Abstract |
Studies of the benthos in nearshore water areas within Protected Areas along the Black Sea are an important ecological task. In this connection, protected habitats can be recognised as intact natural areas, where the entire natural complex is preserved with its typical taxonomic structure and biodiversity features. This study is based on the obtained quantitative data on the biodiversity of benthic assemblages in coastal Protected Areas. It aims to study the structure of polychaete communities as inhabitants of fouling areas at sublittoral (depth 0–3 m) rocky/boulder substrates in four Protected Areas along the Black Sea (Cape Martyan State Nature Reserve, Karadag State Nature Reserve, Opukskiy State Nature Reserve, and Utrish State Nature Reserve). The studied sites differ in their geographic location, orographic shape of the coastal strip, features of underwater substrates, the pattern of spatial coverage and species composition of macroalgae foulings and other environmental parameters. The purpose of this study was also to make a prognostic estimation of the expected species richness (Sexp) and comparative analysis of the structural organisation of polychaete complexes on rocky substrates in the selected coastal Protected Areas. The prognostic modelling of polychaete species richness was based on the known extrapolation algorithms (estimators Chao-2, Jackknife-2, and bootstrap). The results show that the expected level of species richness (Sexp) exceeds the actually detected (Sobs) number of species in each of the four Protected Areas by 20–95%, depending on the estimator and total number of collected samples. In all sublittoral water areas, it was found that with an increase in the number of samples the accumulation curves do not reach a horizontal asymptote. This indicates an underestimated species richness of polychaetes. At the rocky sublittoral zone of the Cape Martyan State Nature Reserve, the fastest increase in Sexp values was revealed with an increase in the number of samples taken. This relationship can be determined by heterogeneity of microhabitats in this area, where the increased number of polychaete species is also formed. Considering the other Protected Areas, the extrapolative curves of species richness have a flatter mode and a smaller expected number of species (Sexp), despite the larger number of samples analysed. According to the results of the clustering and ordination analyses, based on the species composition and abundance of polychaetes, almost all samples taken were grouped into separate clusters corresponding to each of the four studied Protected Areas. The reliability of structural differences between polychaete complexes was statistically confirmed by pairwise comparison of three areas (Utrish, Karadag, and Martyan). The comparison of sites conditionally named Utrish and Opukskiy demonstrated that structural differences between polychaete complexes were statistically insignificant probably due to the affinity of the geomorphologic pattern of hard-bottoms and composition of fouling vegetation in sublittoral zones of both Protected Areas. The specific polychaete complexes have been formed on rocky/boulder substrates within each of the investigated Protected Areas, being different in their taxonomic structure and quantitative characteristics of the certain species. For each polychaete complex, the most significant species were identified, which determine both the intracomplex taxonomic structure (indicating species) and structural differences between the complexes (discriminating species). In the polychaete complex of the rocky sublittoral in the Karadag State Nature Reserve such indicating species were Nereis zonata, Platynereis dumerilii, and Syllis gracilis. In the Cape Martyan State Nature Reserve and Utrish State Nature Reserve, the indicating taxa were Polyophthalmus pictus, Nereis sp., and Pholoe inornata. Finally, in the Opukskiy State Nature Reserve, Notomastus latericeus and Pholoe inornata were recognised as indicating species. Most of these species may also be considered as discriminating species. Parameters of their quantitative development reflect the structural features of polychaete complexes formed under the influence of specific environmental factors in the sublittoral zone of each of the four studied coastal Protected Areas. The obtained results of the structural-faunistic analysis of polychaete complexes can be applied for comparative assessment of the biodiversity features of benthic communities during the comprehensive ecological monitoring of the coastal Protected Areas. |
References |
Alexandrov B.G., Minicheva G.G., Zaitsev Yu.P. 2017. Black Sea Network of Marine Protected areas: European approaches and adaptation to expansion and monitoring in Ukraine. In: P.D. Goriup (Ed.): Management of Marine Protected Areas: A network perspective, 1st edition. New York: John Wiley & Sons Ltd. P. 227–246. Boltacheva N.A., Revkov N.K., Bondarenko L.V. Makarov M.V., Kopiy V.G., Timofeev V.A., Mazlumyan S.A. 2010. Macrozoobenthos of water area in the Karadag State Nature Reserve. In: A.L. Morozova (Ed.): Chronicle of the Nature. Vol. 25. Simferopol: N.Orianda. P. 150–174. [In Russian] Boltacheva N.A., Kovaleva M.A., Makarov M.V., Bondarenko L.V. 2015. Macrofauna on the rocks: long-term changes in the upper subtidal zone of Karadag (Black Sea). In: A.V. Gaevskaya, A.L. Morozova (Eds.): 100 years of the T.I. Vyazemsky Karadag research station: Proceedings. Simferopol: N.Orianda. P. 530–548. [In Russian] Bondarenko L.V., Boltacheva N.A., Kopiy V.G., Timofeev V.A. 2016. Macrozoobenthic organisms of soft bottom sediments biotope of Opuk Nature Reserve. Ekosistemy 7(37): 19–25. [In Russian] Chao A. 1987. Estimating the population-size for capture-recapture data with unequal catchability. Biometrics 43(4): 783–791. DOI: 10.2307/2531532 Chatfield C., Collins A. 1980. Introduction to multivariate analysis. London: Chapman & Hill. 329 p. Chikina M.V., Spiridonov V.A., Naumov A.D. 2020. Modiolus modiolus Communities of Onega Bay, White Sea: How Stable Are They over Time and Space? Biology Bulletin 47(9): 1099–1114. DOI: 10.1134/S1062359020090058 Clarke K.R., Gorley R.N. 2006. PRIMER v6: User Manual. Plymouth: PRIMER-E Ltd. 190 p. Clarke K.R., Warwick R.M. 2001. Change in marine communities: an approach to statistical analysis and interpretation, 2nd edition. Plymouth: PRIMER-E Ltd. 154 p. Colwell R.K., Coddington J.A. 1994. Estimating terrestrial biodiversity through extrapolation. Philosophical transactions of the Royal Society of London. Series B, Biological sciences 345(1311): 101–118. DOI: 10.1098/rstb.1994.0091 Denisenko N.V. 2009. Composition and distribution of sublittoral zoobenthos in Kemskaya Bay of the White Sea. Oceanology 49(6): 788–800. DOI: 10.1134/S0001437009060058 Escaravage V., Herman M.J., Merckx B., Włodarska-Kowalczuk M., Amouroux J.M., Degraer S., Grémare A., Heip C.H.R., Hummel H., Karakassis I., Labrune C., Willems W. 2009. Distribution patterns of macrofaunal species diversity in subtidal soft sediments: biodiversity-productivity relationships from the MacroBen database. Marine Ecology Progress Series 382: 253–264. DOI: 10.3354/meps08008 Field J.G., Clarke K.R., Warwick R.M. 1982. A practical strategy for analyzing multispecies distribution patterns. Marine Ecology Progress Series 8: 37–52. Foggo A., Attrill M.J., Frost M.T., Rowden A.A. 2003. Estimating marine species richness: an evaluation of six extrapolative techniques. Marine Ecology Progress Series 248: 15–26. DOI: 10.3354/meps248015 Hellmann J.J., Fowler G.W. 1999. Bias, precision and accuracy of four measures of species richness. Ecological Applications 9(3): 824–834. DOI: 10.1890/1051-0761(1999)009[0824:BPAAOF]2.0.CO;2 Kiseleva M.I. 1981. Soft-bottom benthos of the Black Sea. Kiev: Naukova Dumka. 163 p. [In Russian] Kiseleva G.A., Kolova K.A., Molchanova Yu.V. 2010. Polychaete in the tangle of the algae on the aquatorium of Karadag. Ecosystems, Their Optimization and Conservation 3: 42–49. [In Russian] Kolyuchkina G.A., Semin V.L., Basin A.B., Kuznetsova A.V., Grigorenko K.S., Lyubimov I.V., Simakova U.V. 2017. Current status of macrozoobenthos of soft sediments in the Utrish State Nature Reserve. In: Abrau: terrestrial and adjacent marine ecosystems: structure, biodiversity and conservation. Vol. 4. P. 228–240. [In Russian] Kolyuchkina G.A., Syomin V.L., Simakova U.V., Mokievsky V.O. 2018. Presentability of the Utrish Nature Reserve's benthic communities for the North Caucasian Black Sea Coast. Nature Conservation Research 3(4): 1–16. DOI: 10.24189/ncr.2018.065 Kopiy V.G. 2017. Macrozoobenthos in the pseudolittoral zone of the Crimean reserves. Proceedings of the T.I. Vyazemsky Karadag research station – Natural Reserve RAS 1(3): 29–37. [In Russian] Kopiy V.G. 2019. Polychaetes of fouling substrates at the costal marine reserves of the Crimea and the Caucasus. Biota and Environment of Natural Areas 2: 48–65. DOI: 10.25808/26186764.2019.92.22.004 [In Russian] Kopiy V.G., Bondarenko L.V. 2016. Macrozoobenthos of splash zone of the eastern coast of Crimea. In: I.S. Turbanov (Ed.): Prospective and Problems of Modern Hydrobiology. Yaroslavl: Filigran. P. 89–90. [In Russian] Korzhenevskiy V.V., Bokov V.A., Dulitskiy A.I. (Eds.). 1997. Biodiversity of the Crimea: assessment and conservation needs. Materials for international workshop (November, 1997, Gurzuf). Simpheropol: SONAT. 131 p. [In Russian] Kostenko N.S., Dikiy E.A., Alekseeva S.P. 2004. Phytobenthos of the South-Eastern part of Crimean coast of the Black Sea. In: Karadag: hydrobiological investigations. Book 2. Simpheropol: Sonat. P. 66–84. [In Russian] Kovalevа M.A., Boltacheva N.A, Makarov M.V., Bondarenko L.V. 2014. The fouling community on the natural hard substrata of the Karadag State Nature Reserve (Black Sea). Ecosystems, Their Optimization and Conservation 10: 77–81. [In Russian] Kuftarkova E.A., Kovrigina N.P., Bobko N.I. 2004. Hydrochemical characteristics of the waters of the Sudak-Karadag beach. Karadag: hydrobiological research. In: Karadag: hydrobiological investigations. Book 2. Simpheropol: Sonat. P. 12–27. [In Russian] Kukharev I.L., Sheremetyev A.V. 2013. Utrish State Nature Reserve. Atlas. Proceedings of the Utrish State Nature Reserve 2: 16–19. [In Russian] Losovskaya G.V. 1977. Polychaete ecology of the Black Sea. Kiev: Naukova Dumka. 92 p. [In Russian] Makarov M.V., Kovaleva M.A., Boltacheva N.A., Kopiy V.G., Bondarenko L.V. 2015. Macrozoobenthos of natural hard substrates in water areas adjoining to the Kerch peninsula. Scientific Notes of Ternopіl National Pedagogical University. Series Biology 3–4(64): 425–428. [In Russian] Makarov M.V., Bondarenko L.V., Kopiy V.G., Kovaleva M.A. 2020. Macrozoobenthos of the natural hard fouling substrates (boulders) in water area adjacent to the Utrish State Nature Reserve (Black Sea). Proceedings of the Utrish State Nature Reserve 5: 233–239. [In Russian] Maslov I.I., Kraynyuk E.S., Sarkina I.S., Kostin S.Yu., Sergeyenko A.L. 2011. Main directions and results of scientific and environmental activities of the Nature Conservation Department of the NBG, Cape Martyan State Nature Reserve (1973–2010). Bulletin of the State Nikitsky Botanical Gardens 100: 29–39. [In Russian] Milchakova N.A., Aleksandrov V.V., Bondareva L.V., Pankeeva T.V., Chernysheva E.V. 2015. Marine Protected Areas of the Crimea. Scientific Handbook. Simferopol: N. Orianda. 312 p. [In Russian] Morozova A.L., Smirnova Yu.D., Gasnikov S.V. 2008. The influence of natural and anthropogenic factors on the transformation of marine coastal ecosystems of the Karadag State Nature Reserve. In: B.N. Panov (Ed.): Modern ecological problems of the Azov-Black Sea Region. Kerch: YugNIRO. P. 61–67. [In Russian] Nevrova E.L., Snigireva A.A., Petrov A.N., Kovaleva G.V. 2015. Guidelines for quality control of the Black Sea. Microphytobenthos. Simferopol: N. Orianda. 176 p. [In Russian] Petrov A.N. 1998. Near-shore water areas of the Crimea: working out of principles and criteria for creation of new protected areas upon development of municipal and recreation activities. Ecologiya Morya 47: 17–23. [In Russian] Petrov A.N. 2002. Near-shore water areas. In: V.A. Bokov (Ed.): Perspectives of organization of the unified network of Protected Areas in Crimea. Simferopol: Krymuchpedgiz. P. 170–181. [In Russian] Petrov A.N. 2005. Assessment of long-term recreational and marifarming impact upon nearshore seascapes and benthic community (case study: Laspi bay, Crimea, the Black Sea). In: P. Lasserre, P. Viaroli, P. Campostrini (Eds.): Lagoons and Coastal Wetlands in the Global Change Context: Impacts and Management Issues. Dordrecht: Springer Netherlands. P. 293–300. Petrov A.N., Nevrova E.L. 2004. Comparative analysis of taxocene structures of benthic diatoms (Bacillariophyta) in regions with different level of technogenic pollution (Black Sea, Crimea). Marine Ecological Journal 3(2): 72–83. [In Russian] Petrov A.N., Nevrova E.L. 2013. Extrapolative estimation of benthic diatoms (Bacillariophyta) species diversity in different marine habitats of the Crimea (Black Sea). International Journal of Biodiversity 2013: 975459. DOI: 10.1155/2013/975459 Petrov A.N., Nevrova E.L. 2016. Species richness estimation of benthos assemblages at different sediment pollution level and sampling efforts. In: A.V. Gaevskaya (Ed.): Marine Biological Research: Achievements and Perspectives. Vol. 2. Sevastopol: EKOSI-Gidrofizika. P. 385–389. [In Russian] Simakova U.V. 2009. Influence of the sea bottom relief on the Cystoseira communities of the North Caucasian coast of the Black Sea. Oceanology 49(5): 672–680. DOI: 10.1134/S0001437009050087 Sinegub I.A. 2004. Macrofauna of upper rocky sublittoral in the Black Sea near Karadag. In: Karadag: hydrobiological investigations. Book 2. Simpheropol: Sonat. P. 121–132. [In Russian] Ugland K.I., Gray J.S. 2004. Estimation of species richness: analysis of the methods developed by Chao and Karakassis. Marine Ecology Progress Series 284: 1–8. DOI: 10.3354/meps284001 Vershinin A.O., Pankov S.L. 2008. Concerning the necessity of organization of marine reserve within framework of the Utrish Reserve project. In: V.V. Kovalev, S.A. Trepet (Eds.): Prospective for development of Protected Areas and tourism at Northern Caucasus. Maikop: Kachestvo. P. 31–36. [In Russian] Walther B.A., Morand S. 1998. Comparative performance of species richness estimation methods. Parasitology 116(4): 395–405. DOI: 10.1017/S0031182097002230 Walther B.A., Moore J.L. 2005. The concepts of bias, precision and accuracy, and their use in testing the performance of species richness estimators, with a literature review of estimator performance. Ecography 28(6): 815–829. DOI: 10.1111/j.2005.0906-7590.04112.x WoRMS. 2018. World register of Marine Species. Available from: https://www.marinespecies.org/ |