Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

Estimation of standing stock of chlorophylla and primary production from remote-sensed ocean color in the Oyashio region, the Western subarctic Pacific, during the spring bloom in 1997

In order to examine the applicability of remotely-sensed ocean color for the estimation of phytoplankton biomass and primary production in the Oyashio region, the western subarctic Pacific, vertical distributions of chlorophylla concentration and primary production were observed in April and May 1997. Spring bloom was observed in both April and May, and the surface concentration of chlorophylla exceeded 40 mg m⁻³. The relationship between the standing stocks of chlorophylla within the layer from the sea surface to one optical depth (0–1/k layer) and the surface chlorophylla concentration is expressed as a Michaelis-Menten equation. The mean ratio of the standing stock of chlorophylla in the euphotic layer to that in the 0–1/k layer was 4.41, this ratio did not significantly differ from 4.61 which was obtained at homogeneous distribution of chlorophylla within the euphotic layer. These facts suggest that the distribution of chlorophylla could be assumed to be homogeneous in the euphotic layer during the spring bloom. Results of primary production measurements by simulatedin situ method were compared with those by an algorithm with two variables; chlorphylla and non-spectral PAR. Daily primary production in the euphotic layer estimated by the algorithm varied in a range of 38–274% of that estimated by incubation, although the primary productions by the algorithm agreed with those by the incubation at a half of stations. Primary production within the euphotic layer calculated using simply the surface data was the same as that estimated using vertical distribution of chlorophylla. These results show that the primary production in the euphotic layer may be estimated from the remote sensed measurements during the spring bloom in the Oyashio region.     

Has the 1998 regime shift also occurred in the oceanographic conditions and lower trophic ecosystem of the Oyashio region?

To examine whether the regime shift in 1998 that has been variously reported to have occurred in the oceanographic conditions of the central and eastern North Pacific also occurred in the Oyashio region, western North Pacific, we compared data over the period 1990–2003. Oceanographic conditions were compared before 1997 with those after 1998, using the A-line dataset (1990–2003) obtained by the oceanographic surveys of the Hokkaido National Fisheries Research Institute, Fisheries Research Agency (HNFRI/FRA). Seasonal changes of the monthly-mean SST (as temperature in the surface layer) show a significant increase in spring after 1998. After 1998, the mean concentration of chlorophyll a at the surface was higher in spring than that before 1997. This was more remarkable in the main current of the Oyashio. These changes suggest that the spring phytoplankton bloom in the Oyashio region after 1998 was larger in magnitude and initiated earlier. Consumption of nutrients during the spring bloom and standing stock of netplankton also shows a distinct difference between the time period before 1997 and after 1998. These results support the occurrence of the regime shift around 1998 in the Oyashio region. The changes of hydrographical conditions accompanying with the 1998 regime shift are discussed. The hydrographic mechanism of enhancement of primary productivity during the spring phytoplankton bloom was not fully clarified, though. Results in this study may support the usefulness of the A-line dataset for analysis of long-term variability in the western subarctic Pacific.     

Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Changes in biomass structure and trophic status of the plankton communities in a highly dynamic ecosystem (Gulf of Venice,Northern Adriatic Sea)

The changes in the plankton biomass structure in relation to nutrient inputs were studied in the Gulf of Venice (Northern Adriatic Sea), an area characterized by a very marked trophic state variability. The investigation was carried out at two stations, in March, May and July 2005 and 2006, considering the whole water column. The size structure (from picoplankton to mesozooplankton) of both autotrophs and heterotrophs was analysed. Signals of diluted waters and nutrient inputs were more marked in 2005 than in 2006. In 2005, the total plankton biomass was almost double (87 ± 37 μg·C·l^?1) that in 2006 (44 ± 26 μg·C·l^?1). The variations were determined mainly by phytoplankton, with a 70% decrease, and a shift from a community dominated by microphytoplankton (49 ± 12%) in 2005 to one dominated by bacteria (43 ± 11%) in 2006 was observed. The relationship between the heterotrophic (H) and autotrophic (A) biomass indicated a rapid decline of the H/A ratio with increasing phytoplankton biomass. This study, although temporally limited, is consistent with the results reported for other marine environments and it seems to confirm the importance of nutrient inputs in structuring the biomass of plankton community.     

Downward carbon transport by diel vertical migration of the copepods Metridia pacifica and Metridia okhotensis in the Oyashio region of the western subarctic Pacific Ocean

Seasonal change in the downward carbon transport due to respiration and mortality through diel vertical migration (DVM) of the calanoid copepods Metridia pacifica and Metridia okhotensis was estimated in the Oyashio region, western subarctic Pacific during six cruises from June 2001 to June 2002. M. pacifica (C4, C5 and adult females) was an active migratory species throughout the year though its DVM amplitude varied among seasons and stages. The mean distribution depths of adult females during the daytime were positively related with the illumination level in the water column, being shallowest in April and deepest in January. M. okhotensis generally showed less-extensive migrations than M. pacifica. Therefore, together with their lower abundance, this species is considered to be a less-important mechanism of downward transport of carbon except for April when their DVM was more active and descended deeper than M. pacifica, which remained in the upper 150 m even during the daytime. The mean migrating biomass of the two Metridia species was 558 mg C m⁻² d⁻¹ and was high during summer to winter (263–1676 mg C m⁻² d⁻¹) and low during spring (59–63 mg C m⁻² d⁻¹). Total downward flux through DVM fluctuated between 1.0 and 20.0 mg C m⁻² d⁻¹ with an annual mean of 8.0 mg C m⁻² d⁻¹. Contribution of the respiratory flux was greater than the mortality flux and accounted for 64–98% of total migratory flux throughout the year except for January when contribution of both fluxes was equal. Overall the annual carbon transport by DVM of Metridia spp. was estimated as 3.0 g C m⁻² year⁻¹, corresponding to 15% of the annual total POC flux at 150 m at the study site, suggesting that DVM is a significant process for carbon export in the subarctic region as well as that in tropical and subtropical oceanic regions. Since DVM in M. pacifica is more active during the non-bloom season when the gravitational flux of particulate matter is low, this species plays an important role in driving the biological pump in the subarctic Pacific during summer to winter.     

Functional diversity of fish and invertebrates in coral and rocky reefs of the Eastern Tropical Pacific

To define the functional groups of fish and macroinvertebrates in the Eastern Tropical Pacific (ETP), visual censuses were performed in 18 areas of four biogeographic provinces: Cortés, Mexican, Panamic and Oceanic Islands. A total of 257 fish and macroinvertebrate species were recorded, and from them, 27 morpho‐functional groups (MFGs) were constructed on the basis of trophic level, maximum size, taxonomy and morphology. Biomass, richness, diversity and evenness of MFGs were calculated for each province and compared statistically; a regression analysis between taxonomic and functional diversity was conducted to observe the relationship between these two indicators. There were significant differences in all ecological indices (p < .002), highlighting the high biomass and richness of MFGs in the Cortés and Oceanic Islands provinces (>400 g/m²), associated with the influence of cold currents in the northernmost region and less fishing pressure in protected areas. A decreasing pattern of MFG richness towards the tropics was observed, which demonstrates that in the ETP, the relationship between habitat heterogeneity and species diversity has been translated into functional complexity. The Mexican province was the most functionally diverse (biomass well distributed in the MFG; H′ = 0.46 ± 0.009). Related to this, it is predicted that biomass is biased towards certain functional groups (i.e., large carnivores), which shows that the H′ index of the MFG is not a good indicator of the conservation status of ETP reefs. Finally, regression analysis suggests that functional diversity increases at low species diversity but eventually reaches an asymptote (almost all possible functions are represented).     

2003年夏季西北冰洋浮游动物群落结构和地理分布研究

根据西北冰洋43个调查站位的浮游动物种类组成和数量资料,分析了浮游动物的群落结构和地理分布特征,探讨了浮游动物群落与环境因子的关系.结果表明,在调查区域存在三种不同的浮游动物群落类型:楚科奇海台和加拿大海盆地区的高纬度深海群落;楚科奇海中部的陆架群落;阿拉斯加沿岸和楚科奇海北部的沿岸过渡群落.深海群落浮游动物数量较少,...     

Reef fish structure assemblages in oceanic islands of the eastern tropical Pacific: Revillagigedo Archipelago and Clipperton atoll

Revillagigedo Archipelago and Clipperton atoll are oceanic islands located in the eastern tropical Pacific. The composition and trophic structure of their fish assemblages have never been characterized in detail, and the aim of this study was therefore to analyse the community structure and assess the similarity of fish assemblages between these two areas. Underwater visual fish censuses were conducted using band transects at two depth levels in three islands of the Revillagigedo Archipelago (Socorro, San Benedicto, and Roca Partida) and at Clipperton atoll. We calculated ecological indices (density, richness, Shannon diversity, Pielou evenness, and taxonomic distinctiveness) for each transect. These were compared in order to determine quantitative differences among the islands and bathymetric levels. Qualitative analyses (ANOSIM, nMDS, and SIMPER) were also performed. Finally, we analysed the trophic structure of the assemblages and the functional diversity of the four studied sites. The results showed significant differences for all indices, except diversity (H′). Roca Partida was the site with greatest abundance and richness due to the presence of pelagic species in large numbers. No significant differences were found in functional diversity, suggesting a high degree of ecological redundancy. Lower‐level carnivores with mean trophic level values of between 3.3 and 3.8 dominated both areas. The ordination analysis did not show a clear difference in fish composition among the islands, although Clipperton atoll has a particular fauna characterized by endemic species (Thalassoma robertsoni and Stegastes baldwini). In conclusion, the fish communities of the studied areas seem quite similar, but their assemblage structure differs. In particular, Clipperton is differentiated by the influence of endemic species.     

Geographical and seasonal variations in biomass and estimated production rates of net zooplankton in Yatsushiro Bay, Japan

The biomass and production rate of net zooplankton were studied at eight stations in Yatsushiro Bay, Japan, monthly from May 2002 to April 2003. Based on environmental conditions, the bay was divided into three regions, viz. northern (average depth, salinity and chlorophyll a concentration: 11 m, 31.8 and 6.5 μg l⁻¹, respectively), central (30 m, 32.8 and 3.2 μg l⁻¹, respectively) and southern (43 m, 33.4 and 1.9 μg l⁻¹, respectively). Net zooplankton biomass was high in warm months and low in cold ones, with annual averages of 20.2, 38.8 and 16.4 mg C m⁻³ in the northern, central and southern regions, respectively. Copepods were the most important constituent (>ca. 70% of net zooplankton biomass) in all regions. The northern region was characterized by the dominance of Oithona spp. in summer and Acartia spp. in winter-spring. In the central region, Microsetella norvegica was most pronounced in summer-fall. In both central and southern regions, Calanus sinicus and Eucalanus spp. dominated in winter-spring and fall, respectively. The annual average net zooplankton secondary production rate was 4.4, 7.5 and 3.9 mg C m⁻³d⁻¹ in the northern, central and southern region, respectively. Combining the results from the present study with those from other collaborative works on microzooplankton allowed us to determine the trophic interactions in Yatsushiro Bay. If the secondary producers depend entirely on phytoplankton for food, their daily carbon requirement is equivalent to 12.5, 21.6 and 19.1% of the phytoplankton biomass in the respective regions.     

福建省主要海湾浮性鱼卵和仔稚鱼的种类组成与丰度分布

根据2007年8月至2009年10月期间在福建省主要海湾的调查资料,对其浮性鱼卵和仔稚鱼的种类组成与数量分布特点进行初步研究和探讨.结果表明,本调查海域共出现鱼卵和仔稚鱼68种,隶属于29科,主要是近海中上层鱼类、浅海底层鱼类和港湾河口型鱼类3个生态类型.各站鱼卵总平均丰度是497.5粒/网,仔稚鱼总平均丰度50.8个/网.优势种明显,占总量比例大;不同季节和不同海湾的种类数相差不大;数量平面分布呈现不同季节不同海湾差别较大,规律性不强,偶然性大等特点.     

Abundance, Biomass, Production and Trophic Roles of Micro- and Net-Zooplankton in Ise Bay, Central Japan, in Winter

We investigated the geographical variations in abundance and biomass of the major taxonomic groups of micro- and net-zooplankton along a transect through Ise Bay, central Japan, and neighboring Pacific Ocean in February 1995. The results were used to estimate their secondary and tertiary production rates and assess their trophic roles in this eutrophic embayment in winter. Ise Bay nourished a much higher biomass of both micro- and net-zooplankton (mean: 3.79 and 13.9 mg C m^–3, respectively) than the offshore area (mean: 0.76 and 4.47 mg C m^–3, respectively). In the bay, tintinnid ciliates, naked ciliates and copepod nauplii accounted for, on average, 69, 18 and 13% of the microzooplankton biomass, respectively. Of net-zooplankton biomass, copepods (i.e. Acartia, Calanus, Centropages, Microsetella and Paracalanus) formed the majority (mean: 63%). Average secondary production rates of micro- and net-zooplankton in the bay were 1.19 and 1.87 mg C m^–3d^–1 (or 23.1 and 36.4 mg C m^–2d^–1), respectively, and average tertiary production rate of net-zooplankton was 0.75 mg C m^–3d^–1 (or 14.6 mg C m^–2d^–1). Available data approximated average phytoplankton primary production rate as 1000 mg C m^–2d^–1 during our study period. The transfer efficiency from primary production to zooplankton secondary production was 6.0%, and the efficiency from secondary production to tertiary production was 25%. The amount of food required to support the zooplankton secondary production corresponded to 18% of the phytoplankton primary production or only 1.7% of the phytoplankton biomass, demonstrating that the grazing impact of herbivorous zooplankton was minor in Ise Bay in winter.     

基于延绳钓观察员数据的热带太平洋中东部食物网拓扑结构分析

热带太平洋是金枪鱼类、旗鱼类等渔业的主要作业海域,但目前针对此海域食物网的研究仍十分缺乏。本文根据2017年6月-11月中国金枪鱼科学观察员在热带太平洋中东部海域（13°09''N -34°27''S, 100°03''-176°17''W）采集的鱼类样品,结合胃含物及其他分析方法以35个物种间的摄食关系为基础对此海域的食物网结构进行了分析。拓扑学指标（节点强度,D;中心性指数,BC和CC;拓扑重要性指数,TI¹和TI³;关键性指数,K、K_t和K_b）和Key-Player算法（KPP-1和KPP-2）用来筛选关键种,并结合体长数据建立了基于关键种的简化食物网。肯德尔相关性分析和聚类分析结果表明,不同拓扑学指标筛选出的关键种是一致的。大多数关键种与太平洋其他海域的研究结果是相同的,如剑鱼（Xiphias gladius）、长鳍金枪鱼（Thunnus alalunga）、小吻四鳍旗鱼（Tetrapturus angustirostris）头足类和鲭属。移除5个关键种后,食物网将被离散为许多互不连接的部分（F=0.632,F^D=0.795,R^D=0.957）,表明这些物种在维持食物网结构和稳定性中的必要性。物种体长是本研究构建简化食物网时的决定性因素。研究结果有助于提高对热带太平洋食物网结构的认识,同时为进一步的生态系统动力学研究提供科学依据。     

Responses of diatoms to iron-enrichment (SEEDS) in the western subarctic Pacific, temporal and spatial comparisons

Phytoplankton species composition was analyzed inside and outside of the iron-enriched patch during the SEEDS experiment. Before the iron-enrichment, the phytoplankton community consisted of similar proportions of pico-, nano- and micro-sized phytoplankton, and the micro-phytoplankton was dominated by the pennate diatom Pseudo-nitzschia turgidula. Although all the diatoms, except the nano-sized Fragilariopsis sp., increased during the two weeks of the observation period, the flora in the patch dramatically changed with the increase of phytoplankton biomass to a centric diatom-dominated community. Neritic diatoms, especially Chaetoceros debilis, showed higher growth rates than other diatoms, without any delay in the initiation of growth after the enrichment, and accounted for 90% of the micro-phytoplankton after day 9. In contrast, the oceanic diatoms showed distinct delays in the initiation of growth. We conclude that the responses of the diatoms to the manipulation of iron concentration were different by species, and the fast and intensive response of the phytoplankton to iron-enrichment resulted from the presence of a small amount of neritic diatoms at the study site. The important factors that determine the dominant species in the bloom are the potential growth rates under an iron-replete condition and the growth lag. Abundant species in the patch are widely distributed in the North Pacific and their relative contributions in the Oyashio area and at Stn KNOT are high from spring to summer. However, a characteristic difference of species composition between the SEEDS bloom and natural blooms was the lack of Thalassiosira and Coscinodiscus species in the patch, which usually account for a major part of the phytoplankton community under blooming conditions in the western North Pacific.     

胶州湾大型底栖动物的丰度、生物量和生产量研究

为了研究胶州湾大型底栖动物的丰度、生物量和生产量,于2002年3月、6月、8月和12月,在胶州湾北部软底区、大沽河口、黄岛养殖区及养殖区邻域选取4个站位进行采样,对大型底栖动物进行了定量研究。共采到大型底栖动物138种,总平均丰度、平均生物量(湿质量)和年生产量(有机碳)分别为1 719个/m2,27 g/m2,2.2 g/(m2.a),初步估算,胶州湾大型底栖动物的总次级生产量为2.8万t/a。与渤海和南黄海大型底栖动物的丰度和生物量比较,丰度和生物量均低于这两个海域,但是胶州湾大型底栖动物的总次级生产量高于渤海。本研究对于了解胶州湾大型底栖动物现状及湾内养殖对大型底栖动物的影响具有重要的意义。