Long-term variability in zooplankton biomass in the subarctic Pacific Ocean

Zooplankton collections from the subarctic Pacific were analyzed from two periods (1956–1962 and 1980–1989). In this report, we document: 1) a positive correlation between the intensity of winter winds and subsequent summer zooplankton biomass in the subarctic gyre within these periods; and 2) a doubling of zooplankton biomass and a similar increase in pelagic fish and squid abundance between these two periods of time. Some possible explanations for these changes are considered.     

Interdecadal variations of plankton biomass and physical environment in the North Pacific

In the central and western subarctic Pacific, zooplankton biomass and chlorophyll concentrations during the mid 1960s to mid 1970s were a few times higher than in the preceding and following decades, corresponding to higher values of the atmospheric Northern Hemisphere Zonal Index (NHZI). In the Alaskan Gyre, however, it was reported that biomass of zooplankton and nekton doubled after the atmospheric regime shift in the mid 1970s. In the subtropical North Pacific, chlorophyll a concentration decreased drastically after 1980, although a decrease of zooplankton biomass was clearly seen only in the northern part of the subtropical gyre. Chlorophyll concentration in the central subarctic Pacific and zooplankton biomass in the Oyashio have been decreasing since the early 1980s. Additionally, chlorophyll concentration in the western subarctic Pacific and eastern Bering Sea, and zooplankton biomass in the central subarctic Pacific and eastern Bering Sea have also been decreasing since the late 1980s. In these regime-shift situations, there is a general tendency for intensification of wind speed or de-stratification to cause plankton biomass to decrease in regions where the upper mixed layer is deep, such as the western subarctic and north-western subtropical water, whereas in relatively stratified areas, such as in the eastern subarctic and south-western subtropical water, the effect is an increase of plankton biomass.     

Change in chum salmon (Oncorhynchus keta) stomach contents associated with fluctuation of pink salmon (O. gorbuscha) abundance in the central subarctic Pacific and Bering Sea

The abundance and stomach contents of salmonids (Oncorhynchus spp.) and the biomass of prey organisms were examined in the central subarctic Pacific and Bering Sea in the summer of 1991 and 1992. Salmonids were caught by surface longline using the same level of fishing effort. Chum (O. keta) and pink (O. gorbuscha) salmon were the predominant species, representing 44% and 36% sof the total catch (n = 1275) in 1991. In 1992, chum salmon composed 85% of the total catch (n = 603), but the catch of pink salmon decreased to 1% of the total catch due to the odd/even year fluctuation of Asian pink salmon abundance in the study area. It was found that chum salmon changed their dominant diet from gelatinous zooplankton (pteropods, appendicular-ians, jellyfishes, chaetognaths, polychaetes and unidentified materials) in 1991, when pink salmon were abundant, to a diet of crustaceans (euphausiids, cope-pods, amphipods, ostracods, mysids and decapods) in 1992, when pink salmon were less abundant. Local crustacean biomass (wet weight; mg m^-3) had significant negative correlation with the CPUE (catch number per 30 hachi) of pink salmon in 1991 (r = -0.586; P = 0.026) and that of chum salmon in 1992 (r =–0.616; P = 0.014). There may be a limitation in the available prey resource for production of salmonids.     

The zooplankton biomass in the Sea of Japan

We examined seasonal, annual variation and horizontal distribution of zooplankton in the Sea of Japan from 1966 to 1990. Zooplankton was most abundant in the spring. The spring maximum appeared in February–March and in April–May in the southern and eastern parts of the study areas, respectively. In the summer and autumn, a secondary peak was most conspicuous in the eastern part. The difference between the estimated biomass at night and day was large in the spring and small in summer and autumn. The biomass in the offshore southern area peaked about every 3 years between 1966 and 1983, and increased abruptly in 1990. The density in the area north of 39°N or 40°N was high. Total biomass estimated in the upper 150 m layer in the Sea of Japan (10⁶ km²) was 9.5 × 10⁶ t in the daytime and 16.6 × 10⁶ t at night.     

Variability in timing and magnitude of spring bloom in the Oyashio region, the western subarctic Pacific off Hokkaido, Japan

The spring bloom of phytoplankton is a well-established, regular, seasonal event in the western subarctic Pacific and is considered one of the most important conditions of massive production of pelagic fishes. A series of 12 cruises was conducted from 1990 to 1992 to examine the timing and magnitude of the spring phytoplankton bloom in the Oyashio region, the western subarctic Pacific off Hokkaido, Japan. An interannual variability in the bloom events was also analysed. On the basis of hydrographical characteristics, the study area was divided into three water masses: the Oyashio Water Mass, the Mixed Water Mass, and the Coastal Water Mass. Spring blooms were observed first in April in the Oyashio and the Coastal Water Masses, and continued to May in 1991 and 1992. However, no bloom was recorded in the Mixed Water Mass. High nutrient supply into the surface mixed layer during winter is likely to be one of the factors supporting an intense spring bloom in the Oyashio Water Mass. A significant positive relationship between log-transformed surface chlorophyll a concentration and maximum density gradient (MDG) within the euphotic layer was obtained in April, indicating the importance of vertical stability of the water column in the initiation of spring blooms in the Oyashio and the Coastal Water Masses. The spring blooms in 1991 were much more extensive and lasted longer than in 1990. It is suggested that meteorological conditions and abundance of grazers were responsible for this interannual difference.     

Multi‐timescale interactions between pink and chum salmon catch per unit effort in the Bering Sea

Based on generalized linear models, interspecific interactions were identified between chum and pink salmon. In addition, the effects of sea surface temperature and location on the variability of catch per unit effort (CPUE) of chum salmon from gill‐net surveys carried out between 1972 and 2010 were investigated. In the optimal model, interspecific interactions between CPUEs of chum and pink salmon on a year scale were positive for approximately half of all years in the central Bering Sea. In addition, interspecific interactions on a multi‐year scale were positive in even‐numbered years. The effects of location on the CPUE of chum salmon were significant variables in the optimal model. The CPUEs of chum salmon located near the continental shelf in the Bering Sea were higher than those of other locations. This study provides new evidence of positive interspecific interactions between the CPUEs of chum and pink salmon. The results also suggest that the standardized CPUE of chum salmon from the gill‐net surveys reflects relative chum salmon abundance in the North Pacific Ocean in the following year.     

卫星遥感北太平洋渔场叶绿素a浓度

海洋叶绿素a(Chla.)含量是了解世界海洋中地球生物化学循环的基础和估算海洋生产力的基本指标,是判断水域的肥瘠程度和评价水域渔业潜在生产力的基本依据。富集浮游植物的海域是海洋食植动物的大密度存在和水产资源丰富存在的基础,许多鱼类如金枪鱼等中上层鱼群集在锋面两侧或涡旋,这些特征与营养盐供给或混合层深度变化相关。北太平洋渔场已成为我国远洋渔业的重要组成部分,卫星遥感技术可以实现大面积同步的观测和一年四季连续的观测,是测定北太平洋渔场海洋叶绿素a的最有效方法。1 材料和方法1.1 数据来源卫星资料采用美国海洋水色卫星的宽视场海洋水色扫描仪(SeaWiFS)资料,全球海上测量资料利用国际海洋水色协调工作组(IOCCG)在世界大洋范围测量的1175个站位光谱数据和对应的叶绿素a浓度,南海测量数据由国家海洋技术中心在南海测量的48个站位光谱数据和叶绿     

Interannual variation in Neocalanus biomass in the Oyashio waters of the western North Pacific

We examined the interannual variation in Neocalanus copepod biomass in the Oyashio waters in spring and summer from 1972 to 1999. In the mid‐1970s, mesozooplankton biomass in spring was high; however, it decreased significantly in the late 1970s. The timing of the decrease in mesozooplankton biomass corresponded to the 1976/77 climatic regime shift. The biomass of N. flemingeri, which dominated the Neocalanus community, was roughly constant from 1980 to 1999. Although species‐level estimates of Neocalanus biomass were not available for the 1970s, a previous study reported that Neocalanus copepods were the predominant mesozooplankton in the Oyashio waters in spring during the 1970s. Neocalanus copepods dominated the mesozooplankton community throughout the 1970s, and their biomass decreased in the late 1970s. Springtime net community production, an index of new production, also decreased in the late 1970s. We suggest that the reduction in new production negatively affected Neocalanus food availability, resulting decreased copepod biomass. New production may have been limited by a combination of subsurface iron supplies, increased vertical density gradient, and reduced vertical water mixing in winter, which resulted in diminished iron entrainment in winter. In summer, mesozooplankton biomass significantly decreased and increased synchronously with the 1976/77 and 1988/89 climatic regime shifts. The biomass of N. plumchrus, which dominated the Neocalanus community, was low in the 1980s and increased in the early 1990s. The biomass of the second‐most dominant copepod, N. cristatus, also increased in the early 1990s. Neocalanus copepods were reported to be a dominant component of the mesozooplankton community in the 1970s; Neocalanus biomass was high in the mid‐1970s and decreased in the late 1970s. Japanese sardine (Sardinops melanostictus), an important predator of Neocalanus copepods, exhibited interannual variation in standing stock that was inversely related to mesozooplankton biomass. At their peak in 1984, sardines consumed 32–138% of the daily Neocalanus production during summer. Therefore, predation pressure on Neocalanus by Japanese sardine is likely to affect interannual variation in mesozooplankton biomass during the summer.     

Ecological effects of regime shifts in the Bering Sea and eastern North Pacific Ocean

Large‐scale shifts occurred in climatic and oceanic conditions in 1925, 1947, 1977, 1989 and possibly 1998. These shifts affected the mix and abundance of suites of coexisting species during each period of relative environmental stability—from primary producers to apex predators. However, the 1989 regime shift was not a simple reversal of the 1977 shift. The regime shifts occurred abruptly and were neither random variations nor simple reversals to the previous conditions. Timing of these anomalous environmental events in the North Pacific Ocean appears to be linked to physical and biological responses in other oceanic regions of the world. Changes in the atmospheric pressure can alter wind patterns that affect oceanic circulation and physical properties such as salinity and depth of the thermocline. This, in turn, affects primary and secondary production. Data from the North Pacific indicate that regime shifts can have opposite effects on species living in different domains, or can affect similar species living within a single domain in opposite ways. Climatic forcing appears to indirectly affect fish and marine mammal populations through changes in the distribution and abundance of their predators and prey. Effects of regime shifts on marine ecosystems are also manifested faster at lower trophic levels. Natural variability in the productivity of fish stocks in association with regime shifts indicates that new approaches to managing fisheries should incorporate climatic as well as fisheries effects.     

Anomalous conditions in the south-eastern Bering Sea, 1997: nutrients, phytoplankton and zooplankton

Anomalies in the regional weather over the south-eastern Bering Sea during spring and summer of 1997 resulted in significant differences in nutrient availability, phytoplankton species composition, and zooplankton abundance over the continental shelf as compared with measurements in the 1980s. Calm winds and the reduction of cloud cover in spring and summer produced a very shallow mixed layer in which nitrate and silicate were depleted after an April diatom bloom. High submarine light levels allowed subsequent phytoplankton growth below the pycnocline and eventual depletion of nitrate from the water column to depths of 70 m or more. Thus, total new production during 1997 may have exceeded that of previous years when nitrate was not depleted below the pycnocline. A bloom of the coccolithophorid, Emiliania huxleyi , was observed in early July in the warm, nutrient-depleted waters over the middle and inner shelf. Emiliania huxleyi concentrations reached 4.5 × 10⁶ cells L^–1 by September, and the bloom persisted through the autumn. There was evidence for increased abundance of some species of copepods in 1997 as compared with data from the middle domain in June 1981. The abundance of adult and juvenile euphausiids in 1997 was statistically similar to values measured in 1980 and 1981. However, near-surface swarms were rarely observed on the inner shelf in August–September 1997. Lack of euphausiid availability in the upper water column may partially explain the August–September mass mortality of planktivorous short-tailed shearwaters ( Puffinus tenuirostris ) observed on the inner shelf.     

东海近岸海域浮游动物群落时空分布

为探讨东海近岸海域浮游动物时空分布特征,根据2013年5月(春季)、8月(夏季)和12月(冬季)东海近岸海域3个航次的浮游动物调查资料,分析了该海域春季、夏季和冬季浮游动物的种类组成、丰度与生物量、优势种季节变化及生态类群。结果显示,3个航次共鉴定浮游动物108种(含未定种),浮游幼体14类,隶属于7门15大类,其中桡足类和水母类的种类和数量占绝对优势,桡足类为最优势类群,共47种,占总种数的43.52%,水母类共19种,占总种数的17.59%。浮游动物种类季节变化较为明显,夏季种类显著高于春季和冬季。各季节浮游动物种类数目与纬度呈负相关。浮游动物平均丰度与平均生物量随季节变化较为明显,其中平均丰度与平均生物量冬季最低,平均生物量夏季最高,平均丰度春季最高。根据浮游动物对水文环境条件耐受性和生活海区的差异性将浮游动物群落分为6个生态类群:近岸低盐种、广温广盐种、高温高盐种、近岸暖温种、暖水种和大洋广布种。其中东海北部近岸海域春、冬季近岸低盐种和广温广盐种占绝对优势,夏季暖水性种群所占比例高;长江口及邻近水域暖水性种群在春、夏、冬三季中均占主导地位;东海中南部近岸海域春、冬季暖水性种群占绝对优势,但夏季高温高盐种、大洋广布种优势明显。各季节中华哲水蚤均为优势种,但其他优势种季节更替较明显,其中春、夏两季优势种更替率为90.9%,夏、秋两季为50%。     

Interannual differences in prey taken by capelin, herring, and red salmon relative to zooplankton abundance during the spring bloom in a southeast Alaskan embayment

Capelin, herring, and red salmon diets were examined in relation to zooplankton abundance and biomass in the water column and surface layer of Auke Bay, Alaska, during the spring bloom period, 1987 and 1988 (April through mid-June). Euphausiid eggs were the dominant prey of capelin in mid-May 1987. Pseu-docalanus spp. and barnacle nauplii dominated during the rest of the season. Capelin consumed Calanus spp. and Metridia spp. in April and Pseudocaknus and Cen-tropages abdominalis in May and June 1988. During May and June 1987, herring were eating primarily barnacle larvae and Oikopleura spp. During April 1988, herring consumed primarily Calanus spp. and barnacle nauplii. In late April and early May they shifted to Pseudocalanus and Thysanoessa raschii, and in late May and June they consumed Centropages abdominalis and barnacle cyprids. Outmigrating red salmon fry consumed primarily Oikopleura during both years, along with substantial quantities of barnacle larvae. These dietary changes roughly correspond with variations in the abundance of prey taxa in the plankton samples. Likelihood measures of niche breadth indicated that capelin sometimes consume prey in approximately equal proportions to its abundance. Niche overlap between the herring and capelin was greatest in April and early May. Niche overlap was also high between herring and red salmon during June, when herring were in shallow water to breed.     

黄海鱼类功能群及其对浮游动物捕食的季节变化

用2006年9月-2007年8月对黄海中南部进行的5次断面调查,通过胃含物分析、聚类分析、功能群划分以及估算摄食量等方法,研究了黄海鱼类群落功能群组成及其对浮游动物摄食的季节变化。结果表明,黄海鱼类群落包括7个功能群,除初春以外,黄海鱼类群落主要以浮游生物食性功能群为主。春夏季的功能群组成简单,秋冬季的功能群组成变得复杂。黄海鱼类群落摄食的浮游动物种类主要有11种,春季摄食的浮游动物量最大,夏秋季逐渐减少,直至冬季摄食的浮游动物量最少,初春摄食的浮游动物量有所回升。不同季节摄食的浮游动物种类,以及摄食量都有较大差异,仅太平洋磷虾在各个月份的食物中均有出现,其余浮游动物饵料种类均是季节性的。     

Pacific herring, Clupea pallasi, recruitment in the Bering Sea and north-east Pacific Ocean, I: relationships among different populations

We examined recruitment and average weight-at-age time series for Pacific herring ( Clupea pallasi ) populations from the Bering Sea and north-east Pacific Ocean to determine similarities. Statistical correlation and multivariate clustering methods indicated Pacific herring populations form large-scale groups. Large year classes occur synchronously among several Pacific herring populations. Multivariate cluster analyses of recruitment and weight-at-age data indicated that Bering Sea herring populations are distinct from north-east Pacific Ocean populations. Within the NE Pacific Ocean, there appear to be three groups of herring populations: a British Columbia group, a south-east Alaska coastal group, and an outer Gulf of Alaska group. Jackknife and randomization tests indicate these groups are robust and not the result of random chance. Deviations from observed herring population groups were examined for indications of anthropogenic perturbations. The Prince William Sound herring populations did not show any strong deviations corresponding to the oil spill of 1989. There might not yet be enough data since the spill to detect changes in the recruitment or weight-at-age data since that time, particularly if oil spill effects were concentrated on the early life history stages.     

Intercalibrating SCOR, NORPAC and bongo nets and the consequences for interpreting decadal-scale variation in zooplankton biomass in the Gulf of Alaska

The Canadian weathership time series of zooplankton wet weight biomass, collected at Ocean Station Papa in the Gulf of Alaska from 1956 to 1981, is one of the most comprehensive of its kind in marine science. In 1966, the sampling gear changed from a white North Pacific (NORPAC) net to a black Scientific Committee on Oceanic Research (SCOR) net and it was recently discovered that insufficient intercalibration samples were collected to understand how their sampling properties differed. A Canada‐GLOBEC project to redo the intercalibration of these net types and to understand how they relate to current sampling gear (bongo net) started in 1997. Seventy replicates of the three net types were collected in deep water in the Gulf of Alaska. The major finding is that all nets have similar sampling characteristics, whereas earlier reports indicated that NORPAC biomass values should be multiplied by a factor of 1.538 to be equivalent to the SCOR net. It now appears that this factor arose because flowmeters were not used in the original 1956–81 sampling (volume filtered was estimated from tow length × mouth area). A positive bias was introduced into the SCOR values because relatively more water passed through the SCOR net (undetected without a flow meter) than through the NORPAC net. This means that the unmetered NORPAC samples from 1956 to 1966 should not be adjusted and the unmetered SCOR values should be reduced by a factor that is related to wire angle. The general effect on the entire series is to lower the average biomass estimates, but more so for the early portion of the series than the later years.     

Potential trophodynamic and environmental drivers of steelhead (Oncorhynchus mykiss) productivity in the North Pacific Ocean

Information on prey availability, diets, and trophic levels of fish predators and their prey provides a link between physical and biological changes in the ecosystem and subsequent productivity (growth and survival) of fish populations. In this study two long‐term data sets on summer diets of steelhead (Oncorhynchus mykiss) in international waters of the central North Pacific Ocean (CNP; 1991–2009) and Gulf of Alaska (GOA; 1993–2002) were evaluated to identify potential drivers of steelhead productivity in the North Pacific. Stable isotopes of steelhead muscle tissue were assessed to corroborate the results of stomach content analysis. We found the composition of steelhead diets varied by ocean age group, region, and year. In both the GOA and CNP, gonatid squid (Berryteuthis anonychus) were the most influential component of steelhead diets, leading to higher prey energy densities and stomach fullness. Stomach contents during an exceptionally warm year in the GOA and CNP (1997) were characterized by high diversity of prey with low energy density, few squid, and a large amount of potentially toxic debris (e.g., plastic). Indicators of good diets (high proportions of squid and high prey energy density) were negatively correlated with abundance of wild populations of eastern Kamchatka pink salmon (O. gorbuscha) in the CNP. In conclusion, interannual variations in climate, abundance of squid, and density‐dependent interactions with highly‐abundant stocks of pink salmon were identified as potential key drivers of steelhead productivity in these ecosystems. Additional research in genetic stock identification is needed to link these potential drivers of productivity to individual populations.     

Pacific herring, Clupea pallasi, recruitment in the Bering Sea and north-east Pacific Ocean, II: relationships to environmental variables and implications for forecasting

Previous studies have shown that Pacific herring populations in the Bering Sea and north-east Pacific Ocean can be grouped based on similar recruitment time series. The scale of these groups suggests large-scale influence on recruitment fluctuations from the environment. Recruitment time series from 14 populations were analysed to determine links to various environmental variables and to develop recruitment forecasting models using a Ricker-type environmentally dependent spawner–recruit model. The environmental variables used for this investigation included monthly time series of the following: southern oscillation index, North Pacific pressure index, sea surface temperatures, air temperatures, coastal upwelling indices, Bering Sea wind, Bering Sea ice cover, and Bering Sea bottom temperatures. Exploratory correlation analysis was used for focusing the time period examined for each environmental variable. Candidate models for forecasting herring recruitment were selected by the ordinary and recent cross-validation prediction errors. Results indicated that forecasting models using air and sea surface temperature data lagged to the year of spawning generally produced the best forecasting models. Multiple environmental variables showed marked improvements in prediction over single-environmental-variable models.     

Anomalous conditions in the south-eastern Bering Sea 1997: linkages among climate, weather, ocean, and Biology

In 1997, the Bering Sea ecosystem, a productive, high-latitude marginal sea, demonstrated that it responds on very short time scales to atmospheric anomalies. That year, a combination of atmospheric mechanisms produced notable summer weather anomalies over the eastern Bering Sea. Calm winds, clear skies, and warm air temperatures resulted in a larger-than-normal transfer of heat to surface waters and the establishment of a shallow mixed layer. In spring, significant new production occurred below the shallow pycnocline over the Middle Shelf, depleting the subpycnocline nutrient reservoir that normally exists during summer. Following the depletion of nitrate and silicate from the system, a sustained (≥ 4 months) bloom of coccolithophores ( Emiliania huxleyi ) was observed – a phenomenon not previously documented in this region. Summer Middle Shelf Domain copepod concentrations were higher for some species in 1997 than in the early 1980s. Warmer surface water and lack of wind mixing also changed the basic distribution of hydrographic regimes on the south-eastern shelf and altered the strength and position of fronts or transition zones where apex predators seek elevated food concentrations. The Inner Front was well inshore of its normal position, and adult euphausiids (the primary prey of short-tailed shearwaters, Puffinus tenuirostris ) were unavailable at, and shoreward of, the front in autumn. High shearwater mortality rates followed the period of low euphausiid availability. Some, but not all, of these anomalous conditions re-occurred in 1998. These observations are another demonstration that the structure and function of marine ecosystems are intimately tied to forcing from the atmosphere. Alteration of climatological forcing functions, expressed as weather, can be expected to have large impacts on this ecosystem and its natural resources.     

Changes in the long‐term distribution of zooplankton in the Humboldt Current Ecosystem off Peru, 1961–2005, and its relationship to regime shifts and environmental factors

We used a dataset assembled by the Peruvian Marine Research Institute (IMARPE) from 1961 to 2005, to examine the influence of seasonal, environmental (temperature and salinity), temporal (day–night and year) and location (north‐south and on‐off shelf) variables on the distribution and biovolume of zooplankton. Classification and regression trees (CART) showed that the zooplankton distribution can be divided into four time regimes: prior to 1974.5, 1974.5–1989.5, 1989.5–1997.5 and 1997.5–2005. These periods differ in overall biovolume but, more importantly, in having different spatial distributions of variables. Offshore‐onshore, seasonal, diel and north‐south patterns are all apparent in the temporal changes observed in zooplankton distribution. In the early period zooplankton biovolume was high throughout the Peruvian Humboldt Current Ecosystem, with enhanced biovolume in all seasons but winter. After 1974.5, biovolume was considerably lower. There was an increased difference between the shelf (lower biovolume) and offshore (higher biovolume), with a significant day–night effect offshore, due to diel migration and possibly diel catchability changes. After 1989.5, seasonal effects were more marked and biovolume appeared to be higher in the winter in the northern region. In other seasons, during the daytime, biovolume was highest farther offshore (greater than 65 km offshore of the 200‐m isobath) and, during the night, higher after 1997.5 than before. We relate these changes in distribution and biovolume to changes in large‐scale current patterns in different regimes and to the possible effect of the fishery in removing nutrients from the system.