COMPOSITION AND DISTRIBUTION OF BENTHOS IN THE INTERTIDAL ZONE OF RUSSKY ISLAND (PETER THE GREAT BAY, SEA OF JAPAN) M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo A.V. Zhirmunsky Institute of Marine Biology, FEB RAS, Vladivostok, 690041 e-mail: [email protected] The intertidal zone of Russky Island (Peter the Great Bay, Sea of Japan) is described, and quantitative characteristics of 23 benthic communities, represented by 50 macrophytic and 181 animal species, are given. It is shown that the proportion of pseudomeiobenthos in meiobenthos is greater than that of eumeiobenthos not only with reference to biomass, but also to population density. When comparing species richness of macrobenthos of the intertidal zone of Russky Island with that of the northern part of Amursky Bay (Peter the Great Bay) adjacent to the mouth of the Razdolnaya River, on the one hand, and with that of the islands of the Far Eastern Marine Biosphere Reserve, situated in the open part of Peter the Great Bay, on the other hand, a biodiversity gradient was revealed, that is a regular increase of macrobenthic species number beginning from the freshened areas of the bay and up to the open sea shores. Key words: intertidal zone, macrobenthos, meiobenthos, pseudomeiobenthos, eumeiobenthos, biodiversity gradient. Описана литораль острова Русский (зал. Петра Великого, Японское море) и дана количественная характеристика 23 сообществ бентоса, представленного 50 видами макрофитов и 181 видом животных. Показано, что в мейобентосе доля псевдомейобентоса больше доли эвмейобентоса не только по биомассе, но и по плотности поселения. При сравнении видового богатства макробентоса литорали острова Русский с таковым северной части Амурского залива (зал. Петра Великого), с одной стороны, и островов Дальневосточного государственного природного морского биосферного заповедника, расположенных в открытой части зал. Петра Великого, с другой, выявлен градиент биоразнообразия, то есть закономерное увеличение числа видов макробентоса от опресненных участков залива к открытым морским берегам. Ключевые слова: литораль, макробентос, мейобентос, псевдомейобентос, эвмейобентос, градиент биоразнообразия. Up to now, biota of the intertidal zone of Russky Island in Peter the Great Bay (Sea of Japan) has not been investigated in detail; nevertheless, publications on the bay biota contain a considerable amount of data on this subject (Zaks, 1927; Derjugin, 1939; Shchapova, 1957; Mokievsky, 1960; Perestenko, 1980; Volova, 1985; Gulbin et al., 1987; Kafanov and Zhukov, 1993; Kostina et al., 1996; Kashenko, 1999; Ivanova et al., 2006, 2008; Kepel, 1999, 2004, 2007). 87 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo The height of the intertidal zone ranges from 0.4 to 1.0 m in various sites of Peter the Great Bay and makes about 0.8 m over 0 depths in the water area adjacent to Vladivostok including Russky Island (Tables…, 2006). The lowest and the highest calculated sea levels, theoretically possible for astronomical reasons and referred to seasonal fluctuations, are accepted as the limits of the intertidal zone. Seasonal variations of the sea level in Peter the Great Bay have a yearly period. With their size reaching 0.3 m, these sea level fluctuations significantly affect vertical distribution of benthic organisms. That is why hydrobiological surveys in Peter the Great Bay consider the summer and winter levels of the low tide maximum, conventionally named by some authors as summer and winter «zeros» of depth (Shchapova, 1957; Mokievsky, 1960; Kudryashov, 1980; Perestenko, 1980). I.G. Zaks (1927) and K.M. Derjugin (1939) distinguished two horizons in the intertidal zone of Peter the Great Bay based on the key floral-faunistic assemblages (communities). K.M. Derjugin, who described chiefly the summer condition of biocenoses, did not probably consider seasonal fluctuations of the sea level and overstated the zero depth mark. If we take into account that the sea level lowers in winter, then «Corallina–Laurencia, Chordaria biocenosis», located according to Derjugin’s scheme (Derjugin, 1939) in transitional levels of the intertidal and subtidal zones, is included into the limits of the intertidal zone. Other authors also divide the intertidal zone of the Sea of Japan into two levels (Mokievsky, 1956, 1960; Shchapova, 1956, 1957; Perestenko, 1980; Kafanov and Zhukov, 1993; Kostina et al., 1996; Kashenko, 1999; Kozhenkova and Galysheva, 2006). So, T.F. Shchapova takes into account the annual sea level fluctuations and correctly determines the intertidal zone height for Putyatin Island (Shchapova et al., 1957), but the height of the intertidal zone of Bolshoy Pelis Island and in other areas of Peter the Great Bay are understated (Shchapova, 1957). Difficulties in distinguishing levels and horizons in the intertidal zone of Peter the Great Bay are connected with the fact that tide levels only slightly exceed seasonal fluctuations in sea level. Following the tradition elaborated in the A.V. Zhirmunsky Institute of Marine Biology FEB RAS (hereafter, IMB) during the intertidal surveys in the Far Eastern seas (Kafanov et al., 2004), we divide the intertidal zone of Russky Island into three levels. The scheme of division of the intertidal zone into three horizons was developed by O.G. Kussakin (1961) for the intertidal zone with irregular semidiurnal tides, as in Peter the Great Bay. During our investigations, the lower horizon remained almost submerged, as low tides did not exceed 0.3–0.5 m. The middle and upper horizon were exposed during our surveys. Material and methods Investigation of intertidal zone of Russky Island was carried out by the littoral group of IMB in August and September of 2007. The following areas were investigated: Ajaks Bay, Paris (Zhitkova) Bay, a shallow backwater near Cape 88 Composition and distribution of benthos in the intertidal zone Akhlestysheva, Karpinsky Bay, an area to the east from Cape Ivantsov, Voevoda Bay (Melkovodnaya and Kruglaya Bays), Rynda and Novik Bays in the area of Capes Staritsky, Ekipazhny and Ermolaev (Figs. 1–13). Fifteen hydrobiological transects have been made (12 routine and 3 recognaissance ones), 63 quantitative and more than 40 qualitative macrobenthos samples, and 58 meiobenthos samples (meiobenthos have not been taken at the 1 and 4 transects) have been collected (Fig. 1). The material was collected using the generally accepted methods of chorological research in the intertidal zone (Kussakin et al., 1974). At the study site, a preliminary visual assessment of intertidal communities’ distribution was made; intertidal Fig. 1. A schematic map showing the location of hydrobiological (1–12) and recognaissance (7a, 8a, 9a) transects in the intertidal zone of Russky Island. 89 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo communities were identified on the basis of dominant species of macrobenthos. The names of communities derive from the Latin names of dominant species. Hydrobiological transects were made perpendicular to the shoreline. Sampling plots were bounded by metal frames (100 and 500 cm2). Samples from communities of small evenly distributed organisms were taken with smaller frames than from communities of relatively large species or species with sparse and scattered distribution. In some cases, samples of meiobenthos were collected with a soil cup (19.6 cm2 cross section and 5 cm high). One or two samples were collected for each community. To remove substrate, samples of benthos were washed through soil sieves with a lower sieve of 59 µm mesh. The samples were sorted; organisms of macrobenthos were enumerated, slightly dried on the filter paper, and weighed on pharmaceutical scale with an accuracy of 10 mg. Meiobenthos was stained with Rose Bengal, sorted, and enumerated under a binocular in Bogorov’s chamber. Biomass of meiobenthic animals was determined using the data of Galtsova and Pavlyuk (1987) calculated from Chislenko’s nomograms (1968). The qualitative data for macro- and meiobenthos were extrapolated to 1 m2. Biomasses of organisms including shells and other skeletal formations are expressed as wet weight values. The collections were fixed with 75% alcohol and 4% formalin. Taxonomy of species was taken from the work Kussakin et al. (1997). In the period of the study, water temperature varied during low tides from 20ºC in Ajaks Bay to 25.5ºC in Melkovodnaya Bay (Voevoda Bay). Water salinity was up to 30‰ in Karpinsky Bay and to 25‰ in Novik and Voevoda Bays. High salinity (35‰) was recorded only in a shallow lagoon at Cape Akhlestysheva. Results and discussion In the intertidal zone of the study area, we have found and identified 50 plant species (32 Rhodophyta, 5 Chlorophyta, 11 Phaeophyceae, and 2 Magnoliophyta) and 181 animal species (32 Gastropoda, 24 Bivalvia, 1 Polyplacophora, 24 Polychaeta, 4 Cirripedia, 34 Amphipoda, 12 Isopoda, 10 Decapoda, 31 Nematoda, 5 Echinodermata, 1 Actiniaria, 1 Nemertea, and 2 Pisces), including 130 species of macrobenthic animals and 51 meiobenthic organisms (Supplement, Table 1). Meiobenthos included Nematoda, Foraminifera, Ostracoda, Copepoda (Harpacticoida, Calanoida), Acarina, Turbellaria, young and adult forms of Polychaeta and Oligochaeta, as well as juvenile individuals of Bivalvia (11 species of them were found only in meiobenthos), Gastropoda (3), Isopoda (4), Amphipoda, Ophiuroidea, and Insecta larvae. Eumeiobenthos was composed of the following groups: Nematoda, Foraminifera, Ostracoda, Copepoda, Acarina, and Turbellaria. Other taxonomic groups of meiobenthos were assigned to pseudomeiobenthos. In the intertidal zone of Russky Island, the highest species diversity of macrobenthos (80 plant and animal species) was found in Stark Strait on the stonyrubble substrate, east of Cape Ivantsov. A somewhat lower diversity (68 species) 90 Composition and distribution of benthos in the intertidal zone was recorded for the middle part of Ajaks Bay on the pebble bottom, and the rocky-rubble intertidal zone in Karpinsky Bay was the third in biodiversity (63 species). The lowest species diversity was observed in a shallow creek near Cape Akhlestysheva in the silted sandy drying site with single stones (22 species), in Novik Bay near Cape Ekipazhny on the crushed stone bottom (20 species), and near Cape Ermolaev on the sand-pebble bottom (19 species). In other investigated areas, the number of recorded plants and animals varied from 25 to 52 species (Supplement, Table 1). A comparative study of species richness of macrobenthos in the intertidal zone of Russky Island, in the northern part of Amursky Bay (Peter the Great Bay) adjacent to the mouth of the Razdolnaya River, and on the islands of the Far Eastern Marine Biosphere Reserve, situated in the open part of Peter the Great Bay, revealed a natural increase of the number of macrobenthic species from freshened areas of the bay towards the open sea coasts. The materials were collected in summer in the indicated three areas of Peter the Great Bay with the use of the same technique, thus the obtained data is quite comparable (see Table), except for the number of algal and sea grass species in the marine reserve. This value for the islands of the Far East Marine Biosphere Reserve is slightly overstated, as it involves the seasonal dynamics, and is calculated for the entire reserve area. Earlier, when studying macrobenthos of the intertidal zone of Shikotan Island (the Lesser Kuril Ridge), we recorded a similar regularity in macrobenthos distribution on the example of 226 species from 16 habitats, and gave it a conventional name «biodiversity gradient» (Ivanova and Tsurpalo, 2007). «Biodiversity gradient» is not a universal feature of biota but typical of macrobenthos; the gradient reflects regularities of macrobenthos distribution, connected with the salinity gradient and the degree of wave activity. An inverse regularity was observed for Biodiversity gradient of macrobenthos in the intertidal zone of Peter the Great Bay Taxon Northern part of Amursky Bay Russky Island Islands of the Far East Marine Biosphere Reserve Crustacea (Cirripedia, Decapoda, Amphipoda, Isopoda) 33 56 85 Mollusca (Gastropoda, Bivalvia, Polyplacophora, Cephalopoda) 21 43 63 Polychaeta 6 22 37 Echinodermata 1 5 9 Actiniaria 1 1 3 32 50 173 Plants (Rhodophyta, Phaeophyceae, Chlorophyta, Magnoliophyta) 91 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo those meiobenthos groups, for which substrate properties and, especially, the presence of fine fractions in bottom sediments are more important for distribution. For example, nematode species were more numerous in the intertidal zone of the northern part of Amursky Bay (45 species) than in the intertidal zone of Russky Island (31 species). On the whole, in the intertidal zone of Russky Island characterized by various substrates, a fairly large number of macrobenthic communities developed, among which 23 plant and animal species dominated by biomass. But every specific habitat included a limited number of communities. The highest number of communities (8 and 7) was registered, respectively, in Stark Strait east from Cape Ivantsov and in Karpinsky Bay. The communities of the barnacle Chthamalus dalli were of most frequent occurrence within the entire vertical range of the intertidal zone (averaged data are presented in Supplement, Table 2). Biomass of the dominant species varied from 140 to 1140 g/m2 in the upper horizon, from 100.7 to 750 g/m2 in the middle horizon, and from 985 to 1270 g/m2 in the lower horizon. The dominant species contributing sometimes nearly 100% of the total biomass in the community reduced in some cases to 56.2%. In a modification of Chthamalus dalli+Littorina mandshurica community, Ch. dalli biomass reduced to 48% of the total value, while subdominant species accounted for 45.8% of the total biomass. Chthamalus dalli community was encountered on rocky, boulder, and stony substrates and on single stones, lying on silted sands (Melkovodnaya Bay). In the upper intertidal horizon, this community included few species, among them gastropods Littorina brevicula, L. mandshurica, Lottia kogamogai, and Lottia versicolor, isopods Dynoides dentisinus, and Cyanobacteria. In the middle horizon, the dominant species was accompanied by a greater number of species: the red algae Gloiopeltis furcata, Masudaphycus irregularis, Ceramium kondoi, and Polysiphonia japonica and different animals. Most frequent among them were L. brevicula and other gastropods (Littorina squalida, L. mandshurica, Falsicingula athera, Lottia tenuisculpta, Epheria turrita and others), crustaceans (Caprella bispinosa, D. dentisinus, Pagurus brachiomastus, Amphibalanus improvisus, Hyale sp. and others). In the lower horizon, the accompanying species included the red calcareous alga Corallina pilulifera, gastropods (Littorina sitkana, L. mandshurica, L. squalida, F. athera, Hima acutidentata, L. kogamogai, L. tenuisculpta), and the barnacle Balanus rostratus. Littorina brevicula community was very frequent in the intertidal zone of Russky Island (averaged data are presented in Supplement, Table 3). As well as the community given above, it occurred within the entire vertical range of the zone. In the upper intertidal horizon, biomass of L. brevicula in some aggregations in rock and stone hollows reached from 150.2 to 7189 g/m2; a lower biomass of 333 to 1984 g/m2 was registered in the middle horizon, and the lowest value of 278 g/m2 92 Composition and distribution of benthos in the intertidal zone was estimated for the lower horizon. Littorina brevicula as a dominant species was responsible for 59.9 to almost 100% of the total biomass of the community. Littorina brevicula community occupied rocks, boulders, blocks, and big pebbles among stones. In the upper intertidal horizon, this community included the alga Gloiopeltis furcata and the animal species Ch. dalli and Falsicingula athera. In the middle horizon, the dominant species was accompanied by the species the listed above and crustaceans (Amphibalanus improvisus, Hyale bassargini, Hemigrapsus sanguineus, Hemigrapsus penicillatus, Gnorimosphaeroma rayi), the limpet Lottia tenuisculpta, and polychaetes (Nereis vexillosa, Nereis multignatha, Capitella capitata and Typosyllis pulchra occidentalis). In the lower horizon, sponges and Ch. dalli were the accompanying species. A community of the red calcareous alga Corallina pilulifera was also very typical of the intertidal zone of Russky Island (averaged data are presented in Supplement, Table 4), the alga was of frequent occurrence in the lower intertidal horizon. The biomass of the dominant species ranged from 270 to 894 g/m2 and made up from 48.4 to 78.1% of the total community biomass. Corallina pilulifera community inhabited rocks, boulders and stony substrates. Various animals and some algae were included into the community as accompanying species: Scytosiphon lomentaria, Dictyota dichotoma, Analipus japonicus and numerous small gastropods (Falsicingula athera, Pusilina plicosa, Ansola angulosa and Ansola angustata, Epheria turrita, juvenile cirripeds, L. brevicula, L. squalida, Homalopoma sangarense, Lucidestea ornatа). Bivalves Turtonia minuta, juvenile Crenomytilus grayanus, Mytilus trossulus kussakini, Venerupis philippinarum occurred less frequently. Besides, polychaetes Nereis multignatha, N. zonata tigrina, Platynereis bicanaliculata, crustaceans Ampithoe zachsi, Ampithoe sp., Hyale sp., Jassa marmorata, Guernea quadrispinosa, as well as Dynoides dentisinus, Holotelson tuberculatus, and Ch. dalli were also registered in the community. In addition, communities dominated by gastropods (Littorina mandshurica, L. squalida, Lottia kogamogai, and Batillaria cumingii) were also recorded as typical of the intertidal zone of Russky Island. In Littorina mandshurica communities, biomass of the dominant species was up to 1467 g/m2 on boulders in the upper intertidal horizon and 463 g/m2 on pebblesandy grounds in the lower horizon. So, the dominant species was responsible for almost 100% of the total community biomass on boulders and up to 68% on pebblesandy ground. Among numerous accompanying species with a substantial biomass, Ch. dalli was recorded in the upper horizon and C. pilulifera, L. squalida, N. vexillosa, and Ch. dalli were common in the lower horizon (Supplement, Table 5). Littorina squalida community was common in the middle intertidal horizon on pebble-sandy silted ground and in the upper part of the lower horizon on stones. L. squalida biomass reached 1495 g/m2 and made up to 58.7% of the total community biomass. In the middle horizon, the red algae Gracilaria gracilis, crustaceans 93 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Eogammarus possjeticus and Pagurus hirsutiusculus, as well as the polychaete Heteromastus filiformis were the accompanying species. In the lower horizon, algae Ralfsia fungiformis, Bossiella compressa, Clathromorphum compactum were attributed to accompanying species. The following animal species inhabited the community: crustaceans Hemigrapsus sanguineus, Caprella sp., and Melita sp., mollusks Venerupis philippinarum, Homalopoma sangarense, Tegula rustica, and Lottia tenuisculpta, and polychaetes N. vexillosa and Harmothoe imbricatа (Supplement, Table 6). In the upper intertidal horizon on boulders and blocks in Karpinsky Bay, a community of the limpet Lottia kogamogai with biomass of the dominant species of 180 g/m2 contributed 63% of the total community biomass. Chthamalus dalli, L. sitkana, Falsicingula athera, and Dynoides dentisinus were accompanying species (Supplement, Table 7). Gastropod Batillaria cumingii colonies were typical of pebble-sandy silted grounds in the middle and lower intertidal horizons. Biomass of the dominant species in this community reached 109 g/m2 and made up nearly 100% of the total community biomass. The only accompanying species here was Nereis sp. juv. The average biomass of B. cumingii made 65.37 g/m2 at population density of 867 ind./m2. In Macoma contabulata+Hima fratercula community, mollusks formed less abundant accumulations. Their biomasses reached 53 and 47 g/m2, 41.6 and 36.9% of the total biomass, respectively. This community occupied silted sand with pebble admixture in the lower intertidal horizon in a shallow creek to the southwest of Cape Ahlestishev. The red algae Polysiphonia japonica and Gracilaria gracilis, as well as amphipods Pontogeneia intermedia were the accompanying species (Supplement, Table 8). Among communities dominated by animals, Hemigrapsus penicillatus+Gnorimosphaeroma rayi community is worth noting. These crustaceans were typical of pebble grounds in the middle horizon, their biomass reached 38.7 and 26.3 g/m2, 33.5 and 22.8% of the total biomass, respectively. Gastropods Epheria turrita, Littorina brevicula, L. squalida, Cecina manchurica, Pusilina plicosa, and Lucidestea ornata, amphipods Pontogeneia sp., and polychaetes Neanthes sp. were of frequent occurrence as accompanying animals (Supplement, Table 9). Among red algae, inhabiting the Russky Island intertidal zone, in addition to the above-mentioned Corallina pilulifera, the following species formed communities: Grateloupia turuturu, Neorhodomela larix aculeata, N. munita, Gracilaria gracilis, Lomentaria hakodatensis and two species of the genus Ceramium in a joint C. japonicum+C. kondoi community. Communities of the red algae Gloiopeltis furcata and Campylaephora crassa inhabited the middle horizon. The highest biomass for red algae, 4200 g/m2, was registered for Grateloupia turuturu that contributed 99% of the total community biomass. This species was accompanied by the epiphyte Polysiphonia japonica and polychaetes Spirorbidae (Supplement, Table 10). 94 Composition and distribution of benthos in the intertidal zone Neorhodomela larix aculeata had a lower biomass not over 3300 g/m2, 87.9% of the total community biomass. This species was accompanied by the mollusks Mytilus trossulus kussakini, Ansola angustata, Falsicingula athera, Epheria turrita, Alaba picta, Littorina squalida, and Pusilina plicosa. Falsicingula athera was the most abundant among them, its population density reached 132800 ind./m2 with 344 g/m2 biomass. The dominant species was accompanied by algae C. pilulifera, Ceramium japonicum, and Gracilaria gracilis. Animals were represented by numerous amphipods (Caprella scaura diceros, C. bispinosa, Ampithoe djakonovi, A. lacertosa, Jassa marmorata and others); hermit crab Pagurus middendorffii, isopod Holotelson tuberculatus, and polychaete Nereis multignatha also occurred (Supplement, Table 11). The community of another species of the genus Neorhodomela, N. munita, formed patches in the C. pilulifera zone. The dominant species biomass in this community reached 1650 g/m2 (73.3% of the total biomass). This species was also accompanied by numerous and various amphipods (Caprella mutica, Pontogeneia intermedia, J. marmorata, Guernea quadrispinosa, Ampithoe sp., A. lacertosa, Ischirocerus sp. and others), hermit crab Pagurus middendorffii, isopod Holotelson tuberculatus, bivalves (Turtonia minuta and M. trossulus kussakini), and numerous gastropods (F. athera, Homalopoma sangarense, Lucidestea ornata, Ansola angulosa, Mitrella burchardi, Pusilina plicosa, and L. squalida) (Supplement, Table 12). Another red alga, Gracilaria gracilis, followed to form a community in the lower intertidal horizon, it attached to small stones on silted sands in Melkovodnaya Bay (Voevoda Bay). The dominant species gave a moderate biomass of 330 g/m2, i.e. 47.2% of the total biomass of the community. The main accompanying species were here mollusks: Batillaria cumingii with 213 g/m2 biomass (30.5%) and Littorina mandshurica (151 g/m2). Amphipods (Eogammarus possjeticus and Monocorophium acherusicum) also occurred in this community. Besides, the epiphyte Polysiphonia japonica and diatoms are also worth mentioning (Supplement, Table 13). Lomentaria hakodatensis formed a community on the upper surface of boulders with a small biomass of the dominant species (260 g/m2, 75.3% of the total biomass). The gastropod F. athera, the amphipods J. marmorata, Ampithoe sp., Ischirocerus sp., and Hyale sp. and the brown alga Dictyota dichotoma were registered here as accompanying species (Supplement, Table 14). The last red alga community in the lower intertidal horizon was formed by Ceramium japonicum and C. kondoi. These species did not develop great biomass, only 94 and 64 g/m2 (30.1 and 20.5% of the total community biomass), respectively. These species were accompanied by other algae (Laurencia nipponica, N. larix aculeata, C. pilulifera, P. japonica, D. dichotoma) and animals (amphipods J. marmorata, P. intermedia, and Ampithoe sp., mollusks F. athera and Lottia tenuisculpta, and the barnacle Ch. dalli) (Supplement, Table 15). 95 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Gloiopeltis furcata community was typical of the middle intertidal horizon, especially in its upper part. This species formed a moderate biomass not over 195 g/m2 (72.2% of the total biomass) on stone blocks and no less than 84.5 g/m2 (65.8% of the total biomass) on boulders. The main accompanying species was Ch. dalli. The isopod Holotelson tuberculatus and the mollusks Tritonalia japonica and L. squalida were also found (Supplement, Table 16). Besides the G. furcata community, the red alga Campylaephora crassa developed communities on block surfaces in the middle horizon. The dominant species made biomass of 620 g/m2 (79.9% of the total biomass). The main accompanying species in this community were Ch. dalli (up to 110 g/m2), amphipods (J. marmorata, Caprella cristibrachium, Ischirocerus sp., Ampithoe sp., Pontogeneia sp.), the isopod Dynoides dentisinus, and gastropods (F. athera, Ansola angustata, and Mitrella burchardi) (Supplement, Table 17). Communities of brown algae were also recorded in the lower and middle horizons of the Russky Island intertidal zone. The highest biomass among brown algae was formed in the lower horizon by Sargassum pallidum (3800 g/m2, 90.6% of the total community biomass). This dominant species was accompanied by the alga Neorhodomela munita with epiphytes Lithophyllum tumidulum, Polysiphonia morrowii, Gelidium vagum, Sphacelaria furcigera, and Cladophora stimpsonii. Besides, algae (Ulva fenestrata, Scytosiphon lomentaria, Grateloupia divaricata, and Ahnfeltia tobuchiensis) and animals (amphipods Ampithoe sp., A. annenkovae, Caprella penantis, C. algaceus, J. marmorata, Platorchestia pachipus, Orchomenella sp., mollusks Turtonia minuta, L. squalida, M. burchardi, F. athera, Lucidestea ornata, Alaba picta, polychaetes Nereis multignatha, and hermit crab Pagurus middendorffii) were also recorded here (Supplement, Table 18). Pebble bottoms in the lower intertidal horizon were occupied by a Coccophora langsdorfii community, biomass of the dominant species made up 240 g/m2 (50.6% of the total biomass). The accompanying species in this community were mollusks (F. athera, L. brevicula, Hima fratercula, Lottia tenuisculpta, L. squalida, Epheria turrita, and M. trossulus kussakini), amphipods (Caprella laevis, Ericthonius tolli, Ampithoe sp.), and the epiphyte Polysiphonia japonica (Supplement, Table 19). Of brown algae, Chordaria flagelliformis and Dictyota dichotoma communities were the most typical of the middle intertidal horizon. Biomass of Ch. flagelliformis reached 2030 g/m2 (99.3% of the total community biomass). The accompanying species in Ch. flagelliformis community were mollusks (F. athera, L. squalida, Pusilina plicosa, Lucidestea ornata, Lirularia iridescens, M. burchardi, Ansola angustata, and Turtonia minuta), isopod Dynoides dentisinus, and amphipods (Caprella penantis, C. laevis, J. marmorata, Calliopius laeviusculus, Paracalliopiella litoralis, Parallorchestes zibellina and Ampithoe sp.) (averaged data see in Supplement, Table 20). 96 Composition and distribution of benthos in the intertidal zone Dictyota dichotoma formed a community on rocks, its biomass was up to 465 g/m2 (39.8% of the total community biomass). Numerous algae (N. larix aculeata, Ch. flagelliformis) with epiphytes (Saundersella simplex, C. pilulifera, Laurencia pinnata), mollusks (F. athera and E. turrita), and amphipods (Ampithoe djakonovi and Hyale sp.) accompanied the dominant species in this community (Supplement, Table 21). Higher plants in the intertidal zone of Russky Island were presented by Phyllospadix iwatensis and Zostera marina communities. These two species inhabited the lower intertidal horizon and the upper subtidal zone. Community of Ph. iwatensis occupied rocks and boulders. Biomass of the dominant species reached 8280 g/m2 (62.5% of the total community biomass). Numerous gastropods (Homalopoma sangarense, Ansola angulosa, Alaba picta, M. burchardi, Pusilina plicosa, L. ornata, Littorina squalida, Lirularia iridescens, F. athera), bivalve T. minuta, and amphipods (Ampithoe sp., Caprella bispinosa, C. penantis, J. marmorata, Paradexamine fraudatrix, and Lysianassidae) were recorded as accompanying species (Supplement, Table 22). Community of the eelgrass Zostera marina developed on soft silted grounds. Biomass of the dominant species was not great and reached 1290 g/m2 (71.7% of the total community biomass). The crustacean Hemigrapsus sanguineus and gastropods B. cumingii and L. squalida were registered as accompanying species (Supplement, Table 23). The zone of the eelgrass Z. marina washed ashore during storms is a special biotope with a peculiar distribution of the biota. This zone supports the highest population density and biomass of meiobenthos in the intertidal zone of Russky Island. The highest pseudomeiobenthos biomass (more than 41 g/m2), contributed mainly by the gastropod mollusks Fartulum bucerius and Brochina derjugini, was recorded for the upper intertidal horizon on sandy ground in the zone of the eelgrass Z. marina cast onto the beach in Paris Bay. The highest eumeiobenthos biomass (more than 2.7 g/m2), formed by abundant Foraminifera, was also registered in the same place. In addition to the mentioned Gastropoda species, various isopods, oligochaetes and other small animals found their shelter under Zostera marina deposits on sandy-pebble beach and formed a considerable for pseudomeiobenthos biomass of about 4 g/m2 in Ajaks Bay and about 3 g/m2 in Rynda Bay (Supplement, Table 24). Especially numerous macrobenthic forms in the zone of Zostera marina washed ashore were the amphipods Platorchestia pachypus and Eogammarus possjeticus, the isopod Gnorimosphaeroma rayi, and Oligochaeta. The polychaete Lycastopsis pontica, gastropod mollusks Epheria turrita, Littorina brevicula, and L. mandshurica, crabs Hemigrapsus sanguineus and H. penicillatus and other animals occurred also there. In general, high values of meiobenthos biomass are not typical of communities, distinguished by dominating species of macrobenthic animals and plants. However, values greater than 1 g/m2 were registered in the communities of Neorhodomela 97 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo larix aculeata (1.80 g/m2), Campylaephora crassa (1.780 g/m2), Littorina squalida (1.303 g/m2), L. mandshurica (1.202 g/m2), L. brevicula (1.077 g/m2), Macoma contabulata+Hima fratercula (1.062 g/m2), and Gloiopeltis furcata (1.051 g/m2). Meiobenthos biomass values close to 1 g/m2 were recorded for the communities of Hemigrapsus penicillatus+Gnorimosphaeroma rayi (0.987 g/m2) and Zostera marina (0.820 g/m2). Some typical features of distribution of pseudomeiobenthos systematic groups were seen in these examples of a relatively high meiobenthic biomass in the communities. Thus, thick growths of Neorhodomela larix aculeata in the inshore block intertidal zone served as a good shelter for Amphipoda juveniles, which formed the bulk of pseudomeiobenthos biomass in this community (1.14 g/m2 of Gammaridea and 0.502 g/m2 of Caprellidea; Supplement, Table 11). In pseudomeiobenthos, Oligochaeta prevailed on the pebble-sandy, sandy and pebble silted grounds in the communities of Littorina squalida (1.1016 g/m2, Supplement, Table 6, the middle horizon), Macoma contabulata+Hima fratercula (0.8262 g/m2, Supplement, Table 8), and Hemigrapsus penicillatus+Gnorimosphaeroma rayi (0.459 g/m2, Supplement, Table 9). On muddy bottoms, in Zostera marina community, Oligochaeta occupied the second place in pseudomeiobenthos biomass (0.1836 g/m2), Polychaeta being the first (0.4131 g/m2, Supplement, Table 23). In a community of Littorina mandshurica formed on the pebble-sandy bottom, Oligochaeta predominated (0.6426 g/m2), and Gastropoda (0.1071 g/m2), represented by Fartulum bucerius, Brochina derjugini, and Epheria sp. species, were the second in the pseudomeiobenthos biomass (Supplement, Table 5). Gastropoda prevailed by biomass in pseudomeiobenthos on rocks, blocks and stones under various conditions. Represented by F. bucerius, B. derjugini, and Pusilina plicosa, Gastropoda were a dominant group in pseudomeiobenthos of the communities of Campylaephora crassa (0.784 g/m2, Supplement, Table 17), Gloiopeltis furcata (0.385 g/m2, Supplement, Table 16), and Littorina brevicula (0.3801 g/m2, Supplement, Table 3, the middle horizon). Communities with meiobenthos biomass below 1 g/m2 revealed practically the same regularities in Amphipoda and Oligochaeta distribution. So, numerous Amphipoda juveniles that took shelter on rocks, rubbles and big stones among thick algal beds prevailed by biomass in pseudomeiobenthos in the communities of Lomentaria hakodatensis (0.408 g/m2, Supplement, Table 14), Dictyota dichotoma (0.36 g/m2, Supplement, Table 21), Corallina pilulifera (0.288 g/m2, Supplement, Table 4), Ceramium japonicum+C. kondoi (0.157 g/m2, Supplement, Table 15), and Grateloupia turuturu (0.104 g/m2, Supplement, Table 10). Oligochaeta prevailed on mudsandy bottom with small stones in pseudomeiobenthos of Gracilaria gracilis community (0.1836 g/m2, Supplement, Table 13). Gastropoda, mainly owing to Ansola angustata and Odostomia culta species, dominated by biomass in pseudomeiobenthos on pebble grounds in a Coccophora langsdorffii community (0.203 g/m2 at the total meiobenthos biomass of 0.52051 g/m2, Supplement, Table 19). 98 Composition and distribution of benthos in the intertidal zone In the community of the barnacle Chthamalus dalli, the meiobenthos biomass varied from 0.1228 to 0.34625 g/m2 (Supplement, Table 2). Juveniles of the dominant species, designated as Balanomorpha (juv.), prevailed in this community in pseudomeiobenthos of the upper intertidal horizon (0.0672 g/m2). In the middle horizon, Balanomorpha (juv.) occupied the second place by biomass (0.0287 g/m2), Gastropoda (0.0392 g/m2) were the first. In the lower horizon, Balanomorpha (juv.) receded to the third place (0.042 g/m2); Polychaeta (0.108 g/m2) and Gastropoda (0.077 g/m2) were on the first and the second places, respectively. The lowest meiobenthos biomass (0.007622 g/m2) was registered in a Lottia kogamogai community in the upper horizon of the rubble-block intertidal zone (Supplement, Table 7). Polychaeta (0.018 g/m2), Gastropoda (0.014 g/m2), Balanomorpha (juv.) (0.014 g/m2), Isopoda (0.014 g/m2), and Bivalvia (0.014 g/m2) were presented in pseudomeiobenthos of this community almost in equal amounts. In the intertidal zone, biomass of pseudomeiobenthos exceeded as a rule that of eumeiobenthos (Ivanova et al., 2006). The percentages of pseudomeiobenthos and eumeiobenthos (mean biomass) in the intertidal zone of the Russky Island are shown in Fig. 14. With rare exceptions, Foraminifera prevailed in eumeiobenthos of the intertidal zone of Russky Island, when the total eumeiobenthos biomass reached 0.013–0.35 g/m2. So, Foraminifera dominated in the following communities: Littorina mandshurica (0.19125 g/m2, Supplement, Table 5), Macoma contabulata+Hima fratercula (0.19125 g/m2, Supplement, Table 8), Lomentaria hakodatensis (0.1825 g/m2, Supplement, Table 14), Gracilaria gracilis (0.1785 g/m2, Supplement, Table 13), Littorina brevicula (0.168937 g/m2 in the middle horizon, Supplement, Table 3), Zostera marina (0.11475 g/m2, Supplement, Table 23), Littorina squalida (0.08925 g/m2 in the middle horizon and 0.006 g/m2 in the lower horizon, Supplement, Table 6), Gloiopeltis furcata (0.105 g/m2, Supplement, Table 16), Campylaephora crassa (0.055 g/m2, Supplement, Table 17), Coccophora langsdorffii (0.0525 g/m2, Supplement, Table 19), Chthamalus dalli (0.02375 g/m2 in the lower horizon, Supplement, Table 2), Corallina pilulifera (0.033333 g/m2, Supplement, Table 4), and Ceramium japonicum+C. kondoi (0.0135 g/m2, Supplement, Table 15). Calanoida prevailed in eumeiobenthos of the communities of Hemigrapsus penicillatus+Gnorimosphaeroma rayi (0.1224 g/m2) and Dictyota dichotoma (0.01 g/m2), where the total biomass of meiobenthos was equal to 0.15433 g/m2 and 0.01268 g/m2, respectively (Supplement, Tables 9 and 21, respectively). At a lower eumeiobenthos biomass (0.00134–0.00594 g/m2), Turbellaria and Harpacticoida were found to prevail in the communities. Thus, Turbellaria prevailed by biomass in eumeiobenthos of a Neorhodomela larix aculeata community (0.00228 g/m2, Supplement, Table 11), whereas eumeiobenthos of Lottia kogamogai community (Supplement, Table 7) and Grateloupia turuturu community (Supplement, Table 10) developed on rocks was dominated by Harpacticoida (0.00112 g/m2 and 0.00096 g/m2, respectively). 99 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo When eumeiobenthic biomass was minimal (0.000615–0.001 g/m2), Nematoda were the dominant group, as, for example, in Littorina brevicula community in the upper and lower intertidal horizons (0.0002 g/m2 and 0.001 g/m2, respectively, Supplement, Table 3). Ostracoda and Acarina were not registered as biomass dominants in eumeiobenthos of the described communities in the Russky Island intertidal zone. The percentage relationships of meiobenthic groups by their mean biomasses are presented in Fig. 15. The population density of the entire meiobenthos varied during our study from 1900 ind./m2 in Lottia kogamogai community to 1201050 ind./m2 in the zone of Zostera marina cast ashore during storm events. Population density of pseudomeiobenthos made up from 980 ind./m2 in a Littorina squalida community to 605880 ind./m2 in the aggregations of Zostera marina cast onto the beach by storms. The population density of eumeiobenthos varied from 720 ind./m2 in a Chthamalus dalli community to 595170 ind./m2 in the zone of Z. marina cast ashore. The percentages of mean population density of pseudomeiobenthos and eumeiobenthos are shown in Fig. 16. Nematoda more often occupied the first place by the population density (up to 497250 ind./m2), sometimes Foraminifera were recorded as the first, and Harpacticoida dominated in the absence of Nematoda and Foraminifera. Gastropoda and Amphipoda dominated in pseudomeiobenthos. The percentage relationship of the meiobenthic groups by their mean densities is presented in Fig. 17. Thus, although the intertidal zone of Russky Island does not extend high, it supports a diverse and abundant biota. On the other hand, as compared to other areas of the Far Eastern seas, especially such as Kuril and Commander Islands, the intertidal zone of Russky Island is poorly populated. Ajaks and Paris Bays, open to Bosfor Vostochny Strait, are subject to considerable anthropogenic pollution caused by waste waters of Vladivostok. The contamination affects the pattern of vegetation in Ajaks Bay, where the green algae Ulva and Codium prevail in the upper subtidal zone (down to 2–3 m deep) and in the subtidal fringe instead of brown and red algae that are usually common in similar habitats but are sensitive to pollution. In Zhitkova Cove (Paris Bay), the beach in the upper supra-intertidal zone (splash zone) above the belt of sea grass masses cast ashore is covered by plastic litter (bottles, bags and other plastic wastes) (Fig. 13). Nevertheless, there was no oil film within the intertidal zone of the island during our survey. Acknowledgements The authors appreciate the help of many specialists, mostly the workers of IMB, in identification of animals and plants collected during the survey; sincere thanks are due to I.R. Levenets (macrophytes), E.E. Kostina (sea anemones), I.L. Davydkova (polychaetes), I.I. Ovsyannikova (cirripedes), L.L. Budnikova (amphipods; TINRO-Centre), O.A. Golovan (isopods), A.V. Chernyshev (gastropods), N.K. Kolotuchina (bivalves), and A.A. Balanov (fishes). The authors 100 Composition and distribution of benthos in the intertidal zone identified the following animal groups: nematodes (L.S. Belogurova), bivalves and chitons (M.B. Ivanova), echinoderms (A.P. Tsurpalo and M.B. Ivanova), and decapods and nemertines (A.P. Tsurpalo). The study was partly supported by the Asia-Pacific Network for Global Change Research (Project ARCP 2007-12 NMY). References Chislenko, L.I., Nomograms for Determination of Weight of Aquatic Organisms Based on Body Dimensions and Form (Marine Mesobenthos and Plankton), Leningrad: Nauka, 1968, 106 p. Derjugin, K.M., Zonen und Biocönosen der Bucht Peter des Grossen (Japanisches Meer), Volume in Honour of Scientific Activity of N.M. Knipovich (1885–1939). Moscow, 1939, pp. 115–142. 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Zaks, I.G., Preliminary data on fauna and flora distribution in the coastal zone of Peter the Great Bay, Sea of Japan, Proceedings of the 1st Conference on Investigation of Productive Forces of the Far East, Vladivostok, 1927, vol. 4, pp. 213–247. Supplement Table 1 The list of plants and animals of the Russky Island intertidal zone Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos 5 5 Imperium Cellulata Dominium Procariota (Bacteria) Regnum Oxyphotobacteriobiontes Phylum Cyanobacteria (=Cyanophyta) 20.0 Dominium Eukaryota Subdominium Metakaryota Regnum Rhodobiontes Phylum Rhodophyta Classis Florideophyceae 102 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos + 2 – Ordo Hildenbrandiales Fam. Hildenbrandiaceae Hildenbrandia rubra (Sommerf., 1826) Meneghini, 1841 Ordo Corallinales Fam. Corallinaceae Bossiella compressa Kloczcova, 1978 4.2 2, 3 – B. cretacea (Postels et Ruprecht, 1840) Johansen, 1969 + 2 – Clathromorphum compactum (Kjellman, 1883) Foslie, 1898 + 2, 3 – Corallina pilulifera Postels et Ruprecht, 1840 10–894.0 1, 3, 6–12 – Lithophyllum tumidulum Foslie, 1901 10.0 1–3, 12 – Lithothamnion phymatodeum (Foslie, 1902) Foslie, 1929 + 1, 3, 8, 9 – Pneophyllum lejolisii (Rosanoff, 1866) Chamberlain, 1983 + 4, 6, 7, 9, 10 – + 8 – + 9 – + 1 – 6.9–37.0 1 – 8.5 1, 2, 10 – 3.2–330.0 5, 6, 8, 8а, 10 – Ordo Acrochaetiales Fam. Acrochaetiaceae Colaconema daviesii (Dillwyn, 1802–1809) Stegenga, 1985 Fam. Rhodophysemataceae Rhodophysema elegans (Crouan et Crouan, 1867) Dixon, 1964 Ordo Palmariales Fam. Palmariaceae Palmaria stenogona (Perestenko, 1973) Perestenko, 1980 Ordo Ahnfeltiales Fam. Ahnfeltiaceae Ahnfeltia tobuchiensis (Kanno et Matsubara, 1932) Makienko, 1970 Ordo Gelidiales Fam. Gelidiaceae Gelidium vagum Okamura, 1934 Ordo Gracilariales Fam. Gracilariaceae Gracilaria gracilis (Stackhouse, 1802) M. Steentoft, L.M. Irvine et W.F. Farnham, 1995 103 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos 340.0 7 – 0.4–195.0 1, 7, 11, 12 + 1 – 9.0 1, 12 – 4200.0 2, 3, 12 – 260.0 12 – Ceramium japonicum Okamura, 1896 10.5–94.0 10 – C. kondoi Yendo, 1920 4.3–64.0 2, 3, 5, 7, 9–12 – 620.0 12 – + 10 – + 6, 7, 7а, 9 – Symphyocladia latiuscula (Harvey ex Gray, 1857) Yamada, 1941 + 3 – Polysiphonia japonica Harvey ex Perry, 1856 0.2–20.0 1–3, 5, 7, 8а, 9–12 – 12.0 1 – Neorhodomela larix (Turner, 1819) Masuda, 1982 subsp. aculeata Perestenko, 1984 10.0–3300.0 2, 3, 10, 12 – N. munita (Perestenko, 1980) Perestenko, 1994 120.0–3450.0 1, 7, 9 – 0.9–44.0 9–12 – 1.0 12 – Ordo Gigartinales Fam. Dumontiaceae Masudaphycus irregularis (Yamada, 1933) Lindstrom, 1988 Fam. Endocladiaceae Gloiopeltis furcata (Postels et Ruprecht, 1840) J. Agardh, 1876 – Fam. Gigartinaceae Ahnfeltiopsis flabelliformis (Harvey ex Gray, 1856) Masuda, 1993 Ordo Halymeniales Fam. Halymeniaceae Grateloupia divaricata Okamura, 1895 G. turuturu Yamada, 1941 Ordo Rhodymeniales Fam. Lomentariaceae Lomentaria hakodatensis Yendo, 1920 Ordo Ceramiales Fam. Ceramiaceae Campylaephora crassa (Okamura, 1930) Nakamura, 1965 C. hypnaeoides J. Agardh, 1851 Fam. Dasyaceae Dasya sessilis Yamada, 1928 Fam. Rhodomelaceae P. morrowii Harvey ex Gray, 1856 Laurencia nipponica Yamada, 1931 L. pinnata Yamada, 1931 104 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos Regnum Chromobiontes Phylum Ochrophyta Classis Phaeophyceae Ordo Ectocarpales Fam. Chordariaceae Chordaria flagelliformis (Mueller ex Flora Danica, 1775) C. Agardh, 1817 90.0–2030.0 1, 7, 7а, 12 – Saundersella simplex (Saunders, 1901) Kylin, 1940 360.0 12 – Colpomenia peregrina (Sauvageau, 1900– 1904) Hamel, 1924–1930 + 1 – Scytosiphon lomentaria (Lyngbye, 1819) J. Agardh, 1848 2.3–445.0 1 – 4.0 9, 10, 12 – 5.0–35.5 2 – 3.0–465.0 2, 9, 10, 12 – 10.0 1, 10 – 240.0 6, 7, 10 – + 12 – 1449.0–3800.0 1, 3, 6, 9 – 0.005–2.2313* – 3, 5–12 Fam. Scytosiphonaceae Ordo Ralfsiales Fam. Heterochordariaceae Analipus filiformis (Ruprecht, 1850) Wynne, 1971 Fam. Ralfsiaceae Ralfsia fungiformis (Gunnerus, Setchell et Gardner, 1924 1772) Ordo Dictyotales Fam. Dictyotaceae Dictyota dichotoma (Hudson, 1762) Lamouroux, 1809 Ordo Sphacelariales Fam. Sphacelariaceae Sphacelaria furcigera Kuetzing, 1855 Ordo Fucales Fam. Sargassaceae Coccophora langsdorfii (Turner, 1819) Greville, 1830 Sargassum miyabei Yendo, 1907 S. pallidum (Turner, 1819) C. Agardh, 1820 Phylum Granuloreticulosea Classis Granuloreticulosea Subclassis Foraminifera 200–89250* Regnum Chlorobiontes Subregnum Thallobionti 105 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos 2.0 4, 10 – + 8 – 2.0 1 – 16.0 1, 3, 9, 10 – + 3 – Phyllospadix iwatensis Makino, 1931 8280.0 1 – Zostera marina Linnaeus, 1753 1290.0 4, 6, 8–10 – + 9, 12 – + 1, 3 – + 2 – Phylum Chlorophyta Ordo Codiales Fam. Bryopsidaceae Bryopsis plumosa (Hudson, 1778) C. Agardh, 1822 Fam. Codiaceae Codium fragile (Suringar, 1867) Hariot, 1889 Ordo Cladophorales Fam. Cladophoraceae Cladophora stimpsonii Harvey, 1859 Ordo Ulvales Fam. Ulvaceae Ulva fenestrata Postels et Ruprecht, 1840 Fam. Ulvellaceae Acrochaete viridis (Reinke, 1879) R. Nielsen, 1979 Subregnum Embryobionti Phylum Magnoliophyta Classis Monocotyledoneae Ordo Najadales Fam. Zosteraceae Regnum Prometazoa Phylum Spongia (=Porifera) Regnum Metazoa Phylum Cnidaria Subphylum Medusozoa Classis Hydrozoa Ordo Leptothecatae Fam. Campanulariidae Subfam. Obeliinae Obelia longissima (Pallas, 1766) Fam. Sertulariidae Sertularella mutsuensis Stechow, 1931 106 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos + 12 – 0.00019– 0.004845* – 2, 3, 5, 8–10, 12 + 12 – Classis Anthozoa Fam. Actiniidae Cnidopus japonicus (Verrill, 1868) Phylum Plathelminthes Classis Turbellaria 100–2550* Phylum Nemertea Fam. Lineidae Lineus torquatus Coe, 1901 Phyllum Annelida Classis Polychaeta Fam. Phyllodocidae Eulalia viridis (Linnaeus, 1767) 100 1.0 1 – Eulalia sp. juv. 510 0.0459 11 11 Gattyana ciliata Moore, 1902 100 1.0 2, 6 – Harmothoe imbricata (Linnaeus, 1767) 200 7.0 3, 6, 9, 10 – 300–400 0.027–0.036 – 12 1000; 80–2040* 8.3; 0.0072–0.1836* 3 2, 7 200 1.0 3 – Nereis multignatha Imajima et Hartman, 1964 100–1300 0.8–5.2 1, 7, 10, 12 – N. vexillosa Grube, 1849 300–1300 21.0–52.0 2, 3, 6, 12 – N. zonata Malmgren, 1867 100–1020* 0.009–0.0918* – 10–12 100 0.3 8, 11, 12 – Nereis sp. juv. 100; 510* 1.0; 0.0459* 4 8 Platynereis bicanaliculata (Baird, 1863) 100–200; 100–2040* 0.5–1.1; 0.009–0.1836* 3, 4, 6, 9, 10 ,12 7, 8, 10, 12 Exogone gemmifera Pagenstecher, 1862 20–1020* 0.0018–0.0918* – 2, 8, 12 Odontosyllis maculata Uschakov, 1950 100–200* 0.009–0.018* – 10 Pionosyllis compacta Malmgren, 1867 1020* 0.0918* – 8, 12 1530* 0.1377 3 6 400–2040* 0.036–0.1836* 3 8, 12 Fam. Polynoidae Fam. Pholoidae Pholoe longa (Mueller, 1776) Fam. Nereidae Lycastopsis pontica (Bobretzky, 1872) Neanthes sp. N. zonata tigrina Zachs, 1933 Fam. Syllidae Sphaerosyllis hirsuta Ehlers, 1897 Typosyllis fasciata Malmgren, 1867 107 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos + 3 – 400 0.4 6 – 200* 0.018* – 12 40* 0.0036* 3 2 2550* 0.2295* 3 – + – 6 200* 0.018* – 6 100; 200–2040* 0.2; 0.018–0.1836* 6 12 Heteromastus filiformis (Claparede, 1864) 100 0.8 5 – Notomastus sp. 400 3.3 5 – + 3, 6, 8, 11, 12 – + 3 – 0.027–0.018* – 12 3600 16.0 + – 510* 0.0459* – 11 100–16320* 0.009–1.4688* 3 2, 3, 5–8, 12 + 1 – 0.5 1–3, 6, 7, 10, 12 – T. orientalis Imajima et Hartman, 1964 T. pulchra occidentalis Buzhinskaja, 1980 Fam. Lumbrineridae Lumbrineris sp. Fam. Dorvilleidae Dorvillea (Schistomeringos) Annenkova, 1937 japonica Fam. Spionidae Polydora sp. juv. Fam. Cirratulidae Dodecaceria concharum Oersted, 1843 Chaetozone setosa Malmgren, 1867 Fam. Capitellidae Capitella capitata (Fabricius, 1780) Fam. Serpulidae Hydroides ezoensis Okuda, 1934 Fam. Sabellidae Eudistylia polymorpha (Johnson, 1901) Chone sp. juv. Fam. Spirorbidae Neodexiospira alveolata (Zachs, 1933) Classis Oligochaeta 200–300* Phylum Mollusca Classis Polyplacophora Subclassis Neoloricata Ordo Chitonida Fam. Ischnochitonidae Ischnochiton hakodadensis Pilsbry, 1892 Classis Gastropoda Subclassis Prosobranchia Fam. Lottiidae Nipponacmea moscalevi Chernyshev et Chernova, 2002 100 108 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 300–1600 16.8–180.0 20–1500 100 Site Macrobenthos Meiobenthos 1, 12 – + 1 – 0.16–13.0 2, 3, 6, 7, 7a, 9–12 – 8.3 1, 12 – 100–300 13.7–86.0 1–3, 7a, 9, 9a, 12 – Tegula rustica (Gmelin, 1790) 100 490.0–800.0 1–4, 9, 11, 12 – Lirularia iridescens (Schrenck, 1863) 100 0.8–1.2 1, 9 – Epheria turrita (A. Adams, 1861) 100–2400 0.4–8.3 3, 6, 7, 7a, 9, 10, 12 – Epheria sp. 40–3060* 0.0028–0.2142* – 3, 7 Littorina brevicula (Phillippi, 1844) 100–36400 2.4–7189.0 1–11 – L. mandshurica Schrenck, 1867 100–1500 150.0–1467.0 1, 3, 6, 7a, 8, 12 – L. sitkana (Phillippi, 1846) 200–4800 6.5–25.5 1, 12 – L. squalida Broderip et Sowerby, 1829 100–4000 0.4–1495.0 1–12 – 100 0.5 3 – 100–132800 0.1–344.0 1, 3, 5–12 – + 10 – 5100* 0.357* – 5 100–300 0.2–1.0 1, 3 – Lottia kogamogai Sasaki et Okutani, 1994 L. persona (Rathke, 1833) L. tenuisculpta Sasaki et Okutani, 1994 L. versicolor (Moskalev in Golikov et Scarlato, 1967) Fam. Turbinidae Homalopoma sangarense (Schrenck, 1867) Fam. Trochidae Fam. Lacunidae Fam. Littorinidae Fam. Pomatiopsidae Cecina manchurica A. Adams, 1861 Fam. Rissoidae Falsicingula athera Bartsch in Golikov et Scarlato, 1967 F. mundana (Yokoyama, 1926) Setia candida A. Adams, 1861 Lucidestea ornata Golikov et Kussakin in Golikov et Scarlato, 1967 Pusilina plicosa (Smith, 1875) 100–20000; 100–2550* 0.1–27.2; 1, 3, 5–7, 9, 10, 12 2, 5, 7, 11, 12 0.007–0.1785* Fam. Barleeidae Ansola angulosa Golikov et Kussakin, 1978 A. angustata (Pilsbry, 1901) 200–8100 0.2–16.0 1020–8160* 0.0714–0.5712* 1, 7a, 12 – 1, 7a, 9, 10, 12 5, 7, 8, 10 – 5, 7 1 3, 5–7, 9, 11, 12 1 3, 6, 7, 12 Fam. Assimineidae Assiminea sp. 200–4080* 0.014–0.2856* Fam. Caecidae Fartulum bucerius Golikov in Golikov et 180–363120* 0.0126–25.4184* Scarlato, 1967 Brochina derjugini Golikov in Golikov et 510–226950* 0.357–15.8865* Scarlato, 1967 109 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 200–3900 Site Macrobenthos Meiobenthos 0.3–18.4 1, 4, 6, 7a, 10 – 200–1400 38.3–300.0 4, 5, 7, 7a, 8 – Hima fratercula (Dunker, 1860) 300–400 28.0–47.0 3, 5, 6, 7, 7a, 9 – H. multigranosa (Dunker, 1847) 200 1.0 4, 6 – 100–1700 2.2–112.0 1, 2, 6, 10–12 – 100 55.0 11 – + 1, 9, 10 – 1000* 0.07* – 7 200 0.014 – 7 100–400* 0.0035–0.014* 11 11, 12 + – 11 0.0035 – 12 1.0–8.0; 0.0035–0.091* 1, 7, 9, 10 2, 6, 7, 9–12 Fam. Litiopsidae Alaba picta A. Adams, 1861 Fam. Batillariidae Batillaria cumingii (Crosse, 1862) Fam. Nassariidae Fam. Columbellidae Mitrella burchardi (Dunker, 1877) Fam. Muricidae Tritonalia japonica (Dunker, 1850) Fam. Thaididae Nucella heyseana (Dunker, 1882) Fam. Turbinellidae Odostomia culta Dall et Bartsch, 1906 Odostomia sp. Classis Bivalvia Subclassis Neotaxodonta Ordo Arcoida Superfam. Arcoidea Fam. Arcidae Arca boucardi Jousseaume, 1894 Subclassis Pteriomorphia Ordo Mytiloida Superfam. Mytiloidea Fam. Mytilidae Subfam. Mytilinae Mytilus (Mytilus) galloprovincialis Lamarck, 1819 100* Mytilus (Mytilus) trossulus kussakini Scarlato et Starobogatov, 1979 100–700; 100–2600* Mytilus (Crassimytilus) coruscus Gould, 1861 100–1500* 0.0035–0.0525* – 10, 12 510* 0.01785* – 11 100; 100–510* 10.0; 0.0035–0.01785* 1, 3, 8, 9, 12 6, 12 Mytilus sp. Crenomytilus grayanus (Dunker, 1853) 110 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 Arvella japonica (Dall, 1897) 600* Musculista senhousia (Benson in Cantor, 1842) 100* Site Macrobenthos Meiobenthos 0.021* – 12 0.0035* – 11 + 1 – Subfam. Crenellinae Vilasina pillula Bartsch in Scarlato, 1960 Subfam. Modiolinae Modiolus modiolus (Linnaeus, 1758) 100* 0.0035* – 9, 12 100–700* 0.0035–0.0245* 3 6, 9, 11, 12 100; 510* 5.0; 0.1785* 2, 3, 6–9 7 Chlamys (Azumapecten) farreri (Jones et Preston, 1904) + 10, 11 – Mizuhopecten yessoensis (Jay, 1857) + 9 – 100* 0.0035* – 12 100* 0.0035* – 12 Mercenaria stimpsoni (Gould, 1861) 200–510* 0.007–0.01785* – 7, 9, 11, 12 Venerupis (Ruditapes) philippinarum (A. Adams et Reeve, 1850) 100–300; 100–4080* 3.0–35.0; 0.0035–0.1428* 2, 4, 6, 8, 12 6, 7, 9–12 100–3600; 40–3200* 1.0–12.0; 0.0014–0.112* 1, 12 12 + – 9, 10 Septifer (Mytilisepta) keenae Nomura, 1936 Ordo Ostreoida Superfam. Ostreoidea Fam. Ostreidae Crassostrea gigas (Thunberg, 1793) Superfam. Pectinoidea Fam. Pectinidae Subclassis Heterodonta Ordo Veneroida Superfam. Lucinoidea Fam. Lucinidae Pillucina pisidium (Dunker, 1860) Superfam. Glossoidea Fam. Kelliellidae Alveinus ojianus (Yokoyama, 1927) Superfam. Veneroidea Fam. Veneridae Fam. Turtoniidae Turtonia minuta (Fabricius, 1780) Superfam. Tellinoidea Fam. Tellinidae Macoma (Macoma) contabulata (Deshayes, 1854) 100 53.0 5 – Macoma sp. 100* 0.0035* – 9 111 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 100; 510* Site Macrobenthos Meiobenthos 1.0–5.0; 0.01785* 3, 4, 6, 8 6 100* 0.0035* – 10 100* 0.0035* – 9 Teredo japonica Clessin in Kuster et Kobelt, 1893 510–1020* 0.01785–0.0357* – 6 Bankia setacea (Tryon, 1863) 100–510* 0.0035–0.01785* – 7 + 5, 8 – 500–7500* 0.00015– 0.00225* – 2, 3, 7 (larvae) 200–4100* 0.007–0.1435* – 2, 5, 6, 9–12 (juv.) 20–70000 0.8–1270.0 1–3, 5–12 + 200 130.0 12 – Ordo Myoida Subordo Myina Superfam. Myoidea Fam. Myidae Mya (Arenomya) arenaria Linnaeus, 1758 Mya sp. juv. Subordo Pholadina Superfam. Pholadoidea Fam. Pholadidae Barnea sp. Fam. Teredinidae Subclassis Anomalodesmata Ordo Pholadomyoida Superfam. Thracioidea Fam. Laternulidae Laternula (Exolaternula) marilina (Reeve, 1863) Phylum Arthropoda Subphylum Crustacea Superclassis Crustacea Classis Maxillopoda Subclassis Thecostraca Infraclassis Cirripedia Superordo Thoracica Ordo Sessilia Subordo Balanomorpha Superfam. Chthamaloidea Fam. Chthamalidae Subfam. Chthamalinae Chthamalus dalli Pilsbry, 1916 Superfam. Balanoidea Fam. Balanidae Subfam. Balaninae Balanus rostratus Hoek, 1883 112 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 300–500 Site Macrobenthos Meiobenthos 0.5–5.0 3, 4, 6, 7, 7a, 8, 8a,10 + + 6 – 0.0056–0.1224* – 2, 3, 6, 8–12 200–12240* 0.00032–0.0196* – 2, 3, 5–7, 9–12 Subfam. Amphibalaninae Amphibalanus improvisus (Darwin, 1854) Fam. Archaeobalanidae Subfam. Archaeobalaninae Hesperibalanus hesperius (Pilsbry, 1916) Subclassis Copepoda Infraclassis Neocopepoda Superordo Gymnoplea Ordo Calanoida 280–6120* Superordo Podoplea Ordo Harpacticoida 60–2550* 0.000018– 0.000765* – 3, 5, 6, 8–12 200–57000* 0.018–1.14* – 2–5, 6–9а, 10–12 300 2.7 1 – 700–1400 1.9–3.0 10, 12 – 400 2.1 1, 5, 6, 10, 12 – + 1 – 300 6.0 1 – 100–2700 0.08–6.3 1–3, 5–7, 10, 12 – Crassicorophium bonelli (Milne–Edwards, 1830) 400 0.008 – 6 Ericthonius tolli Bruggen, 1909 400 0.2 6, 7 – Monocorophium acherusicum (Costa, 1857) 300 0.2 8 – Ischyrocerus sp. 100–2600 0.1–0.5 1, 3, 12 – Jassa marmorata Holmes, 1903 60–1700 0.02–0.8 1–3, 6, 10, 12 – 100 0.1 1 – 100–200 0.1–0.3 1, 10 – Classis Ostracoda Classis Malacostraca Subclassis Eumalacostraca Superordo Peracarida Ordo Amphipoda Subordo Gammaridea Fam. Ampithoidae Ampithoe annenkovae Gurjanova, 1938 A. djakonovi Gurjanova, 1938 A. lacertosa Bate, 1858 A. tarasovi Bulytscheva, 1952 A. zachsi Gurjanova, 1938 Ampithoe sp. Fam. Corophiidae Fam. Ischyroceridae Fam. Dexaminidae Guernea (Guernea) quadrispinosa Stephensen, 1944 Paradexamine fraudatrix Tzvetkova, 1976 113 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 Calliopius laeviusculus (Kroyer, 1838) 100 Paracalliopiella litoralis (Gurjanova, 1938) Site Macrobenthos Meiobenthos 0.1–0.4 1, 10, 12 – 100 0.1 1 – 200–220 0.1–0.12 1, 5, 10 – Fam. Eusiridae Pontogeneia intermedia Gurjanova, 1938 P. rostrata Gurjanova, 1938 Pontogeneia sp. 200 0.2 2, 3 – 100–300 0.1–0.3 3, 6, 10, 12 – 100 0.1 1 – 100–900 1.2–3.2 5, 6, 8, 8a – 200 0.2–0.3 2, 3 – 200 0.6 7 – 100–1200 0.3–1.1 1, 12 – 100 0.1 1 – 100 0.8 1, 3, 6–8, 8a – Fam. Lysianassidae Orchomenella sp. Fam. Anisogammaridae Eogammarus possjeticus Tzvetkova, 1967 Fam. Melitidae Melita sp. Fam. Hyalidae Hyale bassargini Derzhavin, 1937 Hyale sp. Parallorchestes zibellina (Derzhavin, 1937) Fam. Talitridae Platorchestia pachypus (Derzhavin, 1937) Fam. Photidae Protomedeia sp. Subordo Caprellidea 100 0.002 – 6 200–25100 0.004–0.502 – 2, 6, 7, 10–12 200 0.1 1, 3 – 100–1100 0.2–0.7 1, 2, 7, 10 – + 3 – 2.6 12 – + 3, 12 – Fam. Caprellidae Caprella algaceus Vassilenko, 1967 C. bispinosa Mayer, 1890 C. borealis Mayer, 1903 C. cristibrachium Mayer, 1903 4300 C. danilevskii Czerniavski, 1868 C. laevis (Schurin, 1935) 100–500 0.1–0.2 1, 7 – C. mutica Schurin, 1935 1400 1.6 1, 12 – C. penantis Leach, 1814 200 0.1–0.6 1 – C. scaura diceros Mayer, 1890 100–1600 0.1–2.9 3, 6, 10 – Caprella sp. 200–1500 0.1–1.4 2, 3, 6, 10 – 200–6800 0.2–26.3 1, 3, 6, 7, 10 – Ordo Isopoda Subordo Flabellifera Fam. Sphaeromatidae Gnorimosphaeroma rayi Hoestlandt, 1969 114 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon Site N, ind./m2 B, g/m2 Macrobenthos Meiobenthos G. noblei Menzies, 1954 80–90270* 0.0016–1.8054* 1 2, 3, 5, 7, 9, 11, 12 G. ovatum (Gurjanova, 1933) 200–1020* 0.004–0.0204* 5 3, 10 Dynoides dentisinus Shen, 1929 100–600; 400–4080* 0.1–2.0; 0.008–0.0816* 1, 2, 12 2, 7, 12 Holotelson tuberculatus Richardson, 1909 100–800; 400–4080* 0.5–5.4; 0.008–0.0816* 1, 10, 11 – 100–300* 0.002–0.006* – 11, 12 + 3, 4 – 300–1020* 0.006–0.0204* 1 3, 6, 12 300–1000* 0.006–0.02* – 6, 10 1020* 0.0204* – 3, 6 + 1, 12 – 0.1836* – 3 – 3, 5, 12 Subordo Anthuridea Fam. Paranthuridae Paranthura japonica Richardson, 1909 Subordo Valvifera Fam. Idoteidae Idotea ochotensis Brandt, 1851 Subordo Aselota Fam. Janiridae Ianiropsis kincaidi Richardson, 1904 Fam. Munnidae Munna subneglecta Gurjanova, 1936 Munna sp. Subordo Oniscoidea Fam. Ligiidae Ligia cinerascens Budde-Lund, 1828 Fam. Detonidae Detonella papillicornis (Richardson, 1904) Armadilloniscus ellipticus (Harger, 1875) 9180* 1020–13260* 0.0204–0.2652* Ordo Tanaidacea 60–2040* 0.0012–0.0408* – 6, 7, 11, 12 Ordo Cumacea 200–5100* 0.004–0.102* – 2, 6–9, 11, 12 + 9 – + 4 – Superordo Eucarida Ordo Decapoda Fam. Palaemonidae Leander serrifer macrodactylus Rathbun, 1902 Fam. Crangonidae Crangon septemspinosa f. typica Say Subordo Anomura Fam. Paguridae Pagurus brachiomastus (Thallwitz, 1891) 100 30.0 2, 3, 6, 7, 7a, 10 – P. middendorffii Brandt, 1851 200 5.5–40.2 1–3, 6, 7, 10 – P. hirsutiusculus (Dana, 1851) 100 8.7 3–6 – 115 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos + 4 – + 3, 6 – 100–500 0.8–58.0 2–7, 8, 9, 11 – 100 1.5–105.1 1–3, 6, 7, 9, 10, 12 – + 8 – 200–2040* 0.0001–0.00102* – 3, 5, 9–12 200–7140* 0.0002–0.00714* – 6, 7, 9–12 200* 0.0002* – 9 – 6, 7, 12 – 5, 8 – 3, 6, 7 Fam. Callianassidae Upogebia issaeffi (Balss, 1913) Fam. Canceridae Cancer amphioetus Rathbun, 1898 Fam. Grapsidae Hemigrapsus penicillatus (de Haan, 1835) H. sanguineus (de Haan, 1835) Eriocheir japonicus de Haan, 1835 Subphylum Chelicerata Classis Arachnoidea Ordo Acarina Phylum Nemathelminthes Classis Nematoda Subclassis Enoplia Ordo Enoplida Fam. Enoplidae Enoplus anisospiculus Nelson, Hopper et Webster, 1972 Fam. Thoracostomopsidae Enoplolaimus medius Pavlyuk, 1984 Fam. Anticomidae Anticoma possjetica Platonova, Belogurov et Scheenko, 1979 500–85170* 0.0005–0.08517* Fam. Oncholaimidae Admirandus multicavus Belogurov et Belogurova, 1979 510–2550* Oncholaimium domesticum Chitwood et Chitwood, 1938 500–10710* 0.0005–0.01071* O. japonicum Belogurov et Belogurova, 1981 0.00051– 0.00255* 2040* 0.00204* – 8 O. paraolium Belogurov et Fadeeva, 1980 100–71400* 0.0001–0.0714 – 6–9, 9a, 11 O. ramosum Smolanko et Belogurov, 1987 120–63240* 0.00012– 0.06324* – 2, 5–7 O. unicum Belogurov et Belogurova, 1978 300–62730* 0.0003–0.06273* Oncholaimus brachycercus de Man, 1889 40–5610* 0.00004– 0.00561* Pseudoncholaimus furugelmus Belogurov, 1977 200–1020* 0.0002–0.00102* 116 – 6, 7, 12 – 3, 9, 9a, 10, 11 – 6, 11 Composition and distribution of benthos in the intertidal zone Table 1 (continued) Taxon N, ind./m2 B, g/m2 Site Macrobenthos Meiobenthos P. mediocaudatus Pavlyuk, 1991 200–78030* 0.0002–0.07803* – 2, 5–7, 10 P. vesicarius (Wieser, 1959) 500–1600* 0.0005–0.0016* – 2, 5, 12 40–78540* 0.00004– 0.07854* – 6, 9, 9a, 10 800* 0.0008* – 12 1020* 0.00102* – 8 Dorylaimopsis peculiaris Platonova, 1971 2550* 0.00255* – 8 Sabatieria possjetica Platonova, 1971 100* 0.0001* – 2, 5 200–600* 0.0002–0.0006* – 10, 12 400–1020* 0.0004–0.00102* – 8, 12 1530* 0.00153* – 8 1020* 0.00102* – 8 2550–58140* 0.00255– 0.05814* – 5, 6, 8 200* 0.0002* – 10 2040* 0.00204* – 5 Daptonema variasetosa (Pavlyuk, 1984) 200–5100* 0.0002–0.0051* – 2, 3, 5–8, 12 Daptonema sp. 1020–1530* 0.00102– 0.00153* – 3 200–510* 0.0002–0.00051* – 7 2040* 0.00204* – 11 510* 0.00051* – 11 300–3570* 0.0003–0.00357* – 2, 5, 8, 10, 12 Fam. Enchelidiidae Eurystomina alekseevi Pavlyuk, 1991 Fam. Phanodermatidae Phanoderma platonovae Belogurov, 1980 Fam. Oxystominidae Tycnodora rectispiculata Platonova, 1971 Ordo Chromadorida Fam. Comesomatidae S. pulchra (Schneider, 1906) Fam. Chromadoridae Euchromadora robusta Kulikov, 1991 Euchromadora sp. Fam. Selachinematidae Halichoanolaimus possjetiensis Belogurov et Fadeeva, 1980 H. sonorus Belogurov et Fadeeva, 1980 Fam. Cyatholaimidae Paracanthonchus macrodon (Ditlevsen, 1919) Ordo Monhysterida Fam. Sphaerolaimidae Sphaerolaimus limosus Fadeeva, 1983 Fam. Xyalidae Paramonohystera halerba Fadeeva et Belogurov, 1987 Theristus subacer Pavlyuk, 1984 Fam. Linhomoeidae Therschellingia glabricutis Platonova, 1971 Ordo Desmodorida Fam. Monoposthiidae Monoposthia latiannulata Platonova, 1971 117 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 1 (continued) Taxon N, ind./m2 Phylum Bryozoa B, g/m2 + Site Macrobenthos Meiobenthos + – Phylum Echinodermata Classis Echinoidea Ordo Camarodonta Fam. Strongylocentrotidae Strongylocentrotus nudus (A. Agassiz, 1863) + 12 – S. intermedius (A. Agassiz, 1863) + 1, 6, 12 – + 1, 6, 7a, 9 – + 12 – Asterias amurensis Luetken, 1871 + 1, 3, 4, 6, 7a – Classis Ophiuroidea 0.00003– 0.000765* – 2, 3, 6, 8, 12 + 9, 12 – + 3 – + 5 – Classis Asteroidea Ordo Spinulosa Fam. Asterinidae Asterina pectinifera (Mueller et Troschel, 1842) Ordo Forcipulata Fam. Asteriidae Distolasterias nipon (Doderlein, 1902) 100–2550* Phylum Chordata Subphylum Tunicata Сlassis Ascidiacea Ordo Pleorobranchiata Fam. Styelidae Botryllus tuberatus Ritter et Forsyth, 1917 Classis Teleostomi Subclassis Actinopterygii Ordo Perciformes Subordo Blennioidei Fam. Stichaeidae Neozoarces pulcher Steindachner, 1880 Subordo Gobioidei Fam. Gobiidae Luciogobius guttatus Gill, 1859 *Meiobenthos density and biomass. N o t e . Sites mean hydrobiological transects shown in Fig. 1. «+» – qualitative data, «–» – not found. Here and in the tables below: N – population density, ind./m2, B – biomass, g/m2. 118 Composition and distribution of benthos in the intertidal zone Table 2 Composition and quantitative characteristics of Chthamalus dalli community in the rocky and boulder-pebble intertidal zone of the Russky Island Taxon The upper intertidal horizon The middle intertidal horizon The lower intertidal horizon N, ind./m2 N, ind./m2 N, ind./m2 B, g/m2 B, g/m2 B, g/m2 Macrobenthos Plants Corallina pilulifera Masudaphycus irregularis Gloiopeltis furcata Ceramium kondoi Cyanobacteria Polysiphonia japonica – – – – 3.33 – – 48.57 10.36 4.57 – 0.03 112.0 – – – – – Plants, total 3.33 63.53 112.0 Animals Chthamalus dalli Littorina mandshurica Balanus rostratus Falsicingula athera Littorina brevicula Lottia kogamogai Littorina sitkana Littorina squalida Pagurus brachiomastus Lottia tenuisculpta Lottia versicolor Crassostrea gigas Platynereis bicanaliculata Hima acutidentata Epheria turrita Dynoides dentisinus Nipponacmea moskalevi Amphibalanus improvisus Caprella bispinosa Hyale sp. Ansola angulosa Gnorimosphaeroma rayi Calliopius laeviusculus Pusilina plicosa Ampithoe sp. Jassa marmorata 28017 17 – – 83 33 – – – – 17 – – – – 83 – – – – – – 17 – – – 525.33 27.17 – – 23.73 2.05 – – – – 1.38 – – – – 0.33 – – – – – – 0.02 – – – 17700 14 – 329 143 43 – 43 14 214 – 14 – – 286 71 29 43 71 29 29 29 – 14 14 14 427.24 21.43 – 0.74 43.53 2.4 – 7.07 4.29 1.86 – 0.71 – – 0.43 0.07 0.14 0.07 0.06 0.04 0.03 0.03 – 0.01 0.01 0.01 14800 600 100 30000 – 350 2400 50 – 200 – – 100 100 – 50 – – – – – – – – – – 1127.5 140.0 65.0 60.0 – 33.5 12.75 0.2 – 1.0 – – 0.55 0.5 – 0.05 – – – – – – – – – – Macrobenthic animals, total 28267 580.01 19143 510.16 48750 1441.05 Total biomass of macrobenthos 583.34 119 573.69 1553.05 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 2 (continued) The upper intertidal horizon The middle intertidal horizon The lower intertidal horizon N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 Polychaeta Gastropoda Fartulum buceriu Pusilina plicosa Brochina derjugini Epheria sp. Odostomia sp. Balanomorpha (juv.) Gammaridea Oligochaeta Bivalvia Mytilus trossulus kussakini Venerupis philippinarum Septifer keenae Turtonia minuta Isopoda Munna subneglecta Ianiropsis kincaidi Gnorimosphaeroma noblei Paranthura japonica Caprellidea Cumacea Tanaidacea Cirripedia (larvae) Ophiuroidea Cyanobacteria 180 140 – – 140 – – 1920 300 180 60 60 – – – 140 – – 140 – – 40 – – 20 0.0162 0.0098 – – 0.0098 – – 0.0672 0.006 0.0162 0.0021 0.0021 – – – 0.0028 – – 0.0028 – – 0.0008 – – 0.000006 + 200 560 300 140 – 80 40 820 240 160 120 40 40 – 40 180 – – 120 60 240 200 – 420 – – 0.018 0.0392 0.021 0.0098 – 0.0056 0.0028 0.0287 0.0076 0.0144 0.0042 0.0014 0.0014 – 0.0014 0.0036 – – 0.0024 0.0012 0.0048 0.004 – 0.000126 – – 1200 1100 750 350 – – – 1200 960 – 400 300 50 50 – 650 500 150 – – 650 150 150 – – – 0.108 0.077 0.0525 0.0245 – – – 0.042 0.0192 – 0.014 0.0105 0.00175 0.00175 – 0.013 0.01 0.003 – – 0.013 0.003 0.003 – – – Pseudomeiobenthos, total 2980 0.121106 3140 0.117786 6460 0.2922 40 – 180 340 0.001 – 0.000288 0.00034 – 220 380 620 – 0.0044 0.000608 0.00062 950 850 5000 5050 0.02375 0.017 0.008 0.00505 40 0.00004 80 0.00008 2150 0.00215 – 80 40 60 – 20 – – 0.00008 0.00004 0.00006 – 0.00002 – – 60 – – 140 240 100 – 0.00006 – – 0.00014 0.00024 0.0001 850 750 650 400 250 – – 0.00085 0.00075 0.00065 0.0004 0.00025 – – Taxon Meiobenthos Pseudomeiobenthos Eumeiobenthos Foraminifera Calanoida Harpacticoida Nematoda Pseudoncholaimus mediocaudatus Anticoma possjetica Eurystomina alekseevi Daptonema variasetosa Enoplus anisospiculus Oncholaimium unicum Oncholaimium paraolium Pseudoncholaimus vesicarius 120 Composition and distribution of benthos in the intertidal zone Table 2 (continued) Taxon Monopostia latiannulata Pseudoncholaimus furugelmus Turbellaria Ostracoda Acarina Eumeiobenthos, total Meiobenthos, total The upper intertidal horizon The middle intertidal horizon The lower intertidal horizon N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 N, ind./m2 60 40 40 80 40 0.00006 0.00004 0.000076 0.000024 0.00002 – – 180 – – – – 0.000342 – – 720 0.001748 1400 3700 0.122854 4540 Total biomass 583.46 B, g/m2 – – – 500 200 – – – 0.00015 0.0001 0.00597 12560 0.05405 0.123756 19010 0.34625 573.81 1553.40 N o t e . «+» – qualitative data, «–» – not found. Table 3 Composition and quantitative characteristics of Littorina brevicula community in the intertidal zone of Russky Island Taxon The upper intertidal horizons, rocks and blocks The middle intertidal horizon, stones and rocks The lower intertidal horizon, stones N, ind./m2 N, ind./m2 N, ind./m2 B, g/m2 B, g/m2 B, g/m2 Macrobenthos Plants Gloiopeltis furcata 0.17 3.0 – Animals Littorina brevicula Chthamalus dalli Hemigrapsus penicillatus Hemigrapsus sanguineus Nereis vexillosa Mya arenaria Lottia tenuisculpta Nereis multignatha Amphibalanus improvisus Gnorimosphaeroma rayi Hyale bassargini Typosyllis pulchra occidentalis Falsicingula athera Calliopius laeviusculus Capitella capitata 4900 19733 – – – – – – – – – – 67 – – 2588.73 80.0 – – – – – – – – – – 0.07 – – 8420 10720 20 20 200 20 120 20 100 120 40 80 – 20 20 773.2 196.0 11.6 5.16 5.1 1.0 0.72 0.6 0.4 0.32 0.12 0.08 – 0.04 0.04 6100 6500 – – – – – – – – – – – – – 278.0 30.0 – – – – – – – – – – – – – Macrobenthic animals, total 24700 2668.8 19920 991.38 12600 308.0 Total biomass of macrobenthos 2668.97 121 994.38 308.0 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 3 (continued) The upper intertidal horizons, rocks and blocks The middle intertidal horizon, stones and rocks The lower intertidal horizon, stones N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 – – – – 5430 1500 400 50 200 – – – – – 150 – 150 – 100 50 250 250 0.036 0.0045 0.004 – – – – – 0.003 – 0.003 – 0.002 0.001 0.000075 0.000075 1785 1275 4470 685 1247 1020 127 100 1580 1530 – 50 1317 1275 1530 – 0.3801 + + 0.16065 0.11475 0.0894 0.23975 0.043662 0.0357 0.004462 0.0035 0.0316 0.0306 – 0.001 0.02635 0.0255 0.00046 – 200 300 1200 1700 – – – – – – – – – – – – 0.105 + + 0.018 0.027 0.024 0.0595 – – – – – – – – – – – – 1450 0.05065 20594 0.896447 4900 0.2335 Foraminifera Nematoda Oncholaimium paraolium Anticoma possjetica Pseudoncholaimus mediocaudatus Oncholaimium unicum Daptonema variasetosa Enoplus anisospiculus Paramonochystera halebra Eurystomina alekseevi Calanoida Harpacticoida Turbellaria Ostracoda – 200 – – – 0.0002 – – 6982 10327 2677 2295 0.168937 0.010327 0.002677 0.002295 – 1000 800 – – 0.001 0.0008 – Eumeiobenthos, total Taxon Meiobenthos Pseudomeiobenthos Gastropoda Brochina derjugini Fartulum bucerius Polychaeta Oligochaeta Gammaridea Balanomorpha (juv.) Bivalvia Venerupis philippinarum Mercenaria stimpsoni Mytilus coruscus Isopoda Gnorimosphaeroma noblei Munna subneglecta Gnorimosphaeroma ovatum Caprellidea Cumacea Cirripedia (larvae) Ophiuroidea Pseudomeiobenthos, total Eumeiobenthos Meiobenthos, total – – 1785 0.001785 – – – – – – 200 – 200 50 – – – – – 0.0002 – 0.00032 0.000095 – 1657 893 893 127 – 75 – – 75 0.001657 0.000893 0.000893 0.000127 – 0.0015 – – 0.000023 – – – 200 – – – – – – – – 0.0002 – – – – – 450 0.000615 17459 0.180787 1000 0.001 1900 0.051265 38053 1.077234 5900 0.2345 Total biomass 2669.021 N o t e . «+» – qualitative data, «–» – not found. 122 995.457 308.234 Composition and distribution of benthos in the intertidal zone Table 4 Composition and quantitative characteristics of Corallina pilulifera community in the lower horizon of the rocky boulder and stony intertidal zone of Russky Island Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Corallina pilulifera Scytosiphon lomentaria Analipus japonicus Dictyota dichotoma 641.0 111.25 25.0 21.25 Plants, total 798.5 Animals Falsicingula athera Pusilina plicosa Littorina brevicula Littorina squalida Homalopoma sangarense Nereis multignatha Ansola angustata Mytilus trossulus kussakini juv. Turtonia minuta Crenomytilus grayanus juv. Ampithoe zachsi Ansola angulosa Epheria turrita Venerupis philippinarum Chthamalus dalli Hyale sp. Lucidestea ornata Platynereis bicanaliculata Holotelson tuberculatus Ampithoe sp. Nereis zonata tigrina Dynoides dentisinus Jassa marmorata Guernea quadrispinosa 58150 6200 300 1025 25 425 1450 225 400 25 75 1200 1000 25 25 325 25 25 25 125 25 25 75 25 119.2 9.1 7.55 4.85 3.425 2.9 2.8 2.75 2.5 2.5 1.5 1.4 1.0 0.75 0.5 0.425 0.125 0.125 0.125 0.1 0.075 0.05 0.025 0.025 Macrobenthic animals, total 71225 163.8 Total biomass of macrobenthos 962.3 Meiobenthos Pseudomeiobenthos Gammaridea Gastropoda Fartulum bucerius Brochina derjugini 14400 2300 123 0.288 0.161 + + M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 4 (continued) Taxon N, ind./m2 B, g/m2 Pusilina plicosa Polychaeta Caprellidea Bivalvia Mytilus trossulus kussakini Venerupis philippinarum Septifer keenae Turtonia minuta Mercenaria stimpsoni Modiolus modiolus Macoma sp. Barnea sp. Bankia setacea Balanomorpha (juv.) 1767 400 2266 867 733 267 200 67 33 33 33 33 367 + 0.159 0.008 0.079334 0.030333 0.025667 0.009333 0.007 0.002333 0.001167 0.001167 0.001167 0.001167 0.012833 Pseudomeiobenthos, total 21500 0.744167 Foraminifera Calanoida Nematoda Oncholaimium paraolium Pseudoncholaimus vesicarius Enoplus anisospiculus Pseudoncholaimus mediocaudatus Anticoma possjetica Monoposthia latiannulata Daptonema variasetosa Phanoderma platonovae Oncholaimium domesticum Oncholaimium unicum Sabatieria pulchra Eurystomina alekseevi Euchromadora robusta Oncholaimus brachycercus Enoplolaimus medius Harpacticoida Turbellaria Ostracoda Acarina 1333 233 4399 767 533 433 333 333 333 333 267 200 200 200 167 133 100 67 1533 200 433 233 0.033333 0.004667 0.004399 0.000767 0.000533 0.000433 0.000333 0.000333 0.000333 0.000333 0.000267 0.0002 0.0002 0.0002 0.000167 0.000133 0.0001 0.000067 0.002453 0.00038 0.00013 0.000116 Eumeiobenthos, total 8364 0.045478 29864 0.789645 Eumeiobenthos Meiobenthos, total Total biomass 963.090 N o t e . «+» – qualitative data. 124 Composition and distribution of benthos in the intertidal zone Table 5 Composition and quantitative characteristics of Littorina mandshurica community in the intertidal zone of Russky Island Taxon The upper intertidal horizon, boulders and blocks N, ind./m2 B, g/m2 The middle intertidal horizon, pebble-sandy bottom N, ind./m2 B, g/m2 Macrobenthos Plants Corallina pilulifera Polysiphonia japonica Clathromorphum compactum – – – 45.0 1.4 + Plants, total – 46.4 Animals Littorina mandshurica Littorina squqlida Nereis vexillosa Chthamalus dalli Homalopoma sangarense Gnorimosphaeroma rayi Hemigrapsus penicillatus Amphibalanus improvisus Mya arenaria juv. Epheria turrita Melita sp. Caprella spp. Pontogeneia sp. 1150 – – 200 – – – – – – – – – 921.0 – – 6.5 – – – – – – – – – 300 200 1300 4600 100 1300 100 300 100 200 200 500 100 463.0 55.0 52.0 25.0 15.0 5.1 5.0 5.0 1.0 0.6 0.3 0.3 0.1 Macrobenthic animals, total 1350 927.5 9300 627.4 Total biomass of macrobenthos 927.5 673.8 Meiobenthos Pseudomeiobenthos Oligochaeta Gastropoda Fartulum bucerius Brochina derjugini Epheria sp. Gammaridea – – – – – – – – – – – – 7140 1530 510 510 510 5100 0.6426 0.1071 0.0357 0.0357 0.0357 0.102 Pseudomeiobenthos, total – – 13770 0.8517 – – – – – – 7650 6120 12240 0.19125 0.1224 0.0196 Eumeiobenthos Foraminifera Calanoida Harpacticoida 125 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 5 (continued) Taxon The upper intertidal horizon, boulders and blocks The middle intertidal horizon, pebble-sandy bottom N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 Nematoda Oncholaimus brachycercus Daptonema variasetosa Oncholaimium domesticum Oncholaimium paraolium Ostracoda – – – – – – – – – – – – 16320 5610 5100 3570 2040 1530 0.01632 0.00561 0.0051 0.00357 0.00204 0.00046 Eumeiobenthos, total – – 43860 0.35003 Meiobenthos, total – – 57630 1.20173 Total biomass 927.5 675.002 N o t e . «+» – qualitative data, «–» – not found. Table 6 Composition and quantitative characteristics of Littorina squalida community in the intertidal zone of Russky Island Taxon The middle intertidal horizon, pebble-sandy silted bottom N, ind./m2 B, g/m2 The lower intertidal horizon, stones N, ind./m2 B, g/m2 Macrobenthos Plants Gracilaria gracilis Ralfsia fungiformis Bossiella compressa Clathromorphum compactum 45.0 – – – – 35.5 4.2 + Plants, total 45.0 39.7 Animals Littorina squalida Tegula rustica Hemigrapsus sanguineus Homalopoma sangarense Venerupis philippinarum Nereis vexillosa Pagurus hirsutiusculus Harmothoe imbricata Eugammarus possjeticus Lottia tenuisculpta Heteromastus filiformis 100 – – – – – 100 – 900 – 100 79.0 – – – – – 8.7 – 3.2 – 0.8 126 900 100 100 300 300 300 – 200 – 100 – 1495.0 800.0 90.0 59.0 35.0 21.0 – 7.0 – 1.2 – Composition and distribution of benthos in the intertidal zone Table 6 (continued) Taxon The middle intertidal horizon, pebble-sandy silted bottom The lower intertidal horizon, stones N, ind./m2 B, g/m2 N, ind./m2 – – – – 200 200 0.2 0.1 1200 91.7 2700 2508.5 Melita sp. Caprella sp. Macrobenthic animals, total Total biomass of macrobenthos 136.7 B, g/m2 2548.2 Meiobenthos Pseudomeiobenthos Oligochaeta Gammaridea Polychaeta Lycastopsis pontica Dorvillea japonica Exogone gemmifera Syllidae gen. sp. juv. Isopoda Gnorimosphaeroma noblei Gastropoda Pusilina plicosa Cirripedia (larvae) 12240 4080 – – – – – 510 510 – – – 1.1016 0.0816 – – – – – 0.0102 0.0102 – – – – 220 160 80 40 20 20 80 80 100 100 420 – 0.0044 0.0144 0.0072 0.0036 0.0018 0.0018 0.0016 0.0016 0.007 0.007 0.000126 Pseudomeiobenthos, total 16830 1.1934 980 0.027526 Foraminifera Nematoda Pseudoncholaimus mediocaudatus Oncholaimium ramosum Sabatieria possjetica Pseudoncholaimus vesicarius Daptonema variasetosa Monoposthia latiannulata Calanoida Harpacticoida Ostracoda Turbellaria 3570 15810 6120 5100 3570 1020 – – – 2550 2040 – 0.08925 0.0158 0.00612 0.0051 0.00357 0.00102 – – – 0.00408 0.0006 – 240 1080 320 120 100 160 240 140 280 220 60 100 0.006 0.00108 0.00032 0.00012 0.0001 0.00016 0.00024 0.00014 0.0056 0.000362 0.000018 0.00019 Eumeiobenthos, total 23970 0.10973 1980 0.01325 Meiobenthos, total 40800 1.30313 2960 0.040776 Eumeiobenthos Total biomass 138.003 N o t e . «+» – qualitative data, «–» – not found. 127 2548.241 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 7 Composition and quantitative characteristics of Lottia kogamogai community in the upper boulder-block intertidal horizon of Karpinsky Bay (Russky Island) Taxon N, ind./m2 B, g/m2 1600 2800 200 600 100 180.0 98.0 6.5 0.5 0.3 5300 285.3 Polychaeta Gastropoda Balanomorpha (juv.) Isopoda Dynoides dentisinus Paranthura japonica Bivalvia Turtonia minuta Pillucina pisidium Mytilus trossulus kussakini 200 200 400 700 400 300 400 200 100 100 0.018 0.014 0.014 0.014 0.008 0.006 0.014 0.007 0.0035 0.0035 Pseudomeiobenthos, total 1900 0.074 Harpacticoida Nematoda Pseudoncholaimus vesicarius Oncholaimium unicum Acarina 700 1000 700 300 200 0.00112 0.001 0.0007 0.0003 0.0001 Eumeiobenthos, total 1900 0.00222 Meiobenthos, total 3800 0.07622 Macrobenthos Animals Lottia kogamogai Chthamalus dalli Littorina sitkana Dynoides dentisinus Falsicingula athera Macrobenthic animals, total Meiobenthos Pseudomeiobenthos Eumeiobenthos Total biomass 285.376 Table 8 Composition and quantitative characteristics of Macoma contabulata+Hima fratercula community on the silted sand with pebble admixture in the lower intertidal horizon, southwest of Cape Akhlestyshev (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Polysiphonia japonica Gracilaria gracilis 20.0 4.0 Plants, total 24.0 128 Composition and distribution of benthos in the intertidal zone Table 8 (continued) Taxon N, ind./m2 B, g/m2 100 400 400 200 53.0 47.0 3.3 0.2 1100 103.5 Animals Macoma contabulata Hima fratercula Notomastus sp. Pontogeneia intermedia Macrobenthic animals, total Total biomass of macrobenthos 127.5 Meiobenthos Pseudomeiobenthos Oligochaeta Gammaridea 9180 1530 0.8262 0.0306 10710 0.8568 Foraminifera Nematoda Pseudoncholaimus mediocaudatus Halichoanolaimus sonorus Admirandus multicavus Sphaerolaimus limosus Ostracoda 7650 13260 5100 3570 2550 2040 1020 0.19125 0.01326 0.0051 0.00357 0.00255 0.00204 0.000306 Eumeiobenthos, total 21930 0.204816 Meiobenthos, total 32640 1.061616 Pseudomeiobenthos, total Eumeiobenthos Total biomass 128.562 Table 9 Composition and quantitative characteristics of Hemigrapsus penicillatus+Gnorimosphaeroma rayi community on the pebble ground in the middle intertidal horizon of Ajaks Bay (Russky Island) Taxon N, ind./m2 B, g/m2 500 6800 100 100 1400 1000 100 38.7 26.3 29.3 10.9 4.0 2.5 1.0 Macrobenthos Animals Hemigrapsus penicillatus Gnorimosphaeroma rayi Littorina brevicula Littorina squalida Epheria turrita Pusilina plicosa Lucidestea ornata 129 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 9 (continued) Taxon N, ind./m2 Neantes sp. juv. Hemigrapsus sp. juv. Cecina manсhurica Pontogeneia sp. B, g/m2 200 200 100 400 1.0 0.8 0.5 0.4 10900 115.4 5100 2040 1020 510 510 1530 4590 3570 0.459 0.1428 0.0714 0.0375 0.0375 0.1377 0.0918 0.00107 16830 0.83237 Calanoida Harpacticoida Nematoda Oncholaimus brachycercus Oncholaimium domesticum Daptonema variasetosa Daptonema sp. 6120 10710 14790 5610 4590 3570 1020 0.1224 0.01714 0.01479 0.00561 0.00459 0.00357 0.00102 Eumeiobenthos, total 31620 0.15433 Meiobenthos, total 48450 0.9867 Macrobenthic animals, total Meiobenthos Pseudomeiobenthos Oligochaeta Gastropoda Fartulum bucerius Brochina derjugini Epheria sp. Polychaeta Gammaridea Cirripedia (larvae) Pseudomeiobenthos, total Eumeiobenthos Total biomass 116.388 Table 10 Composition and quantitative characteristics of Grateloupia turuturu community in the lower horizon of the rocky intertidal zone of Ajaks Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Grateloupia turuturu Polysiphonia japonica Ralfsia fungiformis (on Littorina spp.) 4200.0 8.0 5.0 Plants, total 4213.0 130 Composition and distribution of benthos in the intertidal zone Table 10 (continued) Taxon N, ind./m2 B, g/m2 Animals Spirorbidae Littorina brevicula 36000 100 16.0 15.0 Macrobenthic animals, total 36100 31.0 Total biomass of macrobenthos 4244.0 Meiobenthos Pseudomeiobenthos Caprellidea Polychaeta Gammaridea Isopoda Gnorimosphaeroma noblei Cirripedia (larvae) 3100 600 2100 200 200 7500 0.062 0.054 0.042 0.004 0.004 0.00225 Pseudomeiobenthos, total 11400 0.16425 Harpacticoida Turbellaria 600 200 0.00096 0.00038 Eumeiobenthos, total 800 0.00134 12200 0.16559 Eumeiobenthos Meiobenthos, total Total biomass 4244.166 Table 11 Composition and quantitative characteristics of Neorhodomela larix aculeata community in the lower horizon of the block intertidal zone southeast of Cape Staritsky in Novik Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Neorhodomela larix aculeata Corallina pilulifera Ceramium japonicum Gracilaria gracilis 3300.0 25.0 10.5 3.2 Plants, total 3338.7 Animals Falsicingula athera Littorina squalida Epheria turrita Pagurus middendorffii Holotelson tuberculatus 132800 200 1400 200 700 131 344.0 32.6 8.3 5.5 5.4 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 11 (continued) Taxon N, ind./m2 Ansola angustata Pusilina plicosa Ampithoe djakonovi Caprella scaura diceros Ampithoe lacertosa Mytilus trossulus kussakini juv. Caprella spp. Nereis multignatha Caprella bispinosa Alaba picta Paradexamine fraudatrix Jassa marmorata Pontogeneia sp. Macrobenthic animals, total B, g/m2 3200 3200 1400 1600 400 700 1500 100 1100 300 200 100 100 4.8 3.2 3.0 2.9 2.1 2.0 1.4 0.8 0.7 0.4 0.3 0.1 0.1 149200 417.6 Total biomass of macrobenthos 3756.3 Meiobenthos Pseudomeiobenthos Gammaridea Caprellidea Bivalvia Mytilus trossulus kussakini Turtonia minuta Mytilus coruscus Venerupis philippinarum Mya sp. Tellinidae gen. sp. Gastropoda Polychaeta Platynereis bicanaliculata Odontosyllis maculata Nereis zonata 57000 25100 2000 1100 500 100 100 100 100 800 300 100 100 100 1.14 0.502 0.07 0.0385 0.0175 0.0035 0.0035 0.0035 0.0035 0.056 0.027 0.009 0.009 0.009 Pseudomeiobenthos, total 85200 1.795 Eumeiobenthos Turbellaria Harpacticoida Nematoda Monoposthia latiannulata Enoplus anisospiculus Sabatieria pulchra Paracanthonchus macrodon Ostracoda Acarina 1200 1100 1300 500 400 200 200 900 500 0.00228 0.00184 0.0013 0.0005 0.0004 0.0002 0.0002 0.00027 0.00025 Eumeiobenthos, total 5000 0.00594 90200 1.80094 Meiobenthos, total Total biomass 3758.101 132 Composition and distribution of benthos in the intertidal zone Table 12 Composition and quantitative characteristics of macrobenthos in the community Neorhodomela munita in the lower intertidal horizon east of Cape Ivantsova (Russky Island) Taxon N, ind./m2 B, g/m2 Plants Neorhodomela munita Sargassum pallidum Corallina pilulifera 1650.0 300.0 115.0 Plants, total 2065.0 Animals Falsicingula athera Pagurus middendorffii Pusilina plicosa Homalopoma sangarense Turtonia minuta Mytilus trossulus kussakini Mitrella burchardi Ampithoe sp. Littorina squalida Holotelson tuberculatus Caprella mutica Ansola angulosa Eulalia viridis Lucidestea ornata Jassa marmorata Ampithoe lacertosa Guernea quadrispinosa Ischyrocerus sp. Pontogeneia intermedia 28800 200 17600 100 3600 500 300 2100 300 200 1400 300 100 100 200 100 100 100 200 64.0 40.2 27.2 18.0 12.0 8.0 4.5 2.9 1.6 1.6 1.6 1.0 1.0 0.9 0.6 0.5 0.1 0.1 0.1 Animals, total 56300 185.9 Total biomass 2250.9 Table 13 Composition and quantitative characteristics of Gracilaria gracilis community in the lower horizon of the silted sand with small stones intertidal zone in Melkovodnaya Bay (Voevoda Bay, Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Gracilaria gracilis Polysiphonia japonica 330.0 4.0 Plants, total 334.0 133 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 13 (continued) Taxon N, ind./m2 B, g/m2 Batillaria cumingii Littorina mandshurica Eogammarus possjeticus Monocorophium acherusicum 200 300 100 300 213.0 151.0 1.2 0.2 Macrobenthic animals, total 900 365.4 Animals Total biomass of macrobenthos 699.4 Meiobenthos Pseudomeiobenthos Oligochaeta Gastropoda Ansola angustata Pusilina plicosa Cumacea Ophiuroidea 2040 2040 1530 510 2550 1020 0.1836 0.1428 0.1071 0.0357 0.051 0.000306 Pseudomeiobenthos, total 7650 0.377706 Foraminifera Nematoda Oncholaimium paraolium Oncholaimium japonicum Daptonema variasetosa Euchromadora sp. Euchromadora robusta Halichoanolaimus possjeticus Turbellaria Ostracoda 7140 13770 6120 2040 2040 1530 1020 1020 2550 2550 0.1785 0.01377 0.00612 0.00204 0.00204 0.00153 0.00102 0.00102 0.004845 0.000765 Eumeiobenthos, total 26010 0.19788 Meiobenthos, total 33660 0.575586 Eumeiobenthos Total biomass 699.976 Table 14 Composition and quantitative characteristics of Lomentaria hakodatensis community on the upper rubble surface in the lower intertidal horizon of Karpinsky Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Lomentaria hakodatensis Dictyota dichotoma 260.0 17.0 Plants, total 277.0 134 Composition and distribution of benthos in the intertidal zone Table 14 (continued) Taxon N, ind./m2 B, g/m2 Falsicingula athera Jassa marmorata Ampithoe sp. Hyale sp. Ischyrocerus sp. Calliopius laeviusculus 20000 1700 600 500 200 100 66.8 0.7 0.4 0.3 0.1 0.1 Macrobenthic animals, total 23100 68.4 Animals Total biomass of macrobenthos 345.4 Meiobenthos Pseudomeiobenthos Gammaridea Caprellidea Polychaeta Pholoe longa Capitella capitata Chone sp. Gastropoda Pusilina plicosa 13600 6800 800 400 200 200 1000 1000 0.272 0.136 0.072 0.036 0.018 0.018 0.07 0.07 Pseudomeiobenthos, total 22200 0.55 Eumeiobenthos Foraminifera Ostracoda Eumeiobenthos, total Meiobenthos, total 7300 400 0.1825 0.00012 7700 0.18262 29900 0.73262 Total biomass 346.133 Table 15 Composition and quantitative characteristics of Ceramium japonicum+Ceramium kondoi community in the lower horizon of the rubble intertidal zone southeast of Cape Staritsky in Novik Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Ceramium japonicum Ceramium kondoi Laurencia nipponica 94.0 64.0 44.0 135 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 15 (continued) Taxon N, ind./m2 Corallina pilulifera Neorhodomela larix aculeata Analipus filiformis Dictyota dichotoma Bryopsis plumosa Polysiphonia japonica B, g/m2 40.0 10.0 4.0 3.0 2.0 1.0 Plants, total 262.0 Animals Falsicingula athera Chthamalus dalli Lottia tenuisculpta Pontogeneia intermedia Ampithoe sp. Jassa marmorata 22080 20 20 220 100 60 49.28 0.8 0.16 0.12 0.08 0.02 Macrobenthic animals, total 22500 50.46 Total biomass of macrobenthos 312.46 Meiobenthos Pseudomeiobenthos Gammaridea Caprellidea Gastropoda Fartulum bucerius Epheria sp. Bivalvia Turtonia minuta Tanaidacea 5500 2340 220 180 40 40 40 60 0.11 0.0468 0.0154 0.0126 0.0028 0.0014 0.0014 0.0012 Pseudomeiobenthos, total 8160 0.1748 Eumeiobenthos Foraminifera Harpacticoida Turbellaria Nematoda Pseudoncholaimus mediocaudatus Oncholaimus brachycercus Eurystomina alekseevi Ostracoda 540 240 160 180 100 40 40 200 0.0135 0.000384 0.000304 0.00018 0.0001 0.00004 0.00004 0.00006 Eumeiobenthos, total 1320 0.014428 Meiobenthos, total 9480 0.189228 Total biomass 312.649 136 Composition and distribution of benthos in the intertidal zone Table 16 Composition and quantitative characteristics of Gloiopeltis furcata community on blocks and rubbles in the middle intertidal horizon of Russky Island Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Gloiopeltis furcata Chthamalus dalli Tritonalia japonica Littorina squalida Holotelson tuberculatus 139.75 Animals Macrobenthic animals, total 1350 50 50 400 28.0 27.5 3.5 0.45 1850 59.45 Total biomass of macrobenthos 199.2 Meiobenthos Pseudomeiobenthos Gastropoda Fartulum bucerius Bivalvia Mytilus trossulus kussakini Septifer keenae Mytilidae gen. sp. Arca boucardi Venerupis philippinarum Mercenaria stimpsoni Crenomytilus grayanus Musculista senhousia Polychaeta Nereis zonata Platynereis bicanaliculata Cumacea Gammaridea Isopoda Gnorimosphaeroma noblei Paranthura japonica 5500 5500 4500 2600 700 700 400 400 200 100 100 400 200 200 600 300 400 300 100 0.385 0.385 0.1572 0.091 0.0245 0.0245 0.014 0.014 0.007 0.0035 0.0035 0.036 0.018 0.018 0.012 0.006 0.008 0.006 0.002 Pseudomeiobenthos, total 11700 0.943 Eumeiobenthos Foraminifera Harpacticoida Nematoda Oncholaimium paraolium Enoplus anisospiculus Ostracoda Acarina 4100 1100 1200 700 500 500 200 0.105 0.00176 0.0012 0.0007 0.0005 0.00015 0.0001 Eumeiobenthos, total 7100 0.10821 18800 1.05121 Meiobenthos, total Total biomass 200.251 137 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 17 Composition and quantitative characteristics of Campylaephora crassa community on blocks in the middle intertidal horizon of Karpinsky Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Campylaephora crassa Dictyota dichotoma 620.0 22.0 Plants, total 642.0 Animals Chthamalus dalli Mitrella burchardi Caprella cristibrachium Falsicingula athera Ischyrocerus sp. Jassa marmorata Ansola angustata Ampithoe sp. Dynoides dentisinus Pontogeneia sp. Macrobenthic animals, total 9900 100 4300 400 2600 600 300 100 100 200 110.0 19.3 2.6 0.7 0.5 0.4 0.3 0.1 0.1 0.1 18600 134.1 Total biomass of macrobenthos 776.1 Meiobenthos Pseudomeiobenthos Gastropoda Fartulum bucerius Brochina derjugini Pusilina plicosa Polychaeta Bivalvia Turtonia minuta Venerupis philippinarum Mytilus coruscus Arvella japonica Mytilus trossulus kussakini Mercenaria stimpsoni Modiolus modiolus Crenomytilus grayanus Mytilus galloprovincialis Arca boucardi Alveinus ojanus Oligochaeta Balanomorpha (juv.) Gammaridea 11200 7400 3300 500 3700 9400 3200 2700 1500 600 500 400 100 100 100 100 100 2100 1000 1300 138 0.784 0.518 0.231 0.035 0.333 0.329 0.112 0.0945 0.0525 0.021 0.0175 0.014 0.0035 0.0035 0.0035 0.0035 0.0035 0.189 0.035 0.026 Composition and distribution of benthos in the intertidal zone Table 17 (continued) Taxon N, ind./m2 Isopoda Paranthura japonica Ophiuroidea B, g/m2 200 200 400 0.004 0.004 0.00012 29300 1.72012 Foraminifera Nematoda Pseudoncholaimus vesicarius Monoposthia latiannulata Oncholaimium unicum Turbellaria Harpacticoida Ostracoda 2200 2400 1200 700 500 900 1000 300 0.055 0.0024 0.0012 0.0007 0.0005 0.00171 0.0016 0.00009 Eumeiobenthos, total 6800 0.0608 36100 1.78092 Pseudomeiobenthos, total Eumeiobenthos Meiobenthos, total Total biomass 777.881 Table 18 Composition and quantitative characteristics of macrobenthos in the community Sargassum pallidum in the lower horizon of the rubble intertidal zone east of Cape Ivantsova (Russky Island) Taxon N, ind./m2 B, g/m2 Plants Sargassum pallidum Neorhodomela munita Ahnfeltia tobuchiensis Ulva fenestrata Polysiphonia morrowii Sphacelaria furcigera Lithophyllum tumidulum Grateloupia divaricata Gelidium vagum Scytosiphon lomentaria Cladophora stimpsoni 3800.0 120.0 37.0 16.0 12.0 10.0 10.0 9.0 8.5 2.3 2.0 Plants, total 4026.8 Animals Mitrella burchardi Falsicingula athera Alaba picta Pagurus middendorffii Ampithoe sp. Ampithoe annenkovae 1700 11000 3900 200 2700 300 139 112.0 20.1 18.4 11.4 6.3 2.7 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 18 (continued) Taxon N, ind./m2 Littorina squalida Nereis multignatha Turtonia minuta Platorchestia pachypus Caprella penantis Lucidestea ornata Jassa marmorata Orchomenella sp. Caprella algaceus Animals, total B, g/m2 200 100 100 100 200 100 300 100 200 1.9 1.4 1.0 0.8 0.6 0.5 0.1 0.1 0.1 21200 177.4 Total biomass 4204.2 Table 19 Composition and quantitative characteristics of Coccophora langsdorfii community in the lower horizon of the pebble intertidal zone of Rynda Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Coccophora langsdorfii Polysiphonia japonica 240.0 4.0 Plants, total 244.0 Animals Falsicingula athera Littorina brevicula Hima fratercula Lottia tenuisculpta Epheria turrita Mytilus trossulus kussakini juv. Littorina squalida Ampithoe sp. Ericthonius tolli Caprella laevis 61600 300 300 800 800 100 800 200 400 500 140.8 51.7 28.0 6.0 1.6 1.0 0.8 0.4 0.2 0.2 Macrobenthic animals, total 65800 230.7 Total biomass of macrobenthos 474.7 Meiobenthos Pseudomeiobenthos Gastropoda Ansola angustata Odostomia culta 2900 1600 1000 140 0.203 0.112 0.07 Composition and distribution of benthos in the intertidal zone Table 19 (continued) Taxon N, ind./m2 B, g/m2 Assiminea sp. Epheria sp. Polychaeta Syllidae gen sp. juv. Lycastopsis pontica Platynereis bicanaliculata Oligochaeta Gammaridea Caprellidea Cirripedia (larvae) 200 100 1000 400 300 300 700 3100 2300 1700 0.014 0.007 0.09 0.036 0.027 0.027 0.063 0.062 0.046 0.00051 Pseudomeiobenthos, total 11700 0.46451 Foraminifera Nematoda Pseudoncholaimus mediocaudatus Oncholaimium paraolium Oncholaimium domesticum Oncholaimium ramosum 2100 3500 1500 1200 500 300 0.0525 0.0035 0.0015 0.0012 0.0005 0.0003 Eumeiobenthos, total 5600 0.056 Eumeiobenthos Meiobenthos, total 17300 Total biomass 0.52051 475.220 Table 20 Composition and quantitative characteristics of macrobenthos in the community Chordaria flagelliformis in the middle horizon of the rubble intertidal zone east of Cape Ivantsov (Russky Island) Taxon N, ind./m2 B, g/m2 Plants Chordaria flagelliformis 1246.0 Animals Falsicingula athera Littorina squalida Ansola angustata Mitrella burchardi Turtonia minuta Lirularia iridescens Dynoides dentisinus Jassa marmorata Pusilina plicosa Ampithoe sp. 4950 300 650 50 150 50 100 450 250 350 141 16.25 2.8 2.25 1.1 0.8 0.6 0.6 0.4 0.3 0.25 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 20 (continued) Taxon N, ind./m2 Calliopius laeviusculus Lucidestea ornata Caprella penantis Caprella laevis Paracalliopiella litoralis Parallorchestes zibellina 50 50 50 50 50 50 Animals, total 7600 Total biomass B, g/m2 0.2 0.1 0.1 0.05 0.05 0.05 25.9 1271.9 Table 21 Composition and quantitative characteristics of Dictyota dichotoma community in the middle horizon of the rocky intertidal zone in Karpinsky Bay (Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Dictyota dichotoma Saundersella simplex Neorhodomela larix aculeata Chordaria flagelliformis Corallina pilulifera Laurencia pinnata 465.0 360.0 130.0 90.0 10.0 1.0 Plants, total 1056.0 Animals Falsicingula athera Ampithoe djakonovi Epheria turrita Hyale sp. 72000 700 100 600 108.0 1.9 0.8 0.7 Macrobenthic animals, total 73400 111.4 Total biomass of macrobenthos 1167.4 Meiobenthos Pseudomeiobenthos Gammaridea Caprellidea Polychaeta Typosyllis fasciata Pholoe longa Capitella capitata Lumbrineris sp. Gastropoda Pusilina plicosa 11800 6200 1100 400 300 200 200 700 700 0.236 0.124 0.099 0.036 0.027 0.018 0.018 0.049 0.049 Pseudomeiobenthos, total 19800 0.508 142 Composition and distribution of benthos in the intertidal zone Table 21 (continued) Taxon N, ind./m2 B, g/m2 Calanoida Harpacticoida Turbellaria 500 1200 400 0.01 0.00192 0.00076 Eumeiobenthos, total 2100 0.01268 21900 0.52068 Eumeiobenthos Meiobenthos, total Total biomass 1167.921 Table 22 Composition and quantitative characteristics of macrobenthos in the community Phyllospadix iwatensis in the lower horizon of the rubble intertidal zone east of Cape Ivantsova (Russky Island) Taxon N, ind./m2 B, g/м2 Plants Phyllospadix iwatensis Neorhodomela munita Sargassum pallidum Ahnfeltia tobuchiensis 8280.0 3450.0 1449.0 6.9 Plants, total 13185.9 Animals Homalopoma sangarense Ansola angulosa Mitrella burchardi Alaba picta Ampithoe sp. Hemigrapsus sanguineus Turtonia minuta Littorina squalida Lucidestea ornata Lirularia iridescens Pusilina plicosa Caprella bispinosa Falsicingula athera Jassa marmorata Paradexamine fraudatrix Caprella penantis Lysianassidae Animals, total Total biomass 200 8100 200 1600 1900 100 500 100 300 100 400 100 100 100 100 200 100 28.2 16.0 5.3 4.7 1.7 1.5 1.3 1.0 0.9 0.8 0.6 0.2 0.1 0.1 0.1 0.1 0.1 14200 61.2 13247.1 143 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 23 Composition and quantitative characteristics of Zostera marina community on the silted bottom in the lower intertidal horizon of Melkovodnaya Bay (Voevoda Bay, Russky Island) Taxon N, ind./m2 B, g/m2 Macrobenthos Plants Zostera marina 1290.0 Animals Batillaria cumingii Hemigrapsus sanguineus Littorina squalida 400 100 400 300.0 105.1 104.0 Macrobenthic animals, total 900 509.1 Total biomass of macrobenthos 1799.1 Meiobenthos Pseudomeiobenthos Polychaeta Platynereis bicanaliculata Exogone gemmifera Pionosyllis compacta Typosyllis fasciata Nereis sp. juv. Oligochaeta Gastropoda Ansola angustata Gammaridea 4590 1530 1020 1020 510 510 2040 1020 1020 1530 0.4131 0.1377 0.0918 0.0918 0.0459 0.0459 0.1836 0.0714 0.0714 0.0306 Pseudomeiobenthos, total 9180 0.6987 4590 6120 2550 2550 510 510 2040 0.11475 0.00612 0.00255 0.00255 0.00051 0.00051 0.000612 Eumeiobenthos, total 12750 0.121482 Meiobenthos, total 21930 0.820182 Eumeiobenthos Foraminifera Nematoda Dorylaimopsis peculiaris Halichoanolaimus sonorus Admirandus multicavus Monoposthia latiannulata Ostracoda Total biomass 1799.920 144 Composition and distribution of benthos in the intertidal zone Table 24 Composition and quantitative characteristics of meiobenthos under Zostera marina washed ashore by storms in the intertidal and supralittoral zones of Russky Island The upper intertidal horizon, sandy bottom Taxon Paris Bay N, ind./m2 Rynda Bay B, g/m2 N, ind./m2 Supralittoral, sandy-pebble bottom Ajaks Bay B, g/m2 N, ind./m2 B, g/m2 Pseudomeiobenthos Gastropoda Fartulum bucerius+ Brochina derjugini Ansola angustata Setia candida Assiminea sp. Epheria sp. Pusilina plicosa Isopoda Gnorimosphaeroma noblei Armadilloniscus ellipticus Detonella papillicornis Dynoides dentisinus Gnorimosphaeroma ovatum Ianiropsis kincaidi Munna sp. Oligochaeta Polychaeta Platynereis bicanaliculata Sphaerosyllis hirsuta Bivalvia Venerupis philippinarum Mytilus trossulus kussakini Teredo japonica Crenomytilus grayanus Septifer keenae Mya arenaria Crassostrea gigas Mercenaria stimpsoni Bankia setacea Bivalvia fam. gen. sp. Gammaridea Tanaidacea Insecta (larvae) Caprellidea Ophiuroidea 590070 41.3049 6120 0.4284 31620 2.2134 590070 41.3049 6120 0.4284 8670 0.6069 – – – – – – – – – – – – – – 3570 2040 1530 4590 510 2040 510 510 510 510 – – – – 5610 – – – 2040 – – – – – – – – – – – – – – 0.1836 0.1836 0.1377 0.16065 0.01785 0.0714 0.01785 0.01785 0.01785 0.01785 – – – – 0.1122 – – – 0.000612 – – – – – 94350 90270 – – 4080 – – – 5610 – – – 7650 4080 510 1020 – – – 510 510 510 510 2040 2040 – – – – – – – – 1.887 1.8054 – – 0.0816 – – – 0.5049 – – – 0.26775 0.1428 0.01785 0.0375 – – – 0.01785 0.01785 0.01785 0.01785 0.0408 0.0408 – – – 8160 5100 4080 3060 2550 27540 1020 13260 9180 – 1020 1020 1020 13260 – – – – – – – – – – – – – – 2040 – 2040 1020 2550 0.5712 0.357 0.2856 0.2142 0.1785 0.5508 0.0204 0.2652 0.1836 – 0.0204 0.0204 0.0204 1.1934 – – – – – – – – – – – – – – 0.0408 – 0.0408 0.0204 0.000765 Pseudomeiobenthos, total 605880 41.891362 117810 3.16965 80070 4.060365 145 M.B. Ivanova, L.S. Belogurova and A.P. Tsurpalo Table 24 (continued) The upper intertidal horizon, sandy bottom Taxon Paris Bay Rynda Bay Supralittoral, sandy-pebble bottom Ajaks Bay N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 N, ind./m2 B, g/m2 Foraminifera Nematoda Anticoma possjetica Eurystomina alekseevi Pseudoncholaimus mediocaudatus Oncholaimium paraolium Oncholaimium ramosum Oncholaimium unicum Halichoanolaimus sonorus Enoplus anisospiculus Daptonema variasetosa Oncholaimus brachycercus Oncholaimium domesticum Harpacticoida Turbellaria Acarina Ostracoda 89250 497250 85170 78540 2.2313 0.49725 0.08517 0.07854 26010 22950 3570 – 0.65025 0.02295 0.00357 – 6120 5100 – – 78030 0.07803 – – – – 71400 63240 62730 58140 – – – – 8670 – – – 0.0714 0.06324 0.06273 0.05814 – – – – 0.0139 – – – 7650 – 4590 – 7140 – – – – – – – 0.00765 – 0.00459 – 0.00714 – – – – – – – – – – – – 2550 1530 1020 – 2550 1530 2040 – – – – – 0.00255 0.00153 0.00102 – 0.0048 0.000765 0.0006 Eumeiobenthos, total 595170 2.74245 48960 0.6732 17340 0.164265 Meiobenthos, total 1201050 44.633812 166770 3.84285 97410 4.22463 Eumeiobenthos N o t e . «–» – not found. 146 0.153 0.0051 – – Fig. 2. Intertidal zone of Ajaks Bay. Fig. 3. Intertidal zone of a shallow lagoon near Cape Akhlestysheva. Fig. 4. Intertidal zone of Paris Bay. Fig. 5. Intertidal zone of Rynda Bay. Fig. 6. Intertidal zone of Voevoda Bay (Melkovodnaya Bay). Fig. 7. Intertidal zone of Novik Bay (Cape Ekipazhny). Fig. 8. Rocky intertidal zone of Karpinsky Bay. Fig. 9. Boulder intertidal zone of Karpinsky Bay. Fig. 10. Intertidal zone of Voevoda Bay. Chthamalus dalli and Littorina brevicula populations on rocks (Transect 9). Fig. 11. Intertidal zone of Voevoda Bay. Corallina pilulifera populations on rocks (Transect 9). Fig. 12. Intertidal zone of Voevoda Bay. Chthamalus dalli, Gloiopeltis furcata and Littorina brevicula populations on rocks (Transect 9). Fig. 13. Pollution of intertidal and supralittoral zones in Zhitkova Bay (Paris Bay). Fig. 14. Percent ratios of pseudomeiobenthos and eumeiobenthos in the intertidal zone of Russky Island by mean biomass. Fig. 15. Percent ratios of the meiobenthos groups in the intertidal zone of Russky Island by mean biomass. «Others» include the following groups: Calanoida, Cumacea, Harpacticoida, Tanaidacea, Turbellaria, Cirripedia larvae, Ostracoda, Acarina, Ophiuroidea. Fig. 16. Percent ratios of pseudomeiobenthos and eumeiobenthos in the intertidal zone of Russky Island by mean population density. Fig. 17. Percent ratios of meiobenthos groups in the intertidal zone of Russky Island by mean population density. «Others» include the following groups: Turbellaria, Cumacea, Tanaidacea, Ophiuroidea, Acarina.
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