Late pleistocene-holocene ostracod assemblages of the Northern Caspian Sea shelf

Analysis of the ostracod assemblages and the distribution of benthic foraminifera Ammonia beccarii in the core section GS 194-08-1 recovered at a depth of 7.5 m near the Volga delta front showed changes in environmental conditions in response to sea-level fluctuations. The pre-Novocaspian ostracod assemblage reflects the conditions that were dominant during the Late Khvalynian transgression, with a depth of about 20–30 m, and a salinity range of 12–13‰. The Early Novocaspian sediments contain an ostracod assemblage that represents a deeper water environment compared to the present, with an inferred depth of 12–16 m. The faunas from the younger Novocaspian sediments with abundant monospecific populations of Cyprideis torosa and co-occurring benthic foraminifera Ammonia beccarii represent modern-like environmental conditions.     

Distribution of some rare and trace elements in modern bottom sediments of the Caspian Sea

The features of the distribution of some rare and trace elements in modern bottom sediments of the Caspian Sea have been studied from samples collected during cruises 35, 39, and 41 of the R/V Rift and the cruise of the R/V Nikifor Shurekov in 2013. It was established that bottom sediments in different areas of the Caspian Sea vary to some degree in the contents of Zr, Hf, Th, V, Cr, Co, Ni, Cu, Sr, and Ba in comparison to suspended matter discharged by the Volga River and rivers originating in the Caucasus. As follows from the results of a comparison of the geochemical features of modern bottom sediments of the Volga River delta and different sedimentary subsystems of the Caspian Sea with the chemical composition of Middle Archean granitoids and Paleozoic and Mesozoic–Cenozoic basalts, which are regarded as geochemical images of such provenance areas as the basement of East European Platform, Urals, and Caucasus, none of these regions is considered the dominant provenance area for all sedimentary subsystems of the Caspian Sea region. Here, the revealed similarity between modern bottom sediments of the northern, central and southern Caspian Sea regions, the Volga River delta, and Post-Archean average Australian shale (PAAS) in some parameters, including REE spectra, assumes that the Volga River discharge plays a dominant role in the formation of the geochemical image of Caspian Sea subsystems. The role of clastics, including the fine-grained fraction, which is supplied to the Caspian Sea from the Caucasus region and Elburz Mountains, is insignificant already in the coastal area, which is determined both by influence of the marginal filter (MF) and large-scale cholestatic current circulation.     

Diatoms in surface sediments of the Sea of Okhotsk and the northwest Pacific Ocean

The present investigation was targeted at diatom composition studies in the surface sediments (0–1 cm) sampled in the Sea of Okhotsk and the northwest Pacific in the depth range from 130 to 6110 m. The taxonomic analysis, as well as the quantitative (the diatom cell abundance per sediment dry weight unit) content and ecological group definition, was applied. Ten diatom taxa are the main body (80–100%) of the diatom assemblages: Bacterosira bathyomphala, Chaetoceros spp. (spores), Actinocyclus curvatulus, Thalassiosira latimarginata (group), T. antarctica (spores), Neodenticula seminae, Rhizosolenia hebetata f. hiemalis, Thalassiothrix longissima, Coscinodiscus marginatus, Coscinodiscus oculus iridis. The relative content of these species reflects the sedimentation conditions for different parts of the sea: the shelf, the continental slope, the open sea, and the ocean. The highest diatom content (45.6.3–60.0 10⁶ per g of dry weight) was found for the surface sediments in the central part of the Sea of Okhotsk and the continental slope of western Kamchatka.     

Benthic foraminiferal assemblages as indicators of the paleoceanographic conditions in the eastern Equatorial Pacific

Five assemblages of benthic foraminifers reflecting the changes in the properties of the intermediate water masses and the biological productivity during different periods of the last glacial-interglacial cycle are defined based on the abundances and relative contents of indicative species in the upper 9 m of giant Core MD02-2529 from the Cocos Ridge. High bioproductivity and low oxygen content in the bottom water layer and sediments are established for the interstadial period based on the high abundance of the species Uvigerina peregrina, U. hispida, and C. pachyderma (Assemblage I) and especially for the Last Glacial Maximum (Assemblage II) with the dominant role of the same species and E. smithi. The transition from the glaciation to the current interglacial (Termination I, Assemblage III) is characterized by a high share of the epifaunal species C. wuellerstorfi and H. subhaidingeri, which indicates enhanced hydrodynamic activity and ventilation of the intermediate water washing the bottom. The end of Termination I was marked by strengthened selective dissolution of carbonate microfossils due to the increased influx of fresh organic matter to the bottom and/or advection of more aggressive intermediate waters. The diverse Early Holocene assemblage (Assemblage IV) points to variable ecological niches, while the abundance of the infaunal U. hispida in Assemblage V indicates low productivity of the surface waters and a low oxygen concentration in the bottom sediments of the Late Holocene.     

Spatial distribution and feeding of dominant zooplankton species in the Ob River estuary

The distribution and feeding of dominant mesozooplankton species were studied in the estuary of the Ob River and adjacent inner Kara Sea shelf waters in September 2013. It was shown that the spatial distributions of Cyclops sp., Senecella siberica, Limnocalanus macrurus, Mysis oculata, Drepanopus bungei, Jashnovia tolli and Pseudocalanus sp. are related to the specific characteristics of the hydrographic regime in the estuarine frontal zone. The distributions of Cyclops sp., Senecella siberica, and Pseudocalanus sp. are mainly limited by salinity, while other species inhabit an area with a wide range of salinity values without clear preferences. Peaks of their abundance could be either consolidated or distanced in space. The populations of Jashnovia tolli, Drepanopus bungei, and Pseudocalanus sp. permanently inhabit the layer under the pycnohalocline; the populations of Cyclops sp. and Mysis oculata inhabit the upper mixed layer. Limnocalanus macrurus demonstrates a different vertical distribution pattern: the copepod undertakes diel vertical migrations in the southern part of the estuarine frontal zone; in its northern part, the population is concentrated below the pycnocline during day and night. The differences in the distributions of the studied species determine their feeding behavior and their role in phytoplankton grazing. The most intense utilization of biomass and production of autotrophic phytoplankton by zooplankton occur in the freshened water zone and the adjacent southern periphery of the estuarine frontal zone: the total daily phytoplankton consumption makes up 10–18% of the biomass and 60–380% of primary production. Daily zooplankton consumption of phytoplankton in the estuarine frontal zone decreases to 2–7% of the biomass and to 14% of primary production; in inner shelf waters, the values do not exceed 1% for both phytoplankton biomass and production.     

Biology and distribution of hyperiids in the Sea of Okhotsk

Long-term studies in the Sea of Okhotsk (1986–2012) demonstrated that hyperiids account for a small portion of zooplankton, making up only from 1.0 to 5.3% by weight. The minimum hyperiid biomass was observed in the coastal zone. The hyperiid biomass in the open water shelf community increased from spring to autumn. In the Sea of Okhotsk, hyperiids are represented by ten species, among which the most abundant in the northern regions (mainly in Shelikhov Gulf) is Themisto libellula, while the most abundant in other regions is T. pacifica. The distribution of T. pacifica in different seasons showed that the maximum biomass was concentrated in the deep-water zone. The range of T. libellula in the Sea of Okhotsk is mostly limited to the Shelikhov Gulf, but in certain years its habitation area can expand. In warm years with low ice coverage, the maximum expansion of T. libellula to the west in the shelf zone of the northern part of the Sea of Okhotsk was observed. Since variability of salinity and temperature in the northern part of the Sea of Okhotsk may be fatal for the cryophilic stenohaline T. libellula species, forecasted changes in thermohaline circulation will make it possible to predict the population dynamics of this important species.     

酸化过程对不同碳酸钙含量的海洋沉积物中总氮及其同位素分析的影响

为节约成本和样品,一些学者同时分析海洋沉积物中的碳、氮及其同位素（TOC、TN、δ¹³C和δ¹⁵N）。分析沉积物中的δ¹³C,需要对样品进行酸化去除无机碳,但是这一酸化过程会使TN和δ¹⁵N的分析结果产生偏差,且偏差范围与沉积物中无机碳含量（CaCO₃）有关。本研究选取了低CaCO₃含量（1-16%）和高CaCO₃含量（20-40%）的海洋沉积物样品,比较了酸化过程对TN和δ¹⁵N的影响。研究结果表明,酸化过程对海洋沉积物中TN和δ¹⁵N的分析结果产生了显著影响。对于低CaCO₃含量的样品,酸化导致样品中TN流失了约0-40%,δ¹⁵N偏移了约0-2‰;而对于高CaCO₃含量的样品,酸化导致样品中TN流失了约10-60%,δ¹⁵N偏移了约1-14‰。表明酸化对TN和δ¹⁵N的影响已经超过了仪器的误差范围0.002%（TN）和0.08‰（δ¹⁵N）,将影响TN和δ¹⁵N的环境指示意义。因此,即使海洋沉积物样品中CaCO₃含量很低,也必须用原样分析TN和δ¹⁵N以避免酸化过程的影响。     

Dynamics of the abundance of some bivalve species in Russian waters of the Sea of Japan and its prognosis

The abundance dynamics of several species of bivalve mollusks spats were studied on scallop collectors situated in Minonosok bay of Pos’eta Gulf for 27 years and for 4 years in Kit bay of the Sea of Japan (Russia). A significant positive relation was found between the species having similar thermopathy: the Japanese scallop Mizuhopecten yessoensis and Swift’s scallop Swiftopecten swifti, as well as between the wrinkled rock borer Hiatella arctica and Swift’s scallop Swiftopecten swifti. A significant reverse relation was found between the bay mussel Mytilus trossulus and the Northern Pacific seastar Asterias amurensis. Some of the studied mollusks of Minonosok bay and the remote Kit bay display a significant reversed interrelation in their abundance dynamics caused by the precipitation regime. The one-way dispersion analysis a revealed significant influence of the water temperature in June and the precipitation abundance in the summer on Swift’s scallop’s dynamic abundance. The two-way dispersion analysis showed a significant influence of the ice period duration and the solar activity expressed in Wolf’s numbers on the Japanese scallop abundance dynamics. The uneven years in the period from 1977 to 1984 were usually productive for M. yessoensis and S. swifti spat. After 1985, the even years became more productive (there was asynchronicity in the abundance dynamics compared with 1977–1984). Such asynchronicity appeared with the advent of the new 22-year solar cycle, which caused a change in the magnet polarity in 1986.     

Fine-scale Microbial Communities Associated with Manganese Nodules in Deep-sea Sediment of the Korea Deep Ocean Study Area in the Northeast Equatorial Pacific

Despite its potential significance for industrial utilization, any activities associated with the mining of manganese (Mn) nodules might have substantial impacts on benthic ecosystems. Because microorganisms respond quickly to changing environmental conditions, a study of microbial communities provides a relevant proxy to assess possible changes in benthic ecosystems associated with mining activities. We investigated fine-scale microbial community composition and diversity inside and on the surface of Mn nodules and in nearby deep-sea sediments in the Korea Deep Ocean Study (KODOS) area located in the Clarion-Clipperton Fracture Zone (CCFZ) of the northeast equatorial Pacific. Although microbial cell density was lower within nodules (3.21 × 10⁶ cells g^-1) than in sediment (2.14 × 10⁸ cells g^-1), nodules provided a unique habitat for microorganisms. Manganese-oxidizing bacteria including Hyphomicrobium and Aurantimonas in Alphaproteobacteria and Marinobacter in Gammaproteobacteria were abundant in nodules, which implied that these bacteria play a significant role in nodule formation. In contrast, Idiomarina in Gammaproteobacteria and Erythrobacter and Sulfitobacter in Alphaproteobacteria were abundant in sediments. Meanwhile, Thaumarchaeota, a phylum that consists of ammonia-oxidizing chemolithoautotrophs, were the predominant archaeal group both in nodules and sediment. Overall, microbial communities in Mn nodules were unique compared to those observed in sediments. Furthermore, the phylogenetic composition of microorganisms in the KODOS area was distinguishable from that in the nodule provinces claimed by China and Germany in the CCFZ and nodule fields in the central South Pacific Gyre, respectively.     

Morphological features,ecology, and distribution of poorly studied molluscan genera of the Colinae subfamily (Bastropoda,Buccinidae) from the far eastern seas of Russia

Data on the distribution of six genera of poorly studied buccinids of the Colinae subfamily (Neogastropoda, Buccinidae), namely, Colus Röding, 1799; Plicifusus Dall, 1902; Retifusus Dall, 1916; Aulacofusus Dall, 1918; Pararetifusus Kosuge, 1967; and Latisipo Dall, 1916, are presented. These mollusks are widely spread in the North Pacific region dwelling predominantly over loose sediments in a wide range of sea depths. Based on the morphology and contents of their digestive tracts, it is assumed that the representatives of the genera studied are predators with diverse diets. It is supposed that the increase in the dwelling depths had no significant influence on the feeding ecology of the species studied. Meanwhile, the lower abundance of preys at greater depths caused the lower population densities and modifications in the proboscis structure of selected taxa.     

东印度洋现生浮游有孔虫

A brief morphometric study of the recent planktonic foraminifera in the eastern Indian Ocean was provided with the taxonomic key to species,synonyms,SEM microphotographs of shells and chamber arrays.By recent classification,currently 20 species representing 13 genera and 6 families(Canderinidae,Heterohelicoidae,Hedbergellidae,Higerigerinoidae,Globigerinoidae,and Globorotaloidae)identified from the planktonic material of the eastern Indian Ocean up to a depth of 200 m.Their distribution in water(0–200 m)also reports on the new range of expansion in the eastern Indian Ocean,with Dentigloborotalia anfracta,Hastigerina pelagica,Streptochilus globigerus,Globigerinella calida,Globigerinella adamsi,Orcadia reidelii,Tenuitella parkerae,Tenuitella compressa,reported for the first time in this study area.In general,only around 50 planktonic species are valid worldwide,more specifically the species,e.g.,H.pelagica,G.calida,G.adamsi,S.globigerus,O.riedeli,T.parkerae,T.compressa,which occur in the eastern Indian Ocean to fill the the paucity of the recent regional taxonomic literature and the problematic identification from the eastern Indian Ocean.This work aims to bridge this gap and help scientists,managers,educators and students to identify plankton foraminifers by using species notes and images.     

Features of the atmospheric boundary layer block for three versions of the WAM wind wave model

The estimated characteristics of the atmospheric boundary layer, obtained by the simulation of wind wave fields using three versions of the WAM numerical model are compared with the well-known empirical dependences of drag coefficient C _d on wind speed U ₁₀ and wave age A, as well as with the dependence of dimensionless roughness height z _n on inverse wave age u*/с р. Calculations carried out for several years in the areas of the Pacific and Indian oceans, based on the ERA-interim and CFSR wind reanalyses have shown good agreement between the model and empirical dependences C _d (U ₁₀) and C _d (A). The range of estimated variability for z _n (u*/с _р ) has been found to be significantly less than empirical. It has been also found that estimated values of wind speed U _10W (t) are overestimated from 5 to 10% in all versions of WAM models compared with the input wind reanalysis U _10R (t) at the moments of appearance maximum values of wind U _10R (t). The reasons for the established features of the WAM model and their dependence on the model version are discussed.     

Seasonal and Interannual Variations of Heat Fluxes in the Barents Sea Region

Seasonal and interannual variations in adjective heat fluxes in the ocean (dQ_oc) and the convergence of advective heat fluxes in the atmosphere (dQ_atm) in the Barents Sea region have been investigated over the period of 1993–2012 using the results of the MIT regional eddy-permitting model and ERA-Interim atmospheric reanalysis. Wavelet analysis and singular spectrum analysis are used to reveal concealed periodicities. Seasonal 2- to 4- and 5- to 8-year cycles are revealed in the dQ_oc and dQ_atm data. It is also found that seasonal variations in dQ_oc are primarily determined by the integrated volume fluxes through the western boundary of the Barents Sea, whereas the 20-year trend is determined by the temperature variation of the transported water. A cross-wavelet analysis of dQ_oc and dQ_atm in the Barents Sea region shows that the seasonal variations in dQ_oc and dQ_atm are nearly in-phase, while their interannual variations are out-of-phase. It is concluded that the basin of the Barents Sea plays an important role in maintaining the feedback mechanism (the Bjerknes compensation) of the ocean–atmosphere system in the Arctic region.     

Spatial Distribution and Seasonal Dynamics of the Chlorophyll <Emphasis Type="Italic">a</Emphasis> Concentration in the Sea of Azov Based on MERIS Images

The chlorophyll a concentration (Cchl a) in the Sea of Azov is estimated by the two-band NIR-red algorithm [34] from MERIS images for 2002–2012. The sea-truth spectrophotometric measurements and MERIS remote estimates of Cchl a are compared. The monthly average Cchl a values are mapped from MERIS data for its lifetime for the first time. The features of the spatiotemporal distribution of Cchl a are ascertained. Differences between the seasonal dynamics of Cchl a in the Sea of Azov according to the literature data and the dynamics derived from MERIS data are found, namely: the summer–autumn phytoplankton growth period is longer than the spring period and is characterized by higher Cchl a values throughout the water area.     

Intrusive layering of the Antarctic slope front

Cascading of cold Antarctic shelf water (ASW) initiates compensatory isopycnic upwelling of the warm Circumpolar Deep Water (CDW). The baroclinic/thermoclinic Antarctic slope front (ASF) is formed, and a mesoscale intrusive structure develops on the shelf edge and slope. Mesoscale processes when the ASF peaks are periodically accompanied by local baroclinic instability, which forms a smaller-scale intrusive structure. Therefore, the ASF is naturally subdivided into two layers according to the intrusion scales (vertical δН and horizontal L) and the horizontal parameters of the front (thermoclinity (T_L)_ρ and baroclinity γ_ρ). Analysis of ASF intrusive layering due to the baroclinic factor supports the following conclusion: the higher the (T_L)_ρ of the ASF, the greater the intrusion intensity |δθ| (temperature anomaly amplitude), while an increase in γ_ρ of the ASF leads to a decrease in intrusion scales δН and L. Frontal intrusions can be distinguished by a development degree. Regardless of the degree of development, all warm intrusions are characterized by vertical density stratification, while cold intrusions are characterized by density quasihomogeneity. According to field data, the ASF instability process is subdivided into four stages. When theASF is baroclinically unstable, the local baroclinic deformation radius Rd_L of the front is close in magnitude to the horizontal scale L of the intrusions that form, and their characteristic vertical scale δH is close to the typical vertical scale of front instability.     

珠江口溶解有机物的遥感动态监测

河口有色溶解有机物（colored dissolved organic matter,CDOM）的分布是各种物理-生物地球化学过程共同作用的结果。为实现河口高动态变化CDOM的监测,遥感是一种重要的手段。由珠江口四个不同季节的航次获得的实测数据,本文构建了一个遥感算法以反演CDOM在400 nm的吸收系数（a_CDOM （400））。该算法使用以波段反射率比值R_rs （667）/R_rs （443）和R_rs （748）/R_rs （412）为自变量。将构建的算法应用于2002-2014年的MODIS/Aqua数据,本文计算了珠江口不同季节的a_CDOM （400）气候态分布。CDOM的分布主要受珠江径流量和区域水下地形特征的影响。沿着垂直于水深梯度的断面,气候态a_CDOM （400）呈指数减少（y=ae^bx,b<0）,但不同季节差异很大。珠江口CDOM主要是河流淡水输运而来。其中,富里酸比例随盐度的增加而降低。基于构建的算法、CDOM保守混合方程和径流量,本文由MODIS/Aqua数据进一步估算了2002-2014年夏季和冬季珠江DOC的有效入海浓度和有效入海通量。珠江的有效入海浓度和有效入海通量都与流量存在正相关关系,且在夏季的相关性更明显,R2分别为0.698和0.9657。     

Seasonal and interannual variation in mesozooplankton community structure off Tongyeong,southeastern coast of Korea,from 2011 to 2014

Mesozooplankton community structure and environmental factors were monitored monthly at a fixed station off Tongyeong, southeastern coast of Korea, from 2011 to 2014 to better understand the variability of the mesozooplankton community in relation to changes in the marine environment. Total mesozooplankton density varied from 747 to 8,945 inds. m^-3 with peaks in summer. The surface water temperature (r = 0.338, p < 0.05) and chlorophyll-a (Chl-a) concentration (r = 0.505, p < 0.001) were parts of the factors that may have induced the mesozooplankton peaks in summer. Copepods accounted for 71% of total mesozooplankton. Total copepod density, particularly cyclopoid copepods, increased during the study period. Cumulative sum plots and anomalies of the cyclopoid copepod density revealed a change of the cyclopoid density from negative to positive in June 2013. A positive relationship between cyclopoid copepods and the Chl-a concentration (r = 0.327, p < 0.05) appeared to be one of the reasons for the increase in cyclopoids. Dominant mesozooplankton species such as Paracalanus parvus s.l., Oikopleura spp., Evadne tergestina, Cirripedia larvae, Corycaeus affinis, Calanus sinicus, and Oithona similis accounted for 60% of total mesozooplankton density. Based on cluster analysis of the mesozooplankton community by year, the seasonal distinction among groups was different in 2014 compared to other years. P. parvus s.l. and its copepodites contributed most in all groups in all four years. Our results suggest that the high Chl-a concentration since 2013 may have caused the changes in mesozooplankton community structure in the study area.     

Virio- and bacterioplankton in the estuary zone of the Ob River and adjacent regions of the Kara Sea shelf

The distribution of structural and functional characteristics of virioplankton in the north of the Ob River estuary and the adjacent Kara Sea shelf (between latitudes 71°44′44″ N and 73°45′24″ N) was studied with consideration of the spatial variations in the number (N _B) and productivity (P _B) of bacteria and water properties (temperature, salinity, density) by analyzing samples taken in September 2013. The number of plankton viruses (N _V), the occurrence of visible infected bacteria cells, virus-induced mortality of bacteria, and virioplankton production in the studied region varied within (214?2917) × 10³ particles/mL, 0.3?5.6% of NB, 2.2?64.4% of P _B, and (6?17248) × 10³ particles/(mL day), respectively. These parameters were the highest in water layers with a temperature of +7.3–7.5°C, salinity of 3.75?5.41 psu, and conventional density (στ) of 2.846?4.144. The number of bacterioplankton was (614?822) × 10³ cells/mL, and the N _V/N _B ratio was 1.1?4.5. A large amount of virus particles were attached to bacterial cells and suspended matter. The data testify to the considerable role of viruses in controlling the number and production of heterotrophic bacterioplankton in the interaction zone of river and sea waters.     

Variability of Particulate Organic Phosphorus in the Northwestern Part of the Black Sea

Based on long-term (1985–1995) monitoring data, the paper considers the peculiarities of seasonal variability in the spatial and vertical distribution of particulate organic phosphorus (Р_POM) in the surface layer and in the photosynthetic zone in the northwestern Black Sea. Regression equations, experimental data, and satellite observations for the chlorophyll a concentration allowed us to evaluate the seasonal longterm (1979–1995) variability in Р_POM in the surface layer and photosynthesis zone. The ratios of the concentrations of particulate organic carbon, nitrogen, phosphorus, and chlorophyll a are calculated and statistical estimates of seasonal changes in the Р_POM in the areas with different degrees of influence of river runoff and water of open seas are obtained. The consistency of intra-annual changes in the concentrations of Р_POM, chlorophyll a, and phytoplankton biomass is shown, which indicates the role of phytoplankton in the formation of Р_POM and in its intra- and interannual variability in the northwestern part of the sea. It is shown that long-term seasonal variations in Р_POM and related changes in the concentration of chlorophyll a depend on the variability of bulk river runoff, the extent of its abundance in the northwestern shelf, and regional hydrometeorological conditions.     

Phylogenetic relationship between symbionts of tubeworm <Emphasis Type="Italic">Lamellibrachia satsuma</Emphasis> and the sediment microbial community in Kagoshima Bay

Vestimentiferan tubeworms acquire their symbionts through horizontal transmission from the surrounding environment. In the present study, we constructed a 16S rRNA gene clone library to investigate the phylogenetic relationship between diverse microbes in the sediment and symbiotic bacteria in the trophosome of the tubeworm, Lamellibrachia satsuma, from Kagoshima Bay, Japan. Two symbiotic bacterial phylotypes belonging to the classes γ- and ε-Proteobacteria were found from this tubeworm trophosome. They were very closely related to the symbionts of several other marine invertebrates. The most predominant bacteria in the sediment were ε-Proteobacteria. A broad diversity of bacteria belonged to non-proteobacterial phyla such as Planctomycetes, Acidobacteria, and Chloroflexi was observed. The presence of sulfur oxidizers (i.e., ε-Proteobacteria and γ-Proteobacteria) and sulfur reducers (i.e., δ-Proteobacteria) may play a significant role in the sulfur cycle in these habitats and provide multiple sources of nutrition to the cold-seep communities. Closely related clones of ε-Proteobacteria symbiont in the species level and of γ-Proteobacteria symbiont in the genus level were found in the surrounding sediment. The similarity of symbiont clones of L. satsuma with other symbionts and free-living bacteria suggests the possibility of opportunistic symbiosis in ε-Proteobacteria and the co-evolution of γ-Proteobacteria having occurred after symbiosis with the tubeworms.