S. A. Piontkovski1,*, A. V. Melnik2, Yu. A. Zagorodnyaya2, Yu. G. Artemov2, E. A. Skripaleva3, E. Yu. Georgieva2
1 Sevastopol State University, Sevastopol, Russia
2 A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS, Sevastopol, Russia
3 Marine Hydrophysical Institute of RAS, Sevastopol, Russia
* e-mail: spiontkovski@mail.ru
Abstract
Spatial heterogeneity of the thermohaline structure, biotope dynamics and organism interactions form layers of high abundance and biomass. Based on expedition data obtained during the summers of 2010–2024, this paper analyses characteristics of surface and subsurface peaks in phytoplankton biomass, chlorophyll a, total suspended matter, zooplankton (prey and gelatinous), bioluminescence intensity and sound-scattering layers (indicators of the abundance of small pelagic organisms) on the shelf off the coast of Crimea. Characteristic parameter values, layer thicknesses and depths are presented. The mechanisms of layering formation and the relationship between the structural and functional properties of the pelagic community are discussed. It was noted that at lower trophic levels, layering was regulated primarily by thermohaline stratification of the water column. At intermediate trophic levels, represented by copepods and small pelagic fishes, the dominant factor regulating stratification was the organisms' motor activity, associated with feeding behavior, reproductive behavior, defense and other behaviors. In terms of the relationship between the structure and function in a pelagic ecosystem, it was noted that the stratified distribution of organisms created vertical heterogeneity in the density of trophic interactions and consequently vertical heterogeneity in the flow of matter and energy within the community. Trophic interactions were most intense in the layers of maximum thickness due to their greater ecological capacity. These include the surface and subsurface biomass peaks of phytoplankton, zooplankton and small pelagic fishes, primarily common anchovy and sprat.
Keywords
Black Sea, coastal shelf, thermohaline waters structure, phytoplankton, chlorophyll a, total suspended matter concentration, zooplankton, bioluminescence, sound scattering layers, pelagic community
Acknowledgments
The work was funded by the IBSS state assignment no. 124030400057-4, no. 124022400148-4-0556-2024-00, no.124030100127-7, SevSU state assignment no. FEFM-2023-0005, and MHI state assignment no. FNNN-2024-0014. The expeditions were carried out onboard R/V Professor Vodyanitsky. Special thanks should be addressed to V. V. Davydov, the research vessel Chief Engineer, with respect to his long-term support of field measurements on board. The figure of station grids was produced by I. A. Minsky.
About the authors
Sergey A. Piontkovski, Leading Researcher, Sevastopol State University (33 Universitetskaya St., 299053, Sevastopol, Russia), DSc (Biol.), ORCID ID: 0000-002-6472-9701, Scopus Author ID: 6602165194, ResearcherID: ABB-9334-2020, spiontkovski@mail.ru
Aleksandr V. Melnik, Senior Researcher, A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS (2 Nakhimova Ave, Sevastopol, 299011, Russia), PhD (Biol.), ORCID ID: 0000-0002-4371-384X, Scopus AuthorID: 57219127014, ResearcherID: X-1393-2019, melnikav@ibss-ras.ru Yulia A. Zagorodnyaya, Leading Researcher, A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS (2 Nakhimova Ave, Sevastopol, 299011, Russia), PhD (Biol.), ORCID ID:0000-0002-9502-4923, Scopus Author ID: 6506214138, ResearcherID: E-3325-2018, artam-ant.yandex.ru
Yuri G. Artemov, Leading Researcher, A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS (2 Nakhimova Ave, Sevastopol, 299011, Russia), PhD (Geogr.), ORCID ID: 0000-0002-4725-1427, Scopus Author ID: 12767058200, ResearcherID: G-1797-2015, yu.g.artemov@gmail.com
Elena A. Skripaleva, Senior Researcher, Marine Hydrophysical Institute of RAS (2 Kapitanskaya St., Sevastopol, 299011, Russia), PhD (Geogr.), ResearcherID: AAC-6648-2020, ORCID ID: 0000-0003-1012-515X, sea-ant@yandex.ru
Elena Yu. Georgieva, Leading Engineer, A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS (2 Nakhimova Ave, Sevastopol, 299011, Russia), ORCID ID: 0000-0002-8177-0781, Scopus Author ID: 57193546928, e-georgieva@mail.ru
For citation
Piontkovski, S.A., Melnik, A.V., Zagorodnyaya, Yu.A., Artemov, Yu.G., Skripaleva, E.A., Georgieva, E.Yu., 2026. Layering of the Spatial Structure of the Crimean Shelf Pelagic Community in the Summer Season. Ecological Safety of Coastal and Shelf Zones of Sea, (1), pp. 27–51.
References
- Behrenfeld, M.J., O’Malley, R.T., Boss, E.S., Westberry, T.K., Graff, J.R., Halsey, K.H., Milligan, A.J., Siegel, D.A. and Brown, M.B., 2016. Revaluating Ocean Warming Impacts on Global Phytoplankton. Nature Climate Change, 6, pp. 323–330. https://doi.org/10.1038/NCLIMATE2838
- Longhurst, A., 1998. Ecological Geography of the Sea. San Diego: Academic Press, 398 p.
- Moriarty, R. and O’Brien, T.D., 2013. Distribution of Mesozooplankton Biomass in the Global Ocean. Earth System Science Data, 5(1), pp. 45–55. https://doi.org/10.5194/essd-5-45-2013
- Piontkovski, S.A., 2005. Multiscale Variability of Mesoplanktonic Fields of the Ocean. Sevastopol: ECOSI-Gidrophizika, 194 р. (in Russian).
- Mikhailovsky, G.E., 1984. [Specificity of Ecological Systems and Problems of their Study]. Zhurnal Obshchei Biologii, 45(1), pp. 66–77 (in Russian).
- Lasker, R., 1975. Field Criteria for Survival of Anchovy Larvae: the Relation Between the Inshore Chlorophyll Layers and Successful First Feeding. Fishery Bulletin, 73(3), pp. 453–462.
- Glushchenko, T.I. and Chashchin, A.K., 2008. Peculiarities of Nutrition of the Black Sea Sprat Sprattus Sprattus Phalericus (Risso) (Pisces: Clupeidae) and Formation of its Feeding Accumulations. Marine Ekological Journal, 7(3), pp. 5–14 (in Russian).
- Klimova, T.N., Subbotin, A.A., Vdodovich, I.V., Zagorodnyaya, Yu.A. and Zabrodin, D.A., 2024. Ichthyoplankton in the Northern Part of the Black Sea under the Prolongation of Summer Hydrological Season in 2020. Inland Water Biology, 17(1), pp. 197–207. https://doi.org/10.1134/S1995082924010085
- Zaika, V.E., 1981. Environment Capacity: the Content of the Notion and Its Application in Ecology. Ecology of the Sea, 7, pp. 3–9 (in Russian).
- Samodurov, A.S. and Chukharev, A.M., 2017. Intensity of Vertical Turbulent Exchange in the Black Sea Summer Pycnocline Around the Crimean Peninsula. Journal of Physics: Conference Series, 899(2), 0220015. https://doi.org/10.1088/1742-6596/899/2/022015
- Piontkovski, S.A., Al-Oufi, H.S. and Al-Abri, N.M., 2016. Fish Landings and Oman Shelf Area. Journal of Agricultural and Marine Sciences, 21(1), pp. 25–32. https://doi.org/10.24200/jams.vol21iss0pp25-32
- Artamonov, Yu.V., Skripaleva, E.A., Fedirko, A.V., Shutov, S.А., Derjushkin, D.V., Shapovalov, R.O., Shapovalov, Yu. I. and Shcherbachenko, S.V., 2020. Waters Circulation in the Northern Part of the Black Sea in Summer – Winter of 2018. Ecological Safety of Coastal and Shelf Zones of Sea, (1), pp. 69–90. https://doi.org/10.22449/2413-5577-2020-1-69-90 (in Russian).
- Ivanov, V.A., Kuznetsov, A.S. and Morozov, A.N., 2019. Monitoring Coastal Water Circulation along the South Coast of Crimea. Doklady Earth Sciences, 485(2), pp. 405–408. https://doi.org/10.1134/S1028334X19040044
- Zagorodnyaya, Yu.A. and Piontkovski, S.A., 2022. Seasonal and Interannual Variations of the Abundance of the Dinoflagellate Noctiluca Scintillans in the Northern Black Sea. Marine Biology Research, 18(1-2), pp. 104–116. https://doi.org/10.1080/17451000.2022.2086701
- Kovalev, A.V., Melnikov, V.V., Ostrovskaya, N.A. and Prusova, I.Yu., 1993. [Macroplankton]. In: A. V. Kovalev and Z. Z. Finenko, eds., 1993. [Plankton of the Black Sea]. Kiev: Naukova Dumka, pp. 183–193 (in Russian).
- Petipa, T.C., 1957. [On Average Weight of Main Zooplankton Forms of the Black Sea]. In: Academy of Sciences of the USSR, 1957. Trudy Sevastopolskoy Biologicheskoy Stantsii, 9, pp. 39–57 (in Russian).
- Latushkin, A.A., Artamonov, Yu.V., Skripaleva, E.A. and Fedirko, A.V., 2022. The Relationship of the Spatial Structure of the Total Suspended Matter Concentration and Hydrological Parameters in the Northern Black Sea According to Contact Measurements. Fundamental and Applied Hydrophysics, 15(2), pp. 124–137. https://doi.org/10.48612/fpg/4heu-kxbn-gg7t (in Russian).
- Melnik, A., Melnik, L., Mashukova, O. and Melnikov, V., 2021. Field Studies of Bioluminescence in the Antarctic Sector of the Atlantic Ocean in 2002 and 2020. Luminescence, 36(8), pp. 1910–1921. https://doi.org/10.1002/bio.4125
- Kuznetsov, A.S. and Ivashchenko, I.K., 2023. Features of Forming the Alongcoastal Circulation of the Coastal Ecotone Waters nearby the Southern Coast of Crimea. Physical Oceanography, 30(2), pp. 171–185.
- Simonova, Yu.V., Stanichny, S.V. and Lemeshko, E.M., 2020. [Characteristics of Upwellings in the Southern coast of Crimea Based on a Comprehensive Analysis of In Situ and Remote Observations]. In: SRI RAS, 2020. Proceedings of the 19th All-Russian Open Conference “Modern Problems of Remote Probing the Earth from Space”. Moscow: Space Research Institute of RAS Publishing, p. 335 (in Russian).
- Kubryakova, E. and Kubryakov, A., 2020. Warmer Winter Causes Deepening and Intensification of Summer Subsurface Bloom in the Black Sea: the Role of Convection and Self-Shading Mechanism, Biogeosciences Discussions. [Preprint]. https://doi.org/10.5194/bg-2020-210
- Drits, A.V., Nikishina, A.B., Sergeeva, V.M. and Solov’ev, K.A., 2013. Feeding, Respiration, and Excretion of the Black Sea Noctiluca scintillans MacCartney in Summer. Oceanology, 53(4), pp. 442–450. https://doi.org/10.1134/S0001437013040036
- Zagorodnyaya, Yu.A. and Moryakova, V.K., 2018. [Holoplankton]. In: N. S. Kostenko, ed., 2018. The Biology of the Black Sea Offshore Area at the South-Eastern Crimea. Simferopol: PP “ARIAL”, pp. 46–59 (in Russian).
- Mikaelyan, A.S., Malej, A., Shiganova, T.A., Turk, V., Sivkovitch, A.E., Musaeva, E.I., Kogovšek, T. and Lukasheva, T.A., 2014. Populations of the Red Tide Forming Dinoflagellate Noctiluca scintillans (Macartney): A Comparison Between the Black Sea and the Northern Adriatic Sea. Harmful Algae, 33, pp. 29–40. https://doi.org/10.1016/j.hal.2014.01.004
- Zagorodnyaya, Yu.A., Piontkovski, S.A. and Gubanov, V.V., 2023. Pelagic Ecosystem of the Black Sea Goes Gelatinous. Marine Biology Research, 19(6-7), pp. 317–326. https://doi.org/10.1080/17451000.2023.2235571
- Zaitsev, Yu.P. and Polishchuk, L.N., 1984. An Increase in the Number of Aurelia aurita (L.) in the Black Sea. Biologiya Morya, 17, pp. 35–46 (in Russian).
- Balykin, P.A., Kutsyn, D.N. and Startsev A.V., 2021. Fishing Under Climate Change: Dynamics of Composition and Structure of Catches in the Russian Black Sea in the XXI Century. Marine Biological Journal, 6(3), pp. 3–14. https://doi.org/10.21072/mbj.2021.06.3.01 (in Russian).
- Brough, T., Rayment, W. and Dawson, S., 2019. Using a Recreational Grade Echosounder to Quantify the Potential Prey Field of Coastal Predators. PLoS ONE, 14(5), e0217013. https://doi.org/10.1371/journal.pone.0217013
- McInnes, A.M., Khoosal, A., Murrell, B., Merkle, D., Lacerda, M., Nyengera, R., Coetzee, J.C., Edwards, L.C., Ryan, P.G. et al. Recreational Fish-Finders – an Inexpensive Alternative to Scientific Echo-Sounders for Unravelling the Links Between Marine Top Predators and Their Prey. PLoS ONE, 10(11), e0140936. https://doi.org/10.1371/journal.pone.0140936
- Artemov, Yu.G., 2006. Software Support for Investigation of Natural Methane Seeps by Hydroacoustic Method. Marine Ekological Journal, 5(1), pp. 57–71.
- Serikova, I.M., 2020. Algorithm for Mathematical Processing of Bioluminescence Profiles for the Study of Small-Scale Agregation of Plankton. Monitoring Systems of Environment, (1), pp. 145–152. https://doi.org/10.33075/2220-5861-2020-1-145-152 (in Russian).
- Melnik, A.V., Georgieva, E.Iu. and Melnik, L.A., 2019. The Variability of the Spatial Distribution of Bioluminescence and Phytoplankton in the Photic Layer of the Black Sea in Summer 2018. Monitoring systems of environment, (3), pp. 120–126. https://doi.org/10.33075/2220-5861-2019-3-120-126 (in Russian).
- Tokarev, Yu.N., Bityukov, E.P., Vasilenko, V.I. and Sokolov, B.G., 2000. The Bioluminescence Field as a Characteristic Index of the Black Sea Plankton Community Structure. Ecology of the Sea, 53, pp. 20–25 (in Russian).
- Serikova, I.M., Evstigneev, V.P., Tokarev, Yu.N. and Suslin, V.V., 2017. Bioluminescence Field of the Black Sea as Indicator of Dinophyta Aggregation, Its Seasonal and Interannual Dynamics. In: SPIE, 2017. Proceedings of SPIE. Vol. 10466: 23rd International Symposium on Atmospheric and Ocean Optics: Atmospheric Physics, 104663X. https://doi.org/10.1117/12.2287964
- Evstigneev, V.P., Serikova, I.M. and Kyrylenko, N.F., 2019. Biotic and Abiotic Influence on Bioluminescence Field in Summer. In: MEDCOAST, 2019. 14th MEDCOAST Congress on Coastal and Marine Sciences, Engineering, Management and Conservation, MEDCOAST 2019. Marmaris, 2019. Vol. 1, pp. 307–318.
- Bityukov, E.P., Evstigneev, P.V. and Tokarev, Yu.N., 1993. Luminescent Dinoflagellata of the Black Sea as Affected by Anthropogenic Factors. Gidrobiologichesky Zhurnal, 29(4), pp. 27–34 (in Russian).
- Serikova, I.M., Bryantseva, Yu.V. and Vasilenko, V.I., 2013. Seasonal Dynamics of the Bioluminescence Field's Structure and Its Connection with the Dinoflagellates Parameters. Marine Ekological Journal, 12(4), pp. 87–95 (in Russian).
- Yunev, O.A., Konovalov, S.K. and Velikova, V., 2019. Anthropogenic Eutrophication in the Black Sea Pelagic Zone: Long-Term Trends, Mechanisms, Consequences. Moscow: GEOS, 194 p. (in Russian).
- Finenko, Z.Z., Churilova, T.Ya. and Lee, R.I., 2005. Vertical Distribution of Chlorophyll and Fluorescence in the Black Sea. Marine Ekological Journal, 4(1), pp. 15–46 (in Russian).
- Piontkovski, S.A. and Seregin, S.A., 2006. The Behavior of Copepods. Sevastopol: Ekosi-Gidrophizika, 148 р. (in Russian).
- Ricour, F., Capet, A., D'Ortenzio, F., Delille, B. and Grégoire, M., 2021. Dynamics of the Deep Chlorophyll Maximum in the Black Sea as Depicted by BGC-Argo Floats. Biogeosciences, 18(2), pp. 755–774. https://doi.org/10.5194/bg-18-755-2021
- Tsou, T., 2011. Determining the Mean-Variance Relationship in Generalized Linear Models – a Parametric Robust Way. Journal of Statistical Planning and Inference, 141(1), pp. 197–203. https://doi.org/10.1016/j.jspi.2010.05.029
- 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
- Dorofeev, V.L. and Sukhikh, L.I., 2023. Analysis of Long-Term Variability of Hydrodynamic Fields in the Upper 200-Meter Layer of the Black Sea Based on the Reanalysis Results. Physical Oceanography, 30(5), pp. 581–593.
- Stelmakh, L.V., 2013. Microzooplankton Grazing Impact on Phytoplankton Blooms in the Coastal Seawater of the Southern Crimea (Black Sea). International Journal of Marine Science, 3(15), pp. 121–127. https://doi.org/10.5376/ijms.2013.03.0015
- Petipa, T.S., Pavlova, E.V. and Mironov, G.N., 1970. [Food Web Structure, Transfer and Use of Matter and Energy in Black Sea Plankton Communities]. Biologiya Morya, 19, pp. 3–43 (in Russian).