. Preprint, MHI of the Academy of Sciences of the Ukrainian SSR (in Russian).
  31. Polonsky, A.B. and Popov, Yu.I., 2011. Conditions of Formation of Cold Intermediate Layer of the Black Sea. Modern Problems of Oceanology. Sevastopol: ECOSI-Gidrofizika. Iss. 8, 54 p. (in Russian).
  32. Ivanov, V.A. and Belokopytov, V.N., 2013. Oceanography of the Black Sea. Sevastopol: ECOSI-Gidrofizika, 210 p.
  33. Melnikov, V., Minkina, N., Melnik, L. and Pollehne, F., 2021. Distribution of Sprattus sprattus phalericus (Risso, 1827) and Zooplankton near the Black Sea Redoxcline. Journal of Fish Biology, 99(4), pp. 1393–1402. https://doi.org/10.1111/jfb.14848
  34. Melnikov V.V., 2023. Anaeroconture of the Black Sea. Izvestia of Samara Scientific Center of the Russian Academy of Sciences, 25(5), pp. 203–218. https://doi.org/10.37313/1990-5378-2023-25-5-203-218 (in Russian).
  35. Vasechkina, E.F. and Melnikov, V.V., 2024. Modeling the Structure Changes of Cold-Water Copepods Calanus euxinus Population under the Influence of the Black Sea Depths Deoxygenation. Journal of Marine Systems, 246. 104017. https://doi.org/10.1016/j.jmarsys.2024.104017
  36. Evstigneeva, I.K. and Nekhoroshev, M.V., 2008. Morpho-Functional Features of Phyllophora Grev. Genus Species in Conditions of Zernov’s Field (the Black Sea). Trudy YUGNIRO = YugNIRO Proceedings. Kerch: YugNIRO Publishers. Vol. 46, pp. 55–63 (in Russian).
  37. Kalugina-Gutnik, A.A. and Evstigneeva, I.K., 1993. Long-Term Dynamics of Species Composition and Structure of Bottom Phytocenoses of the Phyllophorous Zernov’s Field. Ecology of the Sea, 43, pp. 90–97 (in Russian).
  38. Tkachenko, F.P., Tretiak, I.P. and Kostilev, E.F., 2008. Macrophytobenthos of Zernov Phyllophora Field in Present Conditions (Black Sea, Ukraine). Algologia, 18(4), pp. 423–431 (in Russian).
  39. Minicheva, G.G., 2007. Contemporary Morphofunctional Transformation of Seaweed Communities of the Zernov Phyllophora Field. Algology, 17(2), pp. 171–190 (in Russian).
  40. Stepanyan, O.V., 2020. Macrophytobenthos of the Black and Azov Seas: Floristic and Ecological Aspects (Overview). Science in the South of Russia, 16(4), pp. 26–38. https://doi.org/10.7868/S25000640200404 (in Russian).
  41. Stepanyan, O.V., 2019. Zernov’s Phyllophora Field: 110 Years of the Unique Discovery in the Black Sea. Priroda, (6), pp. 62–70. https://doi.org/10.7868/S0032874X19060085 (in Russian).
  42. Borysova, O., Kondakov, A., Paleari, S., Rautalahti-Miettinen, E., Stolberg, F. and Daler, D., 2005. Eutrophication in the Black Sea Region: Impact Assessment and Causal Chain Analysis. Kalmar: University of Kalmar. 62 p.
  43. Vörösmarty, C.J., Sharma, K.P., Fekete, B.M., Copeland, A.H., Holden, J., Marble, J. and Lough, J.A., 1997. The Storage and Aging of Continental Runoff in Large Reservoir Systems of the World. Ambio, 26(4), pp. 210–219. Available at: http://www.jstor.org/stable/4314590 [Accessed: 18.August 2026].
  44. Mikaelyan, A.S., Zatsepin, A.G. and Chasovnikov, V.K., 2013. Long-Term Changes in Nutrient Supply of Phytoplankton Growth in the Black Sea. Journal of Marine Systems, 117–118, pp. 53–64. https://doi.org/10.1016/j.jmarsys.2013.02.012
  45. Konovalov, S.K., Murray, J.W. and Luther, G.W., 2005. Basic Processes of Black Sea Biogeochemistry. Oceanography, 18(2), pp. 28–35. https://doi.org/10.5670/oceanog.2005.39
  46. Capet, A., Grégoire, M., Stanev, E.V., Beckers, J.M. and Murray, J.W., 2016. Decline of the Black Sea Oxygen Inventory. Biogeosciences, 13(4), pp. 1287–1297. https://doi.org/10.5194/bg-13-1287-2016/
  47. Oven, L.S., Shevchenko, N.F. and Volodin, S.V., 1996. Sprats Nutrition in Different Regions of the Black Sea. In: S.M. Konovalov, ed., 1996. The Modern State of Black Sea Ichtyophauna. Sevastopol: IBSS NASU, pp. 34–38 (in Russian).
  48. Shushkina, E.A. and Musaeva, E.I., 1990. Structure of Epipelagic ZooplanktonCommunity and its Changes Related to the Invasion of Mnemiopsis leidyi in the Black Sea. Okeanologiya, 30(2), pp. 306–310 (in Russian).
  49. Vinogradov, M.E., Sapozhnikov, V.V. and Shushkina, E.A., 1992. [Ecosystem of the Black Sea]. Moscow: Nauka, 111 p. (in Russian).
  50. Ignatyev, S.M., Zuev, G.V. and Melnikova, E.B., 2001. The Long-Term Dynamic of the State of Ctenophora, Mnemiopsis leidyi Agassis Population in the Sevastopol Region (the Black Sea). Ecology of the Sea, 56, pp. 8–12 (in Russian).
  51. Shiganova, T., Musaeva, E., Arashkevich, E., Kamburska, L., Stefanova, K., Mihneva, V., Polishchuk, L., Timofte, F., Ustun, F. et al., 2008. The State of Zooplankton. In: T. Oguz, ed., 2008. State of the Environment of the Black Sea (2001–2006/7). Istanbul, 2008. Chapter 6, pp. 201–246. Available at: https://www.blackseacommission.org/Downloads/ SOE27032009-2_compressed.pdf [Accessed: 08 August 2026].
  52. Kideys, A.E., 1994. Recent Dramatic Changes in the Black Sea Ecosystem: the Reason for the Sharp Decline in Turkish Anchovy Fisheries. Journal of Marine Systems, 5(2), pp. 171–181. https://doi.org/10.1016/0924-7963(94)90030-2
  53. Shulman, G.E., Nikolsky, V.N., Yuneva, T.V., Shchepkina, A.M., Bat, L. and Kideys, A.E., 2007. Influence of Global Climatic and Regional Anthropogenic Factors on Small Pelagic Fishes of the Black Sea. Marine Ecological Journal, 6(4), pp. 18–30 (in Russian).
  54. Konsulov, A.S. and Kamburska, L.T., 1998. Ecological Determination of the New Ctenophora – Beroe ovata Invasion in the Black sea. In: BAS, 1998. Proceedings of the Institute of Oceanology. Varna: Bulgarian Academy of Sciences, 2, pp. 195–198.
  55. Vinogradov, M.E. and Shushkina, E.A., 2002. Age Structure Dynamics in Populations of the ctenophores Mnemiopsis leidyi and Beroe ovata in Near Shore Caucasian Region of the Black Sea in 2000. In: A. G. Zatsepin and M. V. Flint, eds., 2002. Multidisciplinary Investigations of the Northeast Part of the Black Sea. Moscow: Nauka, pp. 272–288 (in Russian).
  56. Çokacar, T., 2024. Cold Intermediate Water Formation in the Black Sea Triggered by March 2022 Cold Intrusions. Journal of Marine Science and Engineering, 12(11), 2027. https://doi.org/10.3390/jmse12112027
  57. Melnikov, V.V., Serebrennikov, A.N., Masevich, A.V. and Chudinovskikh, E.S., 2025. The Main Patterns of the Black Sea Ecosystem Long-Term Changes. Ecological Safety of Coastal and Shelf Zones of Sea, (2), pp. 19–35.
  58. Vidnichuk, A.V. and Konovalov, S.K., 2021. Changes in the Oxygen Regime in the Deep Part of the Black Sea in 1980–2019. Physical Oceanography, 28(2), pp. 180–190. https://doi.org/10.22449/1573-160X-2021-2-180-190
  59. Zaitsev, Y. and Mamaev, V., 1997. Marine Biological Diversity in the Black Sea. A Study of Change and Decline. New York: United Nations Publications. Black Sea Environmental Series No. 3. 1997. 208 p.
  60. Öztürk, B., 2021. Non-Indigenous Species in the Mediterranean and the Black Sea. Rome: FAO, 106 p. Series Studies and reviews (General Fisheries Commission for the Mediterranean). No. 87. https://doi.org/10.4060/cb5949en
  61. Terentjev, A.S., 2007. Changes of Mussel Stock in the Black Sea Shelf Area Adjacent to the Kerch Strait. Trudy VNIRO, 147, pp. 335–349 (in Russian).
  62. Stadnichenko, S.V. and Zolotarev, V.N., 2009. Population Structure of Marine Bivalves in the Danube Delta Area in 2007–2008. Ecological Safety of Coastal and Shelf Zones and Comprehensive Use of Shelf Resources, 20, pp. 248–261 (in Russian).
  63. Stanev, E.V., Peneva, E. and Chtirkova, B., 2019. Climate Change and Regional Ocean Water Mass Disappearance: Case of the Black Sea. Journal of Geophysical Research: Oceans, 124(7), pp. 4803–4819. https://doi.org/10.1029/2019JC015076
  64. Melnikov, V.V., 2024. Reasons for the Appearance and Preservation of Boreal Pelagic Biota in the Black Sea. Izvestia of Samara Scientific Center of the Russian Academy of Sciences, 26(5), pp. 109–121. https://doi.org/10.37313/1990-5378-2024-26-5-109-121 (in Russian).
  65. Klimova, T.N., Vdodovich, I.V., Anninsky, B.E., Subbotin, A.A., Podrezova, P.S. and Melnikov, V.V., 2021. Effect of Certain Abiotic and Biotic Factors on Spawning of the European Sprat Sprattus sprattus (Linnaeus, 1758) in the Black Sea in November 2016–2017. Oceanology, 61(1), pp. 58–68. https://doi.org/10.1134/S0001437021010082
  66. Sazhina, L.I., 1973. [Growth of Black Sea Mass Copepods Under Laboratory Conditions]. In: V. A. Vodyanitsky, ed., 1973. Biologiya Morya. Kiev: Naukova Dumka. Iss. 28, pp. 41–51 (in Russian).
  67. Cannaby, H., Fach, B.A., Arkin, S.S. and Salihoglu, B., 2015. Climatic Controls on Biophysical Interactions in the Black Sea under Present Day Conditions and a Potential Future (A1B) Climate Scenario. Journal of Marine Systems, 141, pp. 149–166. https://doi.org/10.1016/j.jmarsys.2014.08.005
  68. Belokopytov, V.N. and Zhuk, E.V., 2024. Climatic Variability of the Black Sea Thermohaline Characteristics (1950–2023). Physical Oceanography, 31(6), pp. 788–801.
  69. Kapkov, V.I., Saburin, M.Yu., Belenikina, O.A. and Blinova, E.I., 2005. Phytocenosis Recovery of Cystoseira Crinita (Phaeophyta) and Seaweeds Growth Dynamics at the Artificial Reefs. Herald of Moscow University. Series 16. Biology, (2), pp. 30–34 (in Russian).
  70. Maximova, O.V. and Moruchkova, N.A., 2005. Long-Term Anthropogenic Transformation and Contemporary State of the North Caucasian Macrophytobenthos (Black Sea). Oceanology, 45(Suppl. 1), pp. 168–175.
  71. Milchakova, N.A., Mironova, N.V., Aleksandrov, V.V. and Ryabogina, V.G., 2011. [A Current State of Zernov’s Phyllophora Field (the Black Sea)]. In: IFNUL, 2011. [Proceedings of the 13th Meeting of the Ukrainian Botanical Society (Lviv, 19–23 September 2011)]. Lviv: Prostir M, p. 306 (in Russian).
  72. Milchakova, N.A., Mironova, N.V. and Ryabogina, V.G., 2011. [Marine Plant Resour-ces]. In: V. N. Eremeev, A. V. Gaevskaya, G. E. Shulman and Ju. A. Zagorodnyaya, eds., 2011. Biological Resources of the Black Sea and Sea of Azov. Sevastopol: ECOSI-Gidrofizika. Ch. 4, pp. 117–139 (in Russian).
  73. Milchakova, N.A., Mironova, N.V. and Aleksandrov, V.V., 2013. Dynamics, Causes of Degradation and the Prognosis of Restoration Zernov Phyllophora Field (The Black Sea)). Issues of Modern Algology, (2), 2 (in Russian).

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