Benthic Diatoms of the Russian Waters of the Sea of Japan and Adjacent Sea Areas

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Benthic Diatoms of the Russian Waters of the Sea of Japan and Adjacent Sea Areas A. A. Beguna,* and L. I. Ryabushkob aZhirmunsky

National Scientific Center of Marine Biology, Far Eastern Branch, Russian Academy of Sciences, Vladivostok, 690041 Russia b Kovalevsky Institute of Biology of the Southern Seas, Russian Academy of Sciences, Sevastopol, 299011 Russia *e-mail: [email protected] Received August 20, 2019; revised December 19, 2019; accepted January 30, 2020

Abstract—This review analyzes the current state of knowledge of the microphytobenthos in the Russian waters of the Sea of Japan and adjacent sea areas. A retrospective analysis of the literature shows that microphytobenthos, with its primary production that often exceeds the production of phytoplankton, plays an important role in coastal marine ecosystems. Benthic diatoms in the coastal waters of Peter the Great Bay (Sea of Japan), which are the primary food object for many marine invertebrates and mammals, are also used as important indicators of the state of the aquatic environment. Despite the great number of publications on the region, knowledge of the microphytobenthos in taxonomic and ecological-floristic aspects is still insufficient. Keywords: microphytobenthos, benthic diatoms, periphyton, epiphyton, Sea of Japan DOI: 10.1134/S1063074020040021

INTRODUCTION The microphytobenthos is a multi-species community of photosynthetic microorganisms that settle on different substrates. Substrates are non-living (pebble, boulders, rocks, and bottom sediments) and living (bottom vegetation and the surface of bodies of mammals and invertebrate animals); moreover, substrates can be any natural and man-made objects that are floating in water or located on the bottom of water bodies [54, 110, 114]. The microphytobenthos is composed mostly of individual diatoms or colonial forms that are capable of forming either macroforms that are visible with the naked eye or microscopic forms of colonies [54, 141]. Diatoms belong to the largest group of unicellular algae according to the number of species, abundance, and prevalence in all biotopes of the World Ocean. To describe the niches in which diatoms occur, Round [120] suggested that one use the terminology accepted for animals and plants. Thus, the following ecotopes are distinguished: epilithon (algae living on the surface of stones), epipelon (algae living on silty sediments), epipsammon (algae living on sands), epiphyton (algae on the surface of bottom vegetation), epizoon (algae on invertebrates and mammals), and periphyton (attached to anthropogenic substrates placed in water by man or that have accidentally fallen into the sea). Diatoms have existed since ancient times. In fossil complexes, diatoms have been known since the Juras-

sic period [107], but in diversity and in a well-preserved state diatoms have occurred since the Cretaceous period [76]. Specialists attribute some findings of diatoms to the Proterozoic Era [126]. Currently, approximately 200000 diatom species belong to allaround microscopic