composition and distribution of benthos in the intertidal zone of

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).
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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
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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.
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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)
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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)
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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
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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
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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).
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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).
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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).
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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.