Assemblages of vertical migratory mesopelagic fish in the transitional region of the western North Pacific

Assemblages of vertical migratory mesopelagic fish are described for the nighttime upper 20 m layer of the transitional and adjacent waters in the western North Pacific. Twenty-three mesopelagic fish species belonging to six families and 16 genera were collected during June 1996. Family Myctophidae was most speciose, represented by 17 species, followed by Sternoptychidae (two spp.) and Bathylagidae (one spp.). Six assemblages are recognized based on their species composition: Subarctic, Northern Transition, Southern Transition, Subtropical, Slope Water and Seamount Assemblages. These assemblages are also characterized by size composition and biomass of the catches. The distributions of the Subarctic, Northern Transition, Southern Transition and Subtropical Assemblages are approximately defined by the Kuroshio and Oyashio fronts and the Subarctic Boundary. Slope Water and Seamount Assemblages are closely related to the slope water around the shelf edge off Japan and waters around the Shatsky Rise and the Emperor Seamount chain, respectively. The importance of the Southern Transition Assemblage area between the Kuroshio front and Subarctic Boundary as spawning and nursery grounds of many subarctic, transitional and subtropical fishes is elucidated, based on the geographical distribution patterns of their larvae and juveniles.     

Distribution patterns of larval myctophid fish assemblages in the subtropical–tropical waters of the western North Pacific

We examined larval myctophid fish assemblages and their distribution patterns, based on discrete depth sampling. Samples were collected at 19 stations along a transect that crossed the subtropical–tropical waters of the western North Pacific. In total, we collected 27 189 larvae of 40 myctophid species or types, belonging to 15 genera. Three assemblages were recognized, based on their species composition: Kuroshio Axis (KuA), Kuroshio Countercurrent (KCC), and Subtropical Countercurrent–North Equatorial Current (SCC–NEC) assemblages. The distributions of these assemblages were well defined by the positions of the KuA and the Subtropical Convergence. Each species had a specific distribution depth, and none showed diel vertical migration. Larvae of the subfamily Lampanyctinae were distributed in shallower waters (0–50 m) than larvae of the subfamily Myctophinae (50–150 m). Larvae of the same species were distributed at lower depths in the SCC–NEC area than in the KCC area. This corresponded to the abundance of chlorophyll a, which would reflect abundance of prey organisms such as copepod nauplii and copepodites.     

Horizontal and vertical distribution patterns of larval myctophid fishes in the Kuroshio Current region

The horizontal and vertical distribution of myctophid fish larvae is described based on discrete depth sampling from the surface down to 200 m at seven stations on a transect crossing the Kuroshio Current on 6–9 July 1977. A total of 7819 larvae was collected. Myctophid larvae accounted for 72% of the total fish larvae and included 18 species or types belonging to 12 genera. Based on horizontal distribution patterns these species (or types) were categorized into three groups, i.e. Kuroshio-axis group ( Myctophum asperum , M. orientale , Lampadena luminosa and Symbolophorus evermanni ), Kuroshio east group ( Hygophum reinhardtii , Lampanyctus alatus , Diogenichthys atlanticus , Lobianchia gemellarii , M. nitidulum , Benthosema suborbitale , Lampadena sp. and Ceratoscopelus warmingii ) and Pan- Kuroshio group [ Diaphus stubby type (mostly D. garmani ) and D. kuroshio ]. Larvae of the subfamily Lampanyctinae were distributed in shallower (0–30 m) waters than those of the subfamily Myctophinae (50–150 m), while this relationship was opposite of the night-time depth distribution of adults of the two subfamilies.     

Diel vertical migration of sablefish (Anoplopoma fimbria)

To investigate their diel vertical migration (DVM), 599 sablefish (Anoplopoma fimbria) were implanted with electronic archival tags in the Gulf of Alaska, Aleutian Islands, and the eastern Bering Sea. Of these tags, 98 were recovered with usable depth data (7,852,773 recordings representing 81,233 days) that we used to identify DVM and to classify DVM into one of two types: normal DVM (rise from the bottom during nighttime) and reverse DVM (rise from the bottom during daytime). The results of our study highlight three important attributes of DVM for sablefish. First, all tagged sablefish carried out DVM, although the occurrence was intermittent (26% of the days with usable data) and most commonly for short durations (10 days or less). Second, bottom depth for normal DVM was about 78 m shallower than for reverse DVM. Third, normal DVM occurred most often in fall and least often in spring, whereas this high/low pattern was shifted about 3 months later for reverse DVM; reverse DVM occurred most often in winter and least often in summer. Normal DVM likely occurred to increase foraging opportunity (e.g., nightly shift to match depth of prey). Reverse DVM more commonly occurred during winter and may represent an increase in foraging by sablefish during the daytime to compensate for decreased pelagic resources. The default foraging strategy for sablefish may be benthic because of the uncertainty of vertically migrating to a location where the occurrence of prey is not guaranteed; sablefish may invoke DVM when the non‐DVM foraging benefit is reduced.     

Distribution patterns of leptocephali in the Kuroshio–Oyashio transitional region of the western North Pacific

Leptocephalus assemblages and distribution patterns were studied in the Kuroshio–Oyashio transitional region of the western North Pacific. A total of 479 leptocephali, representing at least 21 species (types) including eight undescribed types, were collected during a cruise of the T/V Hokuho Maru from 8 to 30 May 2002. The most abundant leptocephali were the congrids Gnathophis nystromi nystromi and Gnathophis nystromi ginanago. The assemblages at each station were clustered into four groups (one group in the Kuroshio region and three in the transition zone) using the ordination and the fuzzy c‐means method based on species composition similarities between stations. Each leptocephalus species was also clustered by the same method into one of three groups. The first group had the greatest numbers of species and mainly occurred at the station group in the Kuroshio region, sharing a similar distribution to G. n. nystromi. The second group had the greatest abundance, especially in the transition zone, but contained only one species, G. n. ginanago, which also occurred at all stations. Moreover discrimination of station groups in the transition zone corresponded well with the assemblage pattern of this species group. The last species group comprised rare and low abundance species. Most leptocephali in the study area were reported from the Kuroshio upstream region, so the discrimination of leptocephali assemblages in the study area may reflect their different ecology. Furthermore, the analytical methods used in this study yielded more meaningful and clearer groups than the other usual methods, validating application of this analysis to complex areas such as the transition zone.     

北太平洋亚热带海域大眼金枪鱼秋季摄食习性的初步研究

中国金枪鱼延绳钓渔业观察员于2018年9-12月在北太平洋亚热带水域(150°W-164°W,30°N-37°N)采集了146尾大眼金枪鱼(Thunnus obesus)样品,并对其胃含物组成进行了研究。结果表明,大眼金枪鱼胃含物中共鉴别出36种饵料种类,隶属于28科,优势饵料为帆蜥鱼(Alepisaurus ferox)、发光柔鱼(Eucleoteuthis luminosa)、魣蜥鱼(Lestidium prolixum)和眶灯鱼(Diaphus sp.),其相对重要性指数百分比(IRI%)分别为25.65%、14.48%、7.56%和6.77%。大眼金枪鱼在叉长为100~110 cm范围内的摄食强度最高,在性腺成熟度Ⅲ期的摄食强度也最高,而Ⅴ期的摄食强度最低,此外,在150~250 m水层中的摄食强度较高。研究结果有助于提高对大眼金枪鱼生物学特征和摄食习性的认识,及进一步了解其在生态系统和食物网结构中所扮演的重要角色。     

2013年冬季南乔治亚岛南极磷虾群昼夜垂直移动研究

关于冬季南极磷虾群昼夜垂直移动方面的研究可为探索其渔场形成机制提供基础数据,并为其越冬策略研究提供参考。基于2013年冬季南乔治亚岛南极磷虾渔业调查期间收集的相关数据,对冬季南乔治亚岛南极磷虾群昼夜垂直移动进行了研究。结果发现,磷虾群平均深度维持在表层以下100～300 m,磷虾群最深出现在日升时分,而最浅则出现在夜间时分;7—9月,随时间的推移,磷虾群所处水层呈加深趋势。夜间、白天和黄昏时段磷虾群平均深度呈现较为明显的月份差异。白天磷虾群多集中在较深水层,夜间会上浮到较浅水层。     

Spatial and temporal patterns of vertical distribution for three planktivorous fishes in Lake Washington

Abstract – We sampled three limnetic fish species: juvenile sockeye salmon (Oncorhynchus nerka), three‐spine stickleback (Gasterosteus aculeatus) and longfin smelt (Spirinchus thaleichthys) in Lake Washington to quantify species‐specific patterns of diel vertical migration (DVM). Catch‐per‐unit‐effort data analysed from 15 years of midwater trawling documented seasonal and diel differences in vertical distributions for each species. These results were consistent with the hypothesis that the patterns of DVM in Lake Washington were affected by life history, size and morphology. Sockeye salmon showed clear DVM in spring but essentially no DVM in fall, remaining in deep water, whereas three‐spine sticklebacks were prevalent at the surface at night in both seasons. In fall, distribution patterns may be explained by differences in thermal performance (e.g., sticklebacks favouring warm water), but the patterns were also consistent with inter‐specific differences in predation risk. Younger sockeye salmon and longfin smelt were present in greater proportions higher in the water column during dusk and night periods than older conspecifics. Compared with sockeye salmon, the greater use by three‐spine sticklebacks of surface waters throughout the diel cycle during weak thermal stratification in spring was consistent with the hypothesis that sticklebacks’ armour reduces predation risk, but use of this warmer, metabolically beneficial stratum may also have promoted growth. This study illustrates variation in the vertical distribution of three sympatric planktivores and offers broader implications for the DVM phenomenon and applied lake ecology.     

Effects of ontogeny, temperature, and light on vertical movements of larval Pacific cod (Gadus macrocephalus)

The role of behavior, especially vertical migration, is recognized as a critical component of realistic models of larval fish dispersion. Unfortunately, our understanding of these behaviors lags well behind our ability to construct three-dimensional flow-field models. Previous field studies of vertical behavior of larval Pacific cod ( Gadus macrocephalus ) were limited to small, preflexion stages (≤11 mm SL) in a narrow range of thermal conditions. To develop a more complete picture of larval behavior, we examined the effects of ontogeny, temperature, and light on vertical responses of larval Pacific cod in experimental columns. While eggs were strictly demersal, yolk-sac larvae displayed a strong surface orientation as early as 1 day post hatch (∼ 5 mm SL). Consistent with field observations, small preflexion larvae (<10 mm SL) showed no response to varying light levels. However, there was a direct effect of temperature on larval behavior: Pacific cod larvae exhibited a stronger surface orientation at 4°C than at 8°C. The behavior of larger, postflexion larvae (>15 mm SL) in experimental columns was consistent with a diel vertical migration and independent of water temperature: fish were more widely distributed in the column, and median positions were consistently deeper at higher light levels. These laboratory observations are combined with observations from discrete-depth (MOCNESS) sampling in the Gulf of Alaska to characterize the vertical distribution of larval Pacific cod and contrast ontogenetic patterns with walleye pollock ( Theragra chalcogramma ). The vertical movements of larval Pacific cod described here will be applied in the development of dispersal projections from Gulf of Alaska spawning grounds.     

Late winter larval mesopelagic fish assemblage in the Kuroshio waters of the western North Pacific

The larval mesopelagic fish assemblage and its distribution patterns were investigated in the Kuroshio region off southern Japan in late winter. A total of 8690 fish larvae was collected, 85.8% of which were mesopelagic fish larvae. Mesopelagic fish larvae were significantly more abundant in the area east of the Kuroshio axis than west of the Kuroshio axis (660.6 versus 194.5 ind 10 m^?2). Sigmops gracile, Bathylagus ochotensis, Notoscopelus japonicus, Diaphus slender type and Myctophum asperum were the five most abundant larvae and accounted for 16.9, 16.4, 15.2, 13.9 and 9.3% of the total catch in numbers, respectively. We conclude that these larvae were transported by the Kuroshio Current to the more productive transition waters, where they spend their juvenile stage from spring to early summer. The possibility of expatriation and southward long‐distance spawning migrations of N. japonicus and B. ochotensis are discussed, based on the geographic distribution patterns of their larvae, juveniles and adults.     

Migration and behavior of juvenile North Pacific albacore (Thunnus alalunga)

Archival tags were used to study the seasonal movements, migration patterns and vertical distribution of juvenile North Pacific albacore (Thunnus alalunga). Between 2001 and 2006, archival tags were deployed in North Pacific albacore in two regions of the Northeast Pacific: (i) off Northern Baja California, Mexico and Southern California, and (ii) off Washington and Oregon. Twenty archival tagged fish were recovered with times at liberty ranging from 63 to 697 days. Tagged albacore exhibited five distinct, seasonal migratory patterns. Depth and temperature data also showed a broad range of vertical behaviors. In certain regions such as off Baja California, Mexico, juvenile albacore make frequent dives to depths exceeding 200 m during the day and remain in the surface mixed layer at night, whereas off Oregon and Washington they remain near the surface both day and night. Water temperatures encountered ranged from 3.3 to 22.7°C. Peritoneal temperatures were significantly higher by an average of approximately 4°C, as expected in these warm‐bodied fish. This study provides a comprehensive examination of horizontal and vertical movements of juvenile albacore in the Northeast Pacific. The results reveal diverse behavior that varies regionally and seasonally as albacore move among different habitats throughout the entire North Pacific.     

Northerly shift of warm‐water copepods in the western subarctic North Pacific: Continuous Plankton Recorder samples (2001–2013)

Spatial and temporal variation in copepod community structure, abundance, distribution and biodiversity were examined in the western subarctic North Pacific (40–53°N, 144–173°E) during 2001–2013. Continuous Plankton Recorder (CPR) observational data during the summer season (June and July) were analyzed. The latitudinal distribution of warm‐water species in June shifted northward after 2011 while no apparent latitudinal shift of cold‐water and other species was observed. Species number and the Shannon–Wiener biodiversity index (H′) in June tended to increase in the northern area after 2011. The warm‐water species abundance and center latitude of warm‐water distribution were positively correlated with sea surface temperature (SST) across sampling locations, whereas no significant correlations with SST were observed for cold‐water species or other species. Warm SSTs in June after 2011 appeared to cause the northward shift of warm‐water species distribution, which in turn contributed to the higher biodiversity in the northern area. This study demonstrated the rapid response of warm‐water species to warm SST variation, whereas cold‐water and other species did not exhibit such clear responses. These findings indicate that the response of copepods to environmental changes differs among copepod species, highlighting the importance of investigating lower trophic levels to the species level to evaluate individual species’ responses to climate change.     

Habitat characteristics of skipjack tuna (Katsuwonus pelamis) in the western North Pacific: a remote sensing perspective

Skipjack tuna habitat in the western North Pacific was studied from satellite remotely sensed environment and catch data, using generalized additive models and geographic information systems. Weekly resolved remotely sensed sea surface temperature, surface chlorophyll, sea surface height anomalies and eddy kinetic energy data were used for the year 2004. Fifteen generalized additive models were constructed with skipjack catch per unit effort as a response variable, and sea surface temperature, sea surface height anomalies and eddy kinetic energy as model covariates to assess the effect of environment on catch per unit effort (skipjack tuna abundance). Model selection was based on significance of model terms, reduction in Akaike’s Information Criterion, and increase in cumulative deviance explained. The model selected was used to predict skipjack tuna catch per unit effort using monthly resolved environmental data for assessing model performance and to visualize the basin scale distribution of skipjack tuna habitat. Predicted values were validated using a linear model. Based on the four‐parameter model, skipjack tuna habitat selection was significantly (P < 0.01) influenced by sea surface temperatures ranging from 20.5 to 26°C, relatively oligotrophic waters (surface chlorophyll 0.08–0.18, 0.22–0.27 and 0.3–0.37 mg m^?3), zero to positive anomalies (surface height anomalies 0–50 cm), and low to moderate eddy kinetic energy (0–200 and 700–2500 cm² s^–2). Predicted catch per unit effort showed a trend consistent with the north–south migration of skipjack tuna. Validation of predicted catch per unit effort with that observed, pooled monthly, was significant (P < 0.01, r² = 0.64). Sea surface temperature explained the highest deviance in generalized additive models and was therefore considered the best habitat predictor.     

采用声学方法研究2016年秋季布兰斯菲尔德海峡南极磷虾群昼夜垂直移动特征及其影响因素

近年来,南极磷虾渔业过于集中于布兰斯菲尔德海峡,这也使得该海区磷虾资源状况及其生态学特征日益受到关注。南极磷虾群具有较为明显的昼夜垂直移动特征,开展此方面的研究可为探索其渔场形成机制提供基础数据,并为磷虾渔业反馈式管理提供参考。基于磷虾渔船上Simrad EK80记录的相关声学数据,使用Echoview软件判别声学数据中的磷虾群体,对2016年秋季布兰斯菲尔德海峡南极磷虾群昼夜垂直移动特征进行分析,并进一步分析影响磷虾群昼夜垂直移动的因素。结果显示,3月和4月磷虾群深度基本维持在250 m以浅,虾群最大深度出现在日升时分的频次最高（22.9%）,而最浅深度出现在夜间时分的频次最高（36.0%）,同时在日升时分,虾群厚度达到最大值;白天磷虾群多集中在较深水层,夜间会上浮到较浅水层。随着月份的推移,磷虾群平均深度总体呈现加深的趋势。光强和海底深度是影响磷虾群深度变化的2个主要因素。     

Habitat differentiation between sei (Balaenoptera borealis) and Bryde's whales (B. brydei) in the western North Pacific

Two closely related baleen whale species, sei and Bryde's whales, in the western North Pacific were studied to identify differences in habitat use. Data were obtained from May to August 2004 and 2005. This study examined the relationship between oceanographic features derived from satellite data and the distribution of sei and Bryde's whales using basic statistics. We investigated oceanographic features including sea surface temperature (SST), sea surface chlorophyll a (Chl‐a), sea surface height anomalies (SSHAs), and depth of the habitat. These two whale species used habitats with different SST, Chl‐a, and SSHA ranges. The 0.25 mg m^?3 Chl‐a contour (similar to the definition of the Transition Zone Chlorophyll Front) was a good indicator that separated the habitats of sei and Bryde's whales. Then generalized linear models were used to model the probabilities that the whale species would be present in a habitat and to estimate their habitat distribution throughout the study area as a function of environmental variables. The potential habitats of the two species were clearly divided, and the boundary moved north with seasonal progression. The habitat partitioning results indicated that SST contributed to the patterns of habitat‐use and might reflect differences in prey species between the two whales. This study showed that the habitats of the sei and Bryde's whales were clearly divided and their potential habitat‐use changed seasonally.     

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.     

Spatial distribution of the Japanese common squid, Todarodes pacificus, during its northward migration in the western North Pacific Ocean

The spatial distribution of Todarodes pacificus in and near the Kuroshio/Oyashio Transition Zone during its northward migration was examined by comparative surveys using two types of mid‐water trawl net and supplementary squid jigging from June to July 2000. The vertical and horizontal distribution patterns varied for different body sizes in relation to the oceanographic structure. Todarodes pacificus of 1–20 cm dorsal mantle length (ML) were widely distributed from the coastal waters of Japan to near 162°E longitude, probably due to transport by the Kuroshio Extension (KE). Todarodes pacificus smaller than 10 cm ML were mainly distributed in temperate surface layers at sea surface temperatures (SSTs) >15°C near the KE meander probably because of their poor tolerance to lower temperatures and limited swimming ability. Squid of 10–15 cm ML were distributed in the offshore waters of 10–15°C SST and in the coastal waters of northern Honshu, and underwent diel vertical migrations between the sea surface at night and deeper layers during the daytime. Squid larger than 15 cm ML were distributed in the coastal feeding grounds of northern Honshu and Hokkaido until they began their southward spawning migration. They also underwent diel vertical migrations, but remained deeper at night than the squid of 10–15 cm ML; this migration pattern closely matched that of their main prey such as euphausiids. We concluded that as T. pacificus grow, they shift their distribution range from the temperate surface layer around the KE toward the colder deeper layers, above 5°C, in the Oyashio and coastal areas.     

What goes up must come down: Diel vertical migration in the deep‐water sablefish (Anoplopoma fimbria) revealed by pop‐up satellite archival tags

The sablefish (Anoplopoma fimbria) is a long‐lived species with wide distribution throughout the North Pacific Ocean. While adult sablefish are considered a deep‐water fish, diet analyses suggest that they undergo vertical migrations that could be related to prey movement and feeding. Pop‐up satellite archival tags (PSATs) were used to observe the fine‐scale depth selection behavior of adult sablefish tagged off the Washington coast during the summer from June to August. Tags were physically retrieved after they surfaced using direction‐finding equipment so that complete datasets over the entire deployment were obtained from 14 tags. PSATs that recorded depth and temperature every 4 min during the deployment confirm that sablefish inhabit depths of 750 m or greater. However, a majority of the tagged fish underwent extensive vertical migrations that averaged 254.4 m overall and occurred at a 24 hr periodicity. Variations were observed among individuals in the amount of the deployment during which vertical migrations occurred, ranging from 12.37% to 63.48% of the time. During the vertical migration, fish ascended towards the surface at night and descended prior to daylight (i.e., diel vertical migration). Sablefish generally inhabited temperatures of 5°C but during the vertical migrations were found at temperatures from 6 to 10°C. Sablefish are opportunistic feeders with a large proportion of their diet being fish, euphausiids and cephalopods. Because these prey items also exhibit diel vertical migrations, it is possible that the vertical migratory behavior displayed by the sablefish was in response to the movements or the location of their prey.     

Albacore (Thunnus alalunga) fishing ground in relation to oceanographic conditions in the western North Pacific Ocean using remotely sensed satellite data

Satellite‐based oceanographic data of sea surface temperature (SST), sea surface chlorophyll‐a concentration (SSC), and sea surface height anomaly (SSHA) together with catch data were used to investigate the relationship between albacore fishing ground and oceanographic conditions and also to predict potential habitats for albacore in the western North Pacific Ocean. Empirical cumulative distribution function and high catch data analyses were used to calculate preferred ranges of the three oceanographic conditions. Results indicate that highest catch per unit efforts (CPUEs) corresponded with areas of SST 18.5–21.5°C, SSC 0.2–0.4 mg m^?3, and SSHA ?5.0 to 32.2 cm during the winter in the period 1998–2000. We used these ranges to generate a simple prediction map for detecting potential fishing grounds. Statistically, to predict spatial patterns of potential albacore habitats, we applied a combined generalized additive model (GAM) / generalized linear model (GLM). To build our model, we first constructed a GAM as an exploratory tool to identify the functional relationships between the environmental variables and CPUE; we then made parameters out of these relationships using the GLM to generate a robust prediction tool. The areas of highest CPUEs predicted by the models were consistent with the potential habitats on the simple prediction map and observation data, suggesting that the dynamics of ocean eddies (November 1998 and 2000) and fronts (November 1999) may account for the spatial patterns of highest albacore catch rates predicted in the study area. The results also suggest that multispectrum satellite data can provide useful information to characterize and predict potential tuna habitats.