Haptophyte communities along the Kuroshio current reveal their geographical sources and ecological traits.

Haptophytes are one of the most ecologically successful phytoplankton groups in the modern ocean and tend to maintain balanced and stable communities across various environments. However, little is known about the mechanisms that enable community stability and ecological success. To reveal the community characteristics and interactions among haptophytes, we conducted comprehensive observations from the upstream to downstream regions of the Kuroshio Current. Haptophyte abundance and taxonomy were assessed using quantitative polymerase chain reaction and metabarcoding of 18S rRNA sequences, respectively. The haptophyte community structure changed abruptly at sites on the shelf-slope of the East China Sea, indicating the strong influence of shelf waters with high phytoplankton biomass on downstream communities. Correlation network analysis combined with the phylogeny suggested that haptophytes can coexist with their close relatives, possibly owing to their nutritional flexibility, thereby escaping from resource competition. Consistently, some noncalcifying haptophyte genera with high mixotrophic capacities such as Chrysochromulina constituted a major component of the co-occurrence network, whereas coccolithophores such as Emiliania/Gephyrocapsa were rarely observed. Our study findings suggest that noncalcifying haptophytes play crucial roles in community diversity and stability, and in sustaining the food web structure in the Kuroshio ecosystems.

Evaluation of origin-depended nitrogen input through atmospheric deposition and its effect on primary production in coastal areas of western Kyusyu, Japan.

Long term monitoring of atmospheric wet and dry depositions and associated nutrients fluxes was conducted on the coast of Japan facing the East China Sea continuously for 1 year and 2 months, with the origin of air mass investigated based on isotope analyses (Sr, Nd, and NO3). During the same period, intensive observations of ocean conditions and the chemical composition of sinking particles collected using sediment traps were conducted to investigate the effects of atmospheric deposition-derived nutrients on phytoplankton blooms. Dry-deposition-derived nutrient inputs to the surface ocean were larger during autumn to spring than in summer due to the effect of continental air mass occasionally carrying Asian dust (yellow sand). However, these nutrients fluxes were limited (1.1-1.5 mg-N m-2 day-1 on average) and didn't appear to cause phytoplankton blooms through the year. Although average dissolved inorganic nitrogen (DIN) concentrations in rainwater were lower in oceanic air masses compared to continental air masses, wet-deposition-derived nutrient inputs to the surface ocean on rainy days during the summer (26.0 mg-N m-2 day-1 on average) were large due to higher precipitation from oceanic air masses. Wet-deposition-derived nutrients significantly increased nutrient concentrations in the surface ocean and seemed to cause phytoplankton blooms in the warm rainy season when nutrients in the surface were depleted due to increased stratification. The increase in phytoplankton biomass was reflected in increased particle sinking into the bottom layer, as well as changing chemical characteristics. The supply of flesh phytoplankton-derived labile organic matter into the bottom layer could be expected to promote rapid bacterial decomposition and contribute to the formation of hypoxic water masses in early summer when the ocean was strongly stratified. Atmospheric deposition-derived nutrients in East Asia will have important impacts on not only the oligotrophic outer ocean but also surrounding coastal areas in the warm rainy season.

Potential oxygen consumption and community composition of sediment bacteria in a seasonally hypoxic enclosed bay.

The dynamics of potential oxygen consumption at the sediment surface in a seasonally hypoxic bay were monitored monthly by applying a tetrazolium dye (2-(4-iodophenyl)-3-(4-nitrophenyl)-5-phenyl-2H-tetrazolium chloride [INT]) reduction assay to intact sediment core samples for two consecutive years (2012-2013). Based on the empirically determined correlation between INT reduction (INT-formazan formation) and actual oxygen consumption of sediment samples, we inferred the relative contribution of biological and non-biological (chemical) processes to the potential whole oxygen consumption in the collected sediment samples. It was demonstrated that both potentials consistently increased and reached a maximum during summer hypoxia in each year. For samples collected in 2012, amplicon sequence variants (ASVs) of the bacterial 16S rRNA genes derived from the sediment surface revealed a sharp increase in the relative abundance of sulfate reducing bacteria toward hypoxia. In addition, a notable shift in other bacterial compositions was observed before and after the INT assay incubation. It was Arcobacter (Arcobacteraceae, Campylobacteria), a putative sulfur-oxidizing bacterial genus, that increased markedly during the assay period in the summer samples. These findings have implications not only for members of Delta- and Gammaproteobacteria that are consistently responsible for the consumption of dissolved oxygen (DO) year-round in the sediment, but also for those that might grow rapidly in response to episodic DO supply on the sediment surface during midst of seasonal hypoxia.

Nitrogen burden from atmospheric deposition in East Asian oceans in 2010 based on high-resolution regional numerical modeling.

East Asian oceans are possibly affected by a high nitrogen (N) burden because of the intense anthropogenic emissions in this region. Based on high-resolution regional chemical transport modeling with horizontal grid scales of 36 and 12 km, we investigated the N burden into East Asian oceans via atmospheric deposition in 2010. We found a high N burden of 2-9 kg N ha-1 yr-1 over the Yellow Sea, East China Sea (ECS), and Sea of Japan. Emissions over East Asia were dominated by ammonia (NH3) over land and nitrogen oxides (NOx) over oceans, and N deposition was dominated by reduced N over most land and open ocean, whereas it was dominated by oxidized N over marginal seas and desert areas. The verified numerical modeling identified that the following processes were quantitatively important over East Asian oceans: the dry deposition of nitric acid (HNO3), NH3, and coarse-mode (aerodynamic diameter greater than 2.5 µm) NO3-, and wet deposition of fine-mode (aerodynamic diameter less than 2.5 µm) NO3- and NH4+. The relative importance of the dry deposition of coarse-mode NO3- was higher over open ocean. The estimated N deposition to the whole ECS was 390 Gg N yr-1; this is comparable to the discharge from the Yangtze River to the ECS, indicating the significant contribution of atmospheric deposition. Based on the high-resolution modeling over the ECS, a tendency of high deposition in the western ECS and low deposition in the eastern ECS was found, and a variety of deposition processes were estimated. The dry deposition of coarse-mode NO3- and wet deposition of fine-mode NH4+ were the main factors, and the wet deposition of fine-mode NO3- over the northeastern ECS and wet deposition of coarse-mode NO3- over the southeastern ECS were also found to be significant processes determining N deposition over the ECS.

Diurnal and seasonal variation of particle and dissolved organic matter release by the coral Acropora tenuis.

Release rates of particulate organic carbon and nitrogen (POC and PON) and dissolved organic carbon (DOC) from the scleractinian coral Acropora tenuis were measured during the day and night in summer and winter seasons. Physiological parameters including calcification, photosynthesis and respiration rates were also measured simultaneously. The release rate of both POC and DOC was significantly higher in summer compared to winter and higher during the day compared to the night. The daily release rate of total organic carbon (POC + DOC) was 1,094 and 219 µmol C cm-2 d-1 for summer and winter, respectively, being 4.9 times higher in summer. The POC:PON ratios of the particulate organic matter released during daytime in both seasons (summer: 12.8 ± 5.7, winter: 12.0 ± 4.1) were significantly higher than those during nighttime (summer: 6.1 ± 2.5, winter: 2.2 ± 1.8). The DOC:POC ratio was 0.5 ± 0.03 during summer and 0.32 ± 0.98 during winter, suggesting higher mucus release in particulate form. Daily net production was estimated to be 199 and 158 µg C cm-2d-1 for summer and winter, respectively, with the amount of carbon released as mucus accounting for 6.5% and 1.6% of the net carbon fixation, respectively. The study reveals diurnal and seasonal changes in the quantity and quality of mucus released from this coral species. Since coral mucus is used as a food source by reef macro-organisms, and can also serve as an energy source for micro-organisms, the observed changes in mucus release rates are expected to influence the seasonal dynamics of organic carbon and nitrogen cycling over coral reefs.

Dynamics of Sulfate-Reducing Bacteria Community Structure in Surface Sediment of a Seasonally Hypoxic Enclosed Bay.

We herein report on the dynamics of a sulfate-reducing bacteria (SRB) community structure in the surface sediment of a seasonally hypoxic enclosed bay for two consecutive years (2012 and 2013). The uppermost (0-5 mm) and subsurface (5-10 mm) layers of sediment were examined with a terminal-restriction fragment length polymorphism (T-RFLP) analysis based on the dissimilatory sulfite reductase (dsrA) gene. The SRB community significantly differed between the two sediment layers over the sampling period. This difference was mainly attributed to operational taxonomic units (OTUs) that were unique to either of the sediment layers. However, nearly 70% of total OTUs were shared between the two layers, with a few predominating. Therefore, no significant shift was observed in the SRB community structure under varying dissolved oxygen (DO) conditions in bottom water overlying the sediment surface. An additional analysis of 16S rRNA gene amplicon sequences, conducted for three uppermost sediment samples (July, August, and September in 2012), revealed that Desulfococcus, a member of SRB with high tolerance to oxygen, was the predominant Deltaproteobacteria across the uppermost sediment samples. Based on the predominance of shared OTUs across the SRB community in the sediment (0-10 mm) regardless of bottom-water DO, some SRB that are physiologically tolerant of a wide range of DO conditions may have dominated and masked changes in responsive SRB to DO concentrations. These results suggest that the SRB community structure in the enclosed bay became stable under repeated cycles of seasonal hypoxia, but may be compromised if the severity of hypoxia increases in the future.

Effects of bottom-water hypoxia on sediment bacterial community composition in a seasonally hypoxic enclosed bay (Omura Bay, West Kyushu, Japan).

The bacterial community strongly drives carbon and other biogeochemical cycles in marine sediment. However, little is known about the impact of dissolved oxygen (DO) availability on bacterial community composition. To fill this gap, we examined diversity, richness and structure of the bacterial population for three consecutive years (2011-2013) in the uppermost (0-5 and 0-7 mm depth) and the subsurface layers (5-10 and 7-14 mm depth) of Omura Bay, Kyushu, Japan, a seasonally hypoxic enclosed bay. Automated ribosomal intergenic spacer analysis revealed a unimodal pattern of diversity indices with DO, peaking at the suboxic (11 µM O2) condition. Shifts in the bacterial communities were also evident in response to the availability of DO. Changes in the operational taxonomic units (OTUs) that were less abundant accounted for a large part of the community dissimilarity. It was further demonstrated that the relative abundance of OTUs affiliated with Gammaproteobacteria was correlated positively with DO, while that with Deltaproteobacteria was inversely correlated with DO. These results strongly suggest that DO availability of bottom water plays a fundamental role in shaping the bacterial community in sediment surfaces of shallow coastal areas.

Highly heterogeneous diazotroph communities in the Kuroshio Current and the Tokara Strait, Japan.

In this study, we used 454-pyrosequencing to report the highly diverse diazotroph communities in the Kuroshio and its adjacent waters along a transect across the Tokara Strait, Japan. Terrestrial input from the islands resulted in a highly heterogeneous diazotroph community within a relatively small geographic region, which was presumably caused by the remarkably different responses of UCYN-A2, UCYN-C and Trichodesmium to the steep environmental gradient. On the other hand, most major cyanobacterial OTUs found in this study were also detected in an unpublished dataset from the upstream Kuroshio, which suggests transportation of diazotrophs by the Kuroshio in large geographic scale. A significant amount of UCYN-C was found in the Kuroshio and offshore stations, suggesting the importance of this potentially overlooked group in the western North Pacific Ocean (WNPO). Moreover, a novel sublineage of UCYN-B was defined, which was predominant in an oligotrophic water sample; and it was also found to be widely distributed in oceanic waters. In addition, the apparent increase in relative abundance of UCYN-A2 from offshore to near-shore water provides evidence for the earlier and under-debating view that UCYN-A2 prefers coastal conditions. Our report provides new knowledge for understanding the phylogeny and ecology of unicellular cyanobacterial diazotrophs in WNPO.

Evidence of chronic anthropogenic nutrient within coastal lagoon reefs adjacent to urban and tourism centers, Kenya: A stable isotope approach.

The source of anthropogenic nutrient and its spatial extent in three fringing reefs with differing human population gradients in Kenya were investigated using stable isotope approaches. Nutrient concentrations and nitrate-δ15N in seepage water indicated that population density and tourism contributed greatly to the extent of nutrient loading to adjacent reefs. Although water-column nutrient analyses did not show any significant difference among the reefs, higher δ15N and N contents in macrophytes showed terrestrial nutrients affected primary producers in onshore areas in Nyali and Bamburi reefs, but were mitigated by offshore water intrusion especially at Nyali. On the offshore reef flat, where the same species of macroalgae were not available, complementary use of δ15N in sedimentary organic matter suggested inputs of nutrients originated from the urban city of Mombasa. If population increases in the future, nutrient conditions in the shallower reef, Vipingo, may be dramatically degraded due to lower water exchange ratio.

Phosphorus as a driver of nitrogen limitation and sustained eutrophic conditions in Bolinao and Anda, Philippines, a mariculture-impacted tropical coastal area.

The dynamics of nitrogen (N) and phosphorus (P) was studied in mariculture areas around Bolinao and Anda, Philippines to examine its possible link to recurring algal blooms, hypoxia and fish kills. They occur despite regulation on number of fish farm structures in Bolinao to improve water quality after 2002, following a massive fish kill in the area. Based on spatiotemporal surveys, coastal waters remained eutrophic a decade after imposing regulation, primarily due to decomposition of uneaten and undigested feeds, and fish excretions. Relative to Redfield ratio (16), these materials are enriched in P, resulting in low N/P ratios (~6.6) of regenerated nutrients. Dissolved inorganic P (DIP) in the water reached 4µM during the dry season, likely exacerbated by increase in fish farm structures in Anda. DIP enrichment created an N-limited condition that is highly susceptible to sporadic algal blooms whenever N is supplied from freshwater during the wet season.

Groundwater dynamics of Fongafale Islet, Funafuti Atoll, Tuvalu.

Geoelectric and hydrologic surveys during spring tides revealed the spatiotemporal distribution of groundwater quality produced by tidal forcing in Fongafale Islet, Funafuti Atoll, Tuvalu. The observed low resistivity showed that saline water largely immersed the surficial Holocene aquifer, indicating that there is no thick freshwater lens in Fongafale Islet, unlike in other atoll islands of comparable size. Half of the islet was constructed by reclaiming the original swamp with porous, highly permeable coral blocks; this reclaimed area should not be considered as part of the islet width for calculation of the expected thickness of the freshwater lens. The degree of aquifer salinization depends on the topographic characteristics and the hydrologic controls on the inland propagation of the tidal forcing. Large changes in bulk resistivity and the electrical conductivity of groundwater from wells indicate that periodic salinization in phase with the semidiurnal tides was occurring widely, especially in areas at lower elevation than the high-tide level and in reclaimed areas with high permeability. Thin sheets of nearly fresh and brackish water were observed in the surficial aquifer in areas above the high-tide level and in taro swamps, respectively. The thinness of the brackish and freshwater sheets suggests that the taro swamps and the fresh groundwater resources of the islet are highly vulnerable to salinization from anticipated sea-level rise. An understanding of the inherent geologic and topographic features of an atoll is necessary to evaluate the groundwater resources of the atoll and assess the vulnerability of its water resources to climate change.

Human impacts on large benthic foraminifers near a densely populated area of Majuro Atoll, Marshall Islands.

Human impacts on sand-producing, large benthic foraminifers were investigated on ocean reef flats at the northeast Majuro Atoll, Marshall Islands, along a human population gradient. The densities of dominant foraminifers Calcarina and Amphistegina declined with distance from densely populated islands. Macrophyte composition on ocean reef flats differed between locations near sparsely or densely populated islands. Nutrient concentrations in reef-flat seawater and groundwater were high near or on densely populated islands. delta(15)N values in macroalgal tissues indicated that macroalgae in nearshore lagoons assimilate wastewater-derived nitrogen, whereas those on nearshore ocean reef flats assimilate nitrogen from other sources. These results suggest that increases in the human population result in high nutrient loading in groundwater and possibly into nearshore waters. High nutrient inputs into ambient seawater may have both direct and indirect negative effects on sand-producing foraminifers through habitat changes and/or the collapse of algal symbiosis.

Physical and topographic factors affecting suspended particulate matter composition in a shallow tropical estuary.

To better understand topography-dependent characteristics of suspended particulate matter (SPM) in a tropical shallow estuary during dry season, the physical factors causing an increase in SPM and the sources of SPM were investigated at the western coast of Thailand. Single and multiple regression analyses using physical parameters as independent variables indicated that periodic tidal current-driven resuspension, episodic wind-driven resuspension, and river-borne inputs were the most important factors controlling SPM increases in areas surrounded by sand bars, areas directly facing the ocean, and areas close to the river mouth, respectively. The assessment of the origin of increased SPM over the background levels estimated from the chemical signatures (delta(13)C and C/N ratio) at all investigated locations and for each event responsible for an increase in SPM confirmed the results of the multiple regression analyses. The results suggested that specific characteristics of SPM at each location were highly contributed by sedimentary materials and could be rather consistent through the season under similar weather conditions.

Erratum to "Effects of intensive urbanization on the intrusion of shallow groundwater into deep groundwater: examples from Bangkok and Jakarta".

Asian megacities have severe pollution problems in both coastal and urban areas. In addition, the groundwater potential has decreased and land subsidence has occurred because of intensive groundwater pumping in urban areas. To prevent the adverse effects of urbanization on groundwater quality, it is necessary to confirm the changes in groundwater flow and contaminant transport caused by urbanization. We examined the effects of urbanization on contaminant transport in groundwater. The research areas were located around Bangkok, Thailand, and Jakarta, Indonesia, cities with populations of approximately 8 and 12 million, respectively. Each metropolitan city is located on a river delta and is adjacent to a bay. We measured the water level and collected water samples at boreholes at multiple depths (100 to 200 m) in 2004 and 2006 in Bangkok and Jakarta, respectively. The current hydraulic potential is below sea level in both cities because of prior excess abstraction of groundwater. As a result, the direction of groundwater flow is now downward in the coastal area. The Cl- concentration and delta18O distributions in groundwater suggest that the decline in hydraulic potential has caused the intrusion of seawater and shallow groundwater into deep groundwater. Concentrations of Mn and NO3--N in groundwater suggest the intrusion of these contaminants from shallow to deep aquifers with downward groundwater flow and implies an accumulation of contaminants in deep aquifers. Therefore, it is important to recognize the possibility of future contaminant transport with the discharge of deep groundwater into the sea after the recovery of groundwater potential in the coastal areas.

Erratum to "Sources of nitrate and ammonium contamination in groundwater under developing Asian megacities".

The status of nitrate (NO3-), nitrite (NO2-) and ammonium (NH4 +) contamination in the water systems, and the mechanisms controlling their sources, pathways, and distributions were investigated for the Southeast Asian cities of Metro Manila, Bangkok, and Jakarta. GIS-based monitoring and dual isotope approach (nitrate delta15N and delta18O) suggested that human waste via severe sewer leakage was the major source of nutrient contaminants in Metro Manila and Jakarta urban areas. Furthermore, the characteristics of the nutrient contamination differed depending on the agricultural land use pattern in the suburban areas: high nitrate contamination was observed in Jakarta (dry fields), and relatively lower nutrients consisting mainly of ammonium were detected in Bangkok (paddy fields). The exponential increase in NO3--delta15N along with the NO3- reduction and clear delta18O/delta15N slopes of NO3- (approximately 0.5) indicated the occurrence of denitrification. An anoxic subsurface system associated with the natural geological setting (e.g., the old tidal plain at Bangkok) and artificial pavement coverage served to buffer NO3- contamination via active denitrification and reduced nitrification. Our results showed that NO3- and NH4+ contamination of the aquifers in Metro Manila, Bangkok, and Jakarta was not excessive, suggesting low risk of drinking groundwater to human health, at present. However, the increased nitrogen load and increased per capita gross domestic product (GDP) in these developing cities may increase this contamination in the very near future. Continuous monitoring and management of the groundwater system is needed to minimize groundwater pollution in these areas, and this information should be shared among adjacent countries with similar geographic and cultural settings.

Underground sources of nutrient contamination to surface waters in Bangkok, Thailand.

Radon-222 is very concentrated in groundwater relative to surface waters and thus serves as an effective groundwater discharge tracer. We observed spikes in radon data from an earlier (2004) survey of the Chao Phraya River that appeared to correspond to locations where major canals ("klongs") enter the river. We returned in 2006 and conducted more detailed surveys along some of the main klongs on the western (Thonburi) side of the Chao Phraya to evaluate this possibility. Our results show that both radon and conductivity are enriched in some areas of the klongs with 3 apparent "end-members," two of which are likely related to groundwater seepage. Furthermore, nutrient analyses conducted during a time-series experiment at a site of suspected high discharge (Wat Intharam, Klong Bangkok Yai) showed that dissolved inorganic nitrogen (DIN) and phosphate correlated significantly to the groundwater tracer, radon. Rough estimates of the nutrient fluxes in this area are orders of magnitude higher than those measured in coastal settings and may represent a significant fraction of the riverine flux. It thus appears very likely that seepage of shallow groundwater is an important pathway for nutrient contamination of the klongs, and thus to the river, and ultimately to the Gulf of Thailand.

Effects of intensive urbanization on the intrusion of shallow groundwater into deep groundwater: examples from Bangkok and Jakarta.

Asian megacities have severe pollution problems in both coastal and urban areas. In addition, the groundwater potential has decreased and land subsidence has occurred because of intensive groundwater pumping in urban areas. To prevent the adverse effects of urbanization on groundwater quality, it is necessary to confirm the changes in groundwater flow and contaminant transport caused by urbanization. We examined the effects of urbanization on contaminant transport in groundwater. The research areas were located around Bangkok, Thailand, and Jakarta, Indonesia, cities with populations of approximately 8 and 12 million, respectively. Each metropolitan city is located on a river delta and is adjacent to a bay. We measured the water level and collected water samples at boreholes at multiple depths (100 to 200 m) in 2004 and 2006 in Bangkok and Jakarta, respectively. The current hydraulic potential is below sea level in both cities because of prior excess abstraction of groundwater. As a result, the direction of groundwater flow is now downward in the coastal area. The Cl(-) concentration and delta(18)O distributions in groundwater suggest that the decline in hydraulic potential has caused the intrusion of seawater and shallow groundwater into deep groundwater. Concentrations of Mn and NO3(-)-N in groundwater suggest the intrusion of these contaminants from shallow to deep aquifers with downward groundwater flow and implies an accumulation of contaminants in deep aquifers. Therefore, it is important to recognize the possibility of future contaminant transport with the discharge of deep groundwater into the sea after the recovery of groundwater potential in the coastal areas.

Sources of nitrate and ammonium contamination in groundwater under developing Asian megacities.

The status of nitrate (NO(3)(-)), nitrite (NO(2)(-)) and ammonium (NH(4)(+)) contamination in the water systems, and the mechanisms controlling their sources, pathways, and distributions were investigated for the Southeast Asian cities of Metro Manila, Bangkok, and Jakarta. GIS-based monitoring and dual isotope approach (nitrate delta(15)N and delta(18)O) suggested that human waste via severe sewer leakage was the major source of nutrient contaminants in Metro Manila and Jakarta urban areas. Furthermore, the characteristics of the nutrient contamination differed depending on the agricultural land use pattern in the suburban areas: high nitrate contamination was observed in Jakarta (dry fields), and relatively lower nutrients consisting mainly of ammonium were detected in Bangkok (paddy fields). The exponential increase in NO(3)(-)-delta(15)N along with the NO(3)(-) reduction and clear delta(18)O/delta(15)N slopes of NO(3)(-) ( approximately 0.5) indicated the occurrence of denitrification. An anoxic subsurface system associated with the natural geological setting (e.g., the old tidal plain at Bangkok) and artificial pavement coverage served to buffer NO(3)(-) contamination via active denitrification and reduced nitrification. Our results showed that NO(3)(-) and NH(4)(+) contamination of the aquifers in Metro Manila, Bangkok, and Jakarta was not excessive, suggesting low risk of drinking groundwater to human health, at present. However, the increased nitrogen load and increased per capita gross domestic product (GDP) in these developing cities may increase this contamination in the very near future. Continuous monitoring and management of the groundwater system is needed to minimize groundwater pollution in these areas, and this information should be shared among adjacent countries with similar geographic and cultural settings.