ecodagЮг России: экология, развитиеSouth of Russia: ecology, development1992-10982413-0958State Institute of Applied Ecology of the Republic of Dagestan10.18470/1992-1098-2020-2-35-47ecodag-1967Research ArticleЭКОЛОГИЯ ЖИВОТНЫХECOLOGY OF ANIMALSАльтернативные условия массового появления сцифоидной медузы Aurelia aurita (Linnaeus, 1758) и гребневика Pleurobrachia pileus (O.F. Muller, 1776) в планктоне Черного моряAlternative conditions of mass appearance of the scyphozoan jellyfish, Aurelia aurita (Linnaeus, 1758), and the ctenophore, Pleurobrachia pileus (O.F. Muller, 1776), in plankton of the Black Seahttps://orcid.org/0000-0001-9614-9392АннинскийБ. Е.AnninskyB. E.

Борис Е. Аннинский, кандидат биологических наук, ведущий научный сотрудник, отдел физиологии животных и биохимии

299011, г. Севастополь, пр. Нахимова, 2.Тел. +79788370828

Boris E. Anninsky, Cand. Sci. (Biol.), Leading Researcher, Department of Animal Physiology and Biochemistry

2 Nakhimov Ave, Sevastopol, 299011.Tel. +79788370828

anninsky_b@mail.ruhttps://orcid.org/0000-0002-3303-0906ФиненкоГ. А.FinenkoG. A.

Галина А. Финенко

Севастополь

Galina A. Finenko

Sevastopol

https://orcid.org/0000-0002-8113-8786ДацыкН. А.DatsykN. A.

Наталия А. Дацык

Севастополь

Natalya A. Datsyk

Sevastopol

Институт биологии южных морей им. А.О. Ковалевского РАНA.O. Kovalevsky Institute of Biology of the Southern Seas, Russian Academy of Sciences20200808202015235‐47Copyright © Аннинский Б.Е., Финенко Г.А., Дацык Н.А., 20202020Аннинский Б.Е., Финенко Г.А., Дацык Н.А.Anninsky B.E., Finenko G.A., Datsyk N.A.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://ecodag.elpub.ru/ugro/article/view/1967

Цель. Цель исследований – выяснить, какие внешние факторы могут быть движущей силой роста и популяционной динамики двух близких по экологии видов желетелого макропланктона (сцифомедуза Aurelia aurita и гребневик Pleurobrachia pileus), играющих важную роль в функционировании пелагической экосистемы Черного моря.Материалы и методы. Состояние популяций желетелых оценивали по данным, полученным в 2000‐2014 гг. в районе внешнего шельфа Севастопольской бухты, где на 3‐х станциях в 2‐х милях от берега (глубина 50‐ 70 м) ежемесячно отбирали количественные пробы мезо‐ и макропланктона. Сопутствующие погодные и гидрологические процессы оценивали по открытым базам данных.Результаты. Установлено, что удельная скорость линейного роста желетелых зависит от погодно‐гидрологических условий в зимне‐весенние месяцы и от года к году колеблется в пределах 0.85‐1.02% сут.‐1 – у A. aurita, и 0.27‐0.47% сут.‐1 – у P. pileus.Заключение. Внешние факторы неодинаково контролируют рост этих видов. Субширотный перенос теплых воздушных масс, сопровождающийся повышением температуры морской воды, усилением речного стока и циркуляции водных масс в море, приводит к увеличению скорости роста гребневика, субмеридиональное поступление холодного и сухого воздуха – к увеличению скорости роста медузы. В обоих случаях на рост влияет система трофических отношений, по‐разному формирующаяся в зависимости от характера погоды в феврале‐мае. Рост гребневика обеспечивает синхронное повышение биомассы рачкового зоопланктона, рост медузы – развитие микропланктона и мезопланктонных микрофагов.

Aim. The aim of the study is to find out what external factors may be the driving force behind the growth and population dynamics of two ecologically similar species of gelatinous macroplankton (scyphomedusa Aurelia aurita and ctenophore Pleurobrachia pileus), which play an important role in the functioning of the pelagic ecosystem of the Black Sea.Material and Methods. The state of zooplankton populations was estimated by data obtained in 2000‐2014 in the outer shelf of Sevastopol Bay, where monthly quantitative samples of meso‐ and macroplankton were taken 2 miles of the coast (depth 50‐70 m). Weather and hydrology related changes were assessed using open databases.Results. It was been established that size‐specific growth rate of these species depends on weather and hydrological conditions in the winter‐spring months and varies from 0.85 to 1.02% day‐1 and from 0.27 to 0.47% day‐1 for A. aurita and P. pileus, respectively.Conclusions. External factors unequally affect the growth of these species. Sub‐ latitudinal transit of warm air masses, accompanied by increased river flow and seawater circulation, activates the growth of the ctenophore, while sub‐meridional propagation of cold and dry air increases the growth rate of the jellyfish. In both cases, somatic growth is influenced by trophic relations, differing depending on the weather in February‐May. The changes in growth of ctenophore occur in parallel with synchronous variations in biomass of crustacea, while that in jellyfish may relate to an abundance of microplankton and its mesoplanktonic consumers.

Желетелый макропланктонгребневик Pleurobrachiaмедуза Aureliaбиомассаскорость ростамежгодовые измененияЧерное мореGelatinous macroplanktonctenophore Pleurobrachiajellyfish Aureliabiomassgrowth rateinterannual changesBlack SeaРабота подготовлена по теме № АААА‐А18118021490093‐4 государственного задания ФГБУН ФИЦ «Институт биологии южных морей имени А.О. Ковалевского РАН» а также гранту РФФИ №18‐44920022The studies were conducted under the State Project of «A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS» No. АААА‐А18‐118021490093‐4 and the grant No. 18‐44‐920022 from the Russian Foundation for Basic Research.ReferencesMutlu E., Bingel F. Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. I. Pleurobrachia pileus // Marine Biology. 1999. V. 135. Iss. 4. P. 589-601. DOI: 10.1007/s002270050660Mutlu E., Bingel F. Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. I. Pleurobrachia pileus. Marine Biology, 1999, vol. 135, iss. 4, pp. 589-601. DOI: 10.1007/s002270050660Anninsky B.E. Organic composition and ecological energetics of jellyfish Aurelia aurita L. (Cnidaria, Scyphozoa) under the Black Sea conditions // Trophic relationships and Food Supply of Heterotrophic Animals in the Pelagic Ecosystem of the Black Sea. Eds. Shulman G.E. et al. Istanbul. Turkey: Black Sea Commission Publications. 2009. P. 99-160.Anninsky B. E. Organic composition and ecological energetics of jellyfish Aurelia aurita L. (Cnidaria, Scyphozoa) under the Black Sea conditions. Trophic relationships and Food Supply of Heterotrophic Animals in the Pelagic Ecosystem of the Black Sea. Eds. Shulman G.E. et al. Istanbul. Turkey: Black Sea Commission Publications, 2009, pp. 99-160.Riisgård H.U., Goldstein J., Lundgreen K., Lüskow F. Jellyfish and Ctenophores in the Environmentally Degraded Limfjorden (Denmark) During 2014 - Species Composition, Population Densities and Predation Impact // Fisheries and Aquaculture Journal. 2015. V. 6. Iss. 3. 137 p. DOI: 10.4172/2150-3508.1000137Riisgård H.U., Goldstein J., Lundgreen K., Lüskow F. Jellyfish and Ctenophores in the Environmentally Degraded Limfjorden (Denmark) During 2014 - Species Composition, Population Densities and Predation Impact. Fisheries and Aquaculture Journal, 2015, vol. 6, iss. 3, 137 p. DOI: 10.4172/2150-3508.1000137Vinogradov M.E., Shushkina E.A., Mikaelyan A.S., Nezlin N.P. Temporal (seasonal and interannual) changes of ecosystem of the open waters of the Black Sea // Environmental Degradation of the Black Sea: Challenges and Remedies. Eds. Besiktepe S., Unluata U., Bologa A.S. Dordrecht et al.: Kluwer Acad. Publ. 1999. NATO ASI Series V. 56. P. 109-129.Vinogradov M.E., Shushkina E.A., Mikaelyan A.S., Nezlin N.P. Temporal (seasonal and interannual) changes of ecosystem of the open waters of the Black Sea. Environmental Degradation of the Black Sea: Challenges and Remedies. Eds. Besiktepe S., Unluata U., Bologa A.S. Dordrecht et al.: Kluwer Acad. Publ., 1999, NATO ASI Series, vol. 56, pp. 109-129.Daufresne M., Lengfellner K., Sommer U. Global warming benefits the small in aquatic ecosystems // Proceedings of the National Academy of Sciences. 2009. V. 106. Iss. 31. P. 12788-12793. DOI: 10.1073/pnas.0902080106Daufresne M., Lengfellner K., Sommer U. Global warming benefits the small in aquatic ecosystems. Proceedings of the National Academy of Sciences, 2009, vol. 106, iss. 31, pp. 12788-12793. DOI: 10.1073/pnas.0902080106Condon R.H., Duarte C.M., Pitt K.A., Robinson K.L., Lucas C.H., Sutherland K.R., Mianzan H.W., Bogeberg M., Purcell J.E., Decker M.B., Uye S., Madin L.P., Brodeur R.D., Haddock S.H.D., Malej A., Parry G.D., Eriksen E., Quinones J., Acha M., Harvey M., Arthur J.M., Graham, W.M. Recurrent jelly fish blooms are a consequence of global oscillations // Proceedings of the National Academy of Sciences. 2013. V. 110. Iss. 3. P. 1000-1005. DOI: 10.1073/pnas.1210920110Condon R.H., Duarte C.M., Pitt K.A., Robinson K.L., Lucas C.H., Sutherland K.R., Mianzan H.W., Bogeberg M., Purcell J.E., Decker M.B., Uye S., Madin L.P., Brodeur R.D., Haddock S.H.D., Malej A., Parry G.D., Eriksen E., Quinones J., Acha M., Harvey M., Arthur J.M., Graham, W.M. Recurrent jelly fish blooms are a consequence of global oscillations. Proceedings of the National Academy of Sciences, 2013, vol. 110, iss. 3, pp. 1000-1005. DOI: 10.1073/pnas.1210920110Kogovšek T., Vodopivec M., Raicich F., Uye S., Malej A. Comparative analysis of the ecosystems in the northern Adriatic Sea and the Inland Sea of Japan: Can anthropogenic pressures disclose jellyfish outbreaks? // Science of the Total Environment. 2018. V. 626. P 982-994. DOI: 10.1016/j.scitotenv.2018.01.011Kogovšek T., Vodopivec M., Raicich F., Uye S., Malej A. Comparative analysis of the ecosystems in the northern Adriatic Sea and the Inland Sea of Japan: Can anthropogenic pressures disclose jellyfish outbreaks? Science of the Total Environment, 2018, vol. 626, pp. 982-994. DOI: 10.1016/j.scitotenv.2018.01.011Stanev E.V., Peneva E., Chtirkova B. Climate change and regional ocean water mass disappearance: Case of the Black Sea // Journal of Geophysical Research: Oceans. 2019. V. 124. Iss. 7. P. 4803-4819. DOI: 10.1029/2019JC015076Stanev E.V., Peneva E., Chtirkova B. Climate change and regional ocean water mass disappearance: Case of the Black Sea. Journal of Geophysical Research: Oceans, 2019, vol. 124, iss. 7, pp. 4803-4819. DOI: 10.1029/2019JC015076Mutlu E. Recent distribution and size structure of gelatinous organisms in the southern Black Sea and their interactions with fish catches // Marine Biology. 2009. V. 156. P. 935-957. DOI: 10.1007/s00227-009-1139-8Mutlu E. Recent distribution and size structure of gelatinous organisms in the southern Black Sea and their interactions with fish catches. Marine Biology, 2009, vol. 156, pp. 935-957. DOI: 10.1007/s00227-009-1139-8Anninsky B.E., Finenko G.A., Datsyk N.A., Ignatyev S.M. Gelatinous macroplankton in the Black Sea in the autumn of 2010 // Oceanology. 2013. V. 53. Iss. 6. P. 676-685. DOI: 10.1134/S0001437013060015Anninsky B.E., Finenko G.A., Datsyk N.A., Ignatyev S.M. Gelatinous macroplankton in the Black Sea in the autumn of 2010. Oceanology, 2013, vol. 53, iss. 6, pp. 676-685. DOI: 10.1134/S0001437013060015Finenko G.A., Anninsky B.E., Datsyk N.A. Mnemiopsis leidyi A. Agassiz, 1865 (Ctenophora: Lobata) in the Inshore Areas of the Black Sea: 25 Years After Its Outbreak // Russian Journal of Biological Invasions. 2018. V. 9. Iss. 1. P. 86-93. DOI: 10.1134/S2075111718010071Finenko G.A., Anninsky B.E., Datsyk N.A. Mnemiopsis leidyi A. Agassiz, 1865 (Ctenophora: Lobata) in the Inshore Areas of the Black Sea: 25 Years After Its Outbreak. Russian Journal of Biological Invasions, 2018, vol. 9, iss. 1, pp. 86-93. DOI: 10.1134/S2075111718010071Заика В.Е. Аллометрия скоростей роста, питания и метаболизма гребневиков и медуз // Экология моря. 2002. Т. 59. С. 42-47.Zaika V.E. Allometry of growth, feeding and metabolic rates in ctenophores and medusae. Ekologiya morya [Ecologiya moray]. 2002, vol. 59, pp. 42-47. (In Russian)Oguz T., Velikova V., Cociasu A., Korchenko A. The state of eutrophication // State of the Environment of the Black Sea (2001-2006/7). Ed. by T. Oguz. Publ. of the Comm. on the Protection of the Black Sea against Pollution (BSC). Istanbul, 2008. P. 83-112.Oguz T., Velikova V., Cociasu A., Korchenko A. The state of eutrophication. State of the Environment of the Black Sea (2001-2006/7). Ed. by T.Oguz. Publ. of the Comm. on the Protection of the Black Sea against Pollution (BSC), Istanbul, 2008, pp. 83-112.Dolinaj D., Leščešen I., Pantelić M., Urošev M., Milijašević Joksimović D. Danube River Discharge at Bezdan Gauging Station (Serbia) and its correlation with Atmospheric Circulation Patterns // Geographica Pannonica. V. 23. Iss. 1. P. 14-22. DOI: 10.5937/gp2318514Dolinaj D., Leščešen I., Pantelić M., Urošev M., Milijašević Joksimović D. Danube River Discharge at Bezdan Gauging Station (Serbia) and its correlation with Atmospheric Circulation Patterns. Geographica Pannonica, vol. 23, iss. 1, pp. 14-22. DOI: 10.5937/gp23-18514Oguz T., Dippner J.W., Kaymaz Z. Climatic regulation of the Black Sea hydro-meteorological and ecological properties at interannual-to-decadal time scales // Journal of Marine Systems. 2006. V. 60. Iss. 3-4. P. 235-254. DOI: 10.1016/j.jmarsys.2005.11.011Oguz T., Dippner J.W., Kaymaz Z. Climatic regulation of the Black Sea hydro-meteorological and ecological properties at interannual-to-decadal time scales. Journal of Marine Systems, 2006, vol. 60, iss. 3-4, pp. 235-254. DOI: 10.1016/j.jmarsys.2005.11.011Иванов В.А., Белокопытов В.Н. Океанография Черного моря. Севастополь: ЭКОСИ-Гидрофизика, 2013. 210 с.Ivanov V.A., Belokopytov V.N. Okeanografiya Chernogo morya [Oceanography of the Black Sea]. Sevastopol, ECOSY-Gidrofizika Publ., 2013, 210 p. (In Russian)Krichak S.O., Kishcha P., Alpert P. Decadal trends of main Eurasian oscillations and the Eastern Mediterranean precipitation // Theoretical and Applied Climatology. 2002. V. 72. P. 209-220. DOI: 10.1007/s007040200021Krichak S.O., Kishcha P., Alpert P. Decadal trends of main Eurasian oscillations and the Eastern Mediterranean precipitation. Theoretical and Applied Climatology, 2002, vol. 72, pp. 209-220. DOI: 10.1007/s007040200021Lynam C.P., Hay S.J., Brierley A.S. Interannual variability in abundance of North Sea jellyfish and links to the North Atlantic Oscillation // Limnology and Oceanography. 2004. V. 49. P. 637-643. DOI: 10.4319/lo.2004.49.3.0637Lynam C.P., Hay S.J., Brierley A.S. Interannual variability in abundance of North Sea jellyfish and links to the North Atlantic Oscillation. Limnology and Oceanography, 2004, vol. 49, pp. 637-643. DOI: 10.4319/lo.2004.49.3.0637Walraven L.V., Langenberg V.T., Dapper R., Witte J.Ij., Zuur A.F., van der Veer H.W. Long-term patterns in 50 years of scyphomedusae catches in the western Dutch Wadden Sea in relation to climatechange and eutrophication // Journal of Plankton Research. 2015. V. 37. Iss. 1. P. 151-167. DOI: 10.1093/plankt/fbu088Walraven L.V., Langenberg V.T., Dapper R., Witte J.Ij., Zuur A.F., van der Veer H.W. Long-term patterns in 50 years of scyphomedusae catches in the western Dutch Wadden Sea in relation to climatechange and eutrophication. Journal of Plankton Research, 2015, vol. 37, iss. 1, pp. 151-167. DOI: 10.1093/plankt/fbu088Costello J.H., Colin S.P. Morphology, fluid motion and predation by the scyphomedusa Aurelia aurita // Marine Biology. 1994. V. 121. P. 327-334. DOI: 10.1007/BF00346741Costello J.H., Colin S.P. Morphology, fluid motion and predation by the scyphomedusa Aurelia aurita. Marine Biology, 1994, vol. 121, pp. 327-334. DOI: 10.1007/BF00346741Anninsky B.E., Finenko G.A., Datsyk N.A., Kideys A.E. Trophic ecology and assessment of the predatory impact of the Moon jellyfish Aurelia aurita (Linnaeus, 1758) on zooplankton in the Black Sea // Cahiers de Biologie Marine. 2020. V. 61. Iss. 1. P. 33-46. DOI: 10.21411/CBM.A.96DD01AAAnninsky B.E., Finenko G.A., Datsyk N.A., Kideys A.E. Trophic ecology and assessment of the predatory impact of the Moon jellyfish Aurelia aurita (Linnaeus, 1758) on zooplankton in the Black Sea. Cahiers de Biologie Marine, 2020, vol. 61, iss. 1, pp. 33-46. DOI: 10.21411/CBM.A.96DD01AAPurcell J.E., Uye S.-I., Lo W.-T. Anthropogenic causes of jellyfish blooms and direct consequences for humans: a review // Marine Ecology Progress Series. 2007. V. 350. P. 153-174. DOI: 10.3354/meps07093Purcell J.E., Uye S.-I., Lo W.-T. Anthropogenic causes of jellyfish blooms and direct consequences for humans: a review. Marine Ecology Progress Series, 2007, vol. 350, pp. 153-174. DOI: 10.3354/meps07093Stoecker D., Michaels A.E., Davis L.H. Grazing by the jellyfish, Aurelia aurita, on microzooplankton // Journal of Plankton Research. 1987. V. 9. Iss. 5. P. 901-915. DOI: 10.1093/plankt/9.5.901Stoecker D., Michaels A.E., Davis L.H. Grazing by the jellyfish, Aurelia aurita, on microzooplankton. Journal of Plankton Research, 1987, vol. 9, iss. 5, pp. 901-915. DOI: 10.1093/plankt/9.5.901Malej A., Turk V., Lučić D., Benović A. Direct and indirect trophic interactions of Aurelia sp. (Scyphozoa) in stratified marine environment (Mljet Lakes, Adriatic Sea) // Marine Biology. 2006. V. 151. P. 827-841. DOI: 10.1007/s00227-006-0503-1Malej A., Turk V., Lučić D., Benović A. Direct and indirect trophic interactions of Aurelia sp. (Scyphozoa) in stratified marine environment (Mljet Lakes, Adriatic Sea). Marine Biology, 2006, vol. 151, pp. 827-841. DOI: 10.1007/s00227-006-0503-1Turk V., Lučić D., Flander-Putrle V., Malej A. Feeding of Aurelia sp. (Scyphozoa) and links to the microbial food web // Marine Ecology. 2008. V. 29. P. 1-11. DOI: 10.1111/j.1439-0485.2008.00250.xTurk V., Lučić D., Flander-Putrle V., Malej A. Feeding of Aurelia sp. (Scyphozoa) and links to the microbial food web. Marine Ecology, 2008, vol. 29, pp. 1-11. DOI: 10.1111/j.1439-0485.2008.00250.xZoccarato L., Celussi M., Pallavicini A., Umani S.F. Aurelia aurita Ephyrae Reshape a Coastal Microbial Community // Frontiers in Microbiology. 2016. V. 7. Article 749. DOI: 10.3389/fmicb.2016.00749Zoccarato L., Celussi M., Pallavicini A., Umani S.F. Aurelia aurita Ephyrae Reshape a Coastal Microbial Community. Frontiers in Microbiology, 2016, vol. 7, article 749. DOI: 10.3389/fmicb.2016.00749Skikne S.A., Sherlock R.E. Robison B.H. Uptake of dissolved organic matter by ephyrae of two species of scyphomedusae // Journal of Plankton Research. 2009. V. 31. N 12. P. 1563-1570. DOI: 10.1093/plankt/fbp088Skikne S.A., Sherlock R.E. Robison B.H. Uptake of dissolved organic matter by ephyrae of two species of scyphomedusae. Journal of Plankton Research, 2009, vol. 31, no. 12, pp. 1563-1570. DOI: 10.1093/plankt/fbp088Aytan U., Feyzioglu A.M., Valente A., Agirbas E., Fileman E. S. Microbial plankton communities in the coastal southeastern Black Sea: biomass, composition and trophic interactions // Oceanologia. 2018. V. 60. Iss. 2. P. 139-152. DOI: 10.1016/j.oceano.2017.09.002Aytan U., Feyzioglu A.M., Valente A., Agirbas E., Fileman E. S. Microbial plankton communities in the coastal southeastern Black Sea: biomass, composition and trophic interactions. Oceanologia, 2018, vol. 60, iss. 2, pp. 139-152. DOI: 10.1016/j.oceano.2017.09.002Ducklow H.W., Hansell D.A., Morgan J.A. Dissolved organic carbon and nitrogen in the Western Black Sea // Marine Chemistry. 2007. V. 105. P. 140-150. DOI: 10.1016/j.marchem.2007.01.015Ducklow H.W., Hansell D.A., Morgan J.A. Dissolved organic carbon and nitrogen in the Western Black Sea. Marine Chemistry, 2007, vol. 105, pp. 140-150. DOI: 10.1016/j.marchem.2007.01.015Casanova J.-P., Barthélémy R.-M., Duvert M., Faure E. Chaetognaths feed primarily on dissolved and fine particulate organic matter, not on prey: implications for marine food webs // Hypotheses in the Life Sciences. 2012. V. 2. Iss. 1. P. 20-29.Casanova J.-P., Barthélémy R.-M., Duvert M., Faure E. Chaetognaths feed primarily on dissolved and fine particulate organic matter, not on prey: implications for marine food webs. Hypotheses in the Life Sciences, 2012, vol. 2, iss. 1, pp. 20-29.Pomeroy L.R., Williams P.J. leB., Azam F., Hobbie J.E. The Microbial Loop // Oceanography. 2007. V. 20. N 2. P. 28-33. DOI: 10.5670/oceanog.2007.45Pomeroy L.R., Williams P.J. leB., Azam F., Hobbie J.E. The Microbial Loop. Oceanography, 2007, vol. 20, no. 2, pp. 28-33. DOI: 10.5670/oceanog.2007.4536.Höhn D.P., Lucas C.H, Thatje S. Respiratory response to temperature of three populations of Aurelia aurita polyps in northern Europe // PLoS ONE. 2017. V. 12. Iss. 5. e0177913. DOI: 10.1371/journal.pone.0177913Höhn D.P., Lucas C.H, Thatje S. Respiratory response to temperature of three populations of Aurelia aurita polyps in northern Europe. PLoS ONE, 2017, vol.12, iss. 5, e0177913. DOI: 10.1371/journal.pone.0177913Shulman G.E., Nikolsky V.N., Yuneva T.V., Yunev O.A., Bat L., Kideys A.E. Influence of global climatic changes and regional anthropogenic factors on the Black Sea sprat and anchovy condition. Trophic relationships and Food Supply of Heterotrophic Animals in the Pelagic Ecosystem of the Black Sea. Eds. Shulman G.E. et al. Istanbul. Turkey: Black Sea Commission Publications, 2009. P. 273-298.Shulman G.E., Nikolsky V.N., Yuneva T.V., Yunev O.A., Bat L., Kideys A.E. Influence of global climatic changes and regional anthropogenic factors on the Black Sea sprat and anchovy condition. Trophic relationships and Food Supply of Heterotrophic Animals in the Pelagic Ecosystem of the Black Sea. Eds. Shulman G.E. et al. Istanbul. Turkey: Black Sea Commission Publications, 2009, pp. 273-298.Аннинский Б.Е., Игнатьев С.М., Финенко Г.А., Дацык Н.А. Желетелый макропланктон открытой пелагиали и шельфа Черного моря: распределение осенью 2016 г. и межгодовые изменения биомассы и численности // Морской биологический журнал. 2019. Т. 4. N 3. С. 3-14. DOI: 10.21072/mbj.2019.04.3.01Anninsky B.E., Ignatyev S.M., Finenko G.A., Datsyk N.A. Gelatinous macroplankton of the open pelagial and shelf of the Black Sea: distribution in in autumn 2016 and interannual changes in biomass and abundance. Marine Biological Journal, 2019, vol. 4, no. 3, pp. 676-685. (In Russian)Klimova T., Podrezova P. Seasonal distribution of the Black Sea ichthyoplankton near the Crimean Peninsula // Regional Studies in Marine Science. 2018. V. 24. P. 260-269. DOI: 10.1016/j.rsma.2018.08.013Klimova T., Podrezova P. Seasonal distribution of the Black Sea ichthyoplankton near the Crimean Peninsula. Regional Studies in Marine Science, 2018, vol. 24, pp. 260-269. DOI: 10.1016/j.rsma.2018.08.013

The authors declare that there are no conflicts of interest present.