Physical oceanography of the North Carolina continental shelf during the fall and winter seasons: implications for the transport of larval menhaden

The circulation over the continental shelf off the southern Middle Atlantic Bight (MAB) and northern South Atlantic Bight (SAB) is examined for the fall and winter periods. Observational data are compared with results from a three-dimensional numerical model to identify the dominant processes on the shelf. By considering wind-forcing, tides, and a specified upstream inflow (into the MAB), the observed and modelled flow fields are in close agreement in the mid- and inner shelf regions. The resulting larval drift indicates a seasonal dependence of transport pathways from spawning grounds to estuarine nursery areas for menhaden larvae and other offshore-spawning estuarine-dependent fish. Specifically, the physical oceanography of the MAB and SAB during the fall and winter months suggests a north-to-south shift in spawning areas providing recruits to the Carolina estuaries, in agreement with the observed migration of the spawning populations.     

Physical oceanography of the North Carolina continental shelf during the fall and winter seasons: implications for the transport of larval menhaden

The circulation over the continental shelf off the southern Middle Atlantic Bight (MAB) and northern South Atlantic Bight (SAB) is examined for the fall and winter periods. Observational data are compared with results from a three-dimensional numerical model to identify the dominant processes on the shelf. By considering wind-forcing, tides, and a specified upstream inflow (into the MAB), the observed and modelled flow fields are in close agreement in the mid- and inner shelf regions. The resulting larval drift indicates a seasonal dependence of transport pathways from spawning grounds to estuarine nursery areas for menhaden larvae and other offshore-spawning estuarine-dependent fish. Specifically, the physical oceanography of the MAB and SAB during the fall and winter months suggests a north-to-south shift in spawning areas providing recruits to the Carolina estuaries, in agreement with the observed migration of the spawning populations.     

Larval transport during winter in the SABRE study area: results of a coupled vertical larval behaviour–three-dimensional circulation model

A three-dimensional circulation model was used in conjunction with larval fish vertical behaviour models to study the interaction between larval vertical distribution, advection and the outcome of larval transport along the central portion of the east coast of the United States. The circulation model was forced by tides, a northern boundary inflow, and winds. Vertical behaviour models were developed for Atlantic menhaden ( Brevoortia tyrannus ) and spot ( Leiostomus xanthurus ). The purpose of this modelling effort was to investigate the transport pathways of Atlantic menhaden and spot larvae from offshore spawning grounds to estuarine nursery habitats. The coupled circulation and behavioural model demonstrated the importance of along-shelf transport in what is generally thought to be a 'cross-shelf' problem. Cross-shelf transport was associated with bathymetric features, such as shoals. Both physical (e.g. wind) and biological (e.g. changes in larval behaviour) events were responsible for many of the observed patterns in larval transport. Overall, larval transport was determined by circulation but was modified by larval vertical distributions.     

Mechanistic links between climate and fisheries along the east coast of the United States: explaining population outbursts of Atlantic croaker (Micropogonias undulatus)

Climate has been linked to variation in marine fish abundance and distribution, but often the mechanistic processes are unknown. Atlantic croaker (Micropogonias undulatus) is a common species in estuarine and coastal areas of the mid‐Atlantic and southeast coasts of the U.S. Previous studies have identified a correlation between Atlantic croaker abundance and winter temperatures in Chesapeake Bay, and have determined thermal tolerances of juveniles. Here we re‐examine the hypothesis that winter temperature variability controls Atlantic croaker population dynamics. Abundance indices were analyzed at four life history stages from three regions along the east coast of the U.S. Correlations suggest that year‐class strength is decoupled from larval supply and is determined by temperature‐linked, overwinter survival of juveniles. Using a relation between air and water temperatures, estuarine water temperature was estimated from 1930 to 2002. Periods of high adult catch corresponded with warm winter water temperatures. Prior studies indicate that winter temperature along the east coast is related to the North Atlantic Oscillation (NAO); variability in catch is also correlated with the NAO, thereby demonstrating a link between Atlantic croaker dynamics, thermal limited overwinter survival, and the larger climate system of the North Atlantic. We hypothesize that the environment drives the large‐scale variability in Atlantic croaker abundance and distribution, but fishing and habitat loss decrease the resiliency of the population to periods of poor environmental conditions and subsequent weak year classes.     

From spawning grounds to the estuary: using linked individual-based and hydrodynamic models to interpret patterns and processes in the oceanic phase of Atlantic menhaden Brevoortia tyrannus life history

Present theory suggests that population regulation in marine fishes cannot be resolved until an understanding of the processes involved in shaping the overall distribution (the number of populations, geographical extent, mean abundance and temporal changes in abundance) is developed. Here, we present a step toward understanding Atlantic menhaden population patterns, by studying processes in the Middle and South Atlantic Bights, which shape those patterns. We use individual-based and hydrodynamic models to reinterpret the 'mechanics' of the menhaden life history, and put forward several potentially testable hypotheses. The success of the menhaden reproductive strategy seems to depend on the seasonal changes in the mean flow field of the Middle and South Atlantic Bights, suggesting that their life history may have been strongly structured by the regional physics of the system. Because the annual menhaden migration is size-based and spawning occurs throughout the population's range, the size distribution of the adult population may influence the supply of larvae to particular estuaries along the coast. Recruitment of larvae into Delaware and Chesapeake Bays may be dependent on spawning to the north of the bays' mouths, owing to coastline shape and orientation in the vicinity of the bays. Our results suggest that management of this resource might be improved by consideration of the spatial and temporal variability in both the biological and the physical system.     

Spawning and transport dynamics of Atlantic menhaden: inferences from characteristics of immigrating larvae and predictions of a hydrodynamic model

We examined patterns of abundance, age and spawning date distributions of Atlantic menhaden Brevoortia tyrannus larvae immigrating during two seasons through three North Carolina inlets—Oregon, Ocracoke and Beaufort—to elucidate their spawning and transport dynamics. These patterns were examined in conjunction with corresponding predictions from a three-dimensional, wind-and tide-driven hydrodynamic model. Larvae immigrating through different inlets showed consistent similarities as well as marked differences in temporal patterns of abundance, spawning dates and transport times. Intraseasonal patterns in abundance and spawning date distributions among inlets suggest that, in both study years, the spatio-temporal dynamics of menhaden immigration were driven by large-scale patterns along the Atlantic coast, rather than by localized variation in spawning activity. Interannual differences in the temporal patterns of spawning dates and larval immigration indicate interannual differences in transport dynamics and/or the spatial-temporal distribution of spawning. When the spawning locations predicted by the hydrodynamic model are interpreted in conjunction with advanced very high resolution radiometer sea-surface temperature information, the results are consistent with the limited historical information available on spatio-temporal distribution of Atlantic menhaden eggs and larvae. The transport model also predicted distributions of arrival times for immigrating larvae that were comparable in range and variability with observed patterns. Our use of data from immigrating larvae, coupled with a hydrodynamic transport model and sea-surface temperatures, allowed us to uncover relationships between spatio-temporal patterns of Atlantic menhaden spawning and transport dynamics that could not have been identified by either approach alone.     

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.     

The role of oceanographic conditions and plankton availability in larval fish assemblages off the Catalan coast (NW Mediterranean)

In the northwestern Mediterranean, most fish species reproduce in early summer and fewer in the autumn mixing period. This study analyses and compares larval fish assemblages (LFA) in both seasons, and is the first attempt to characterize LFA structure for the autumn period. We analyze horizontal and vertical distribution of fish larvae and the micro- and mesozooplankton biomass and abundance of the main zooplankton groups. The oceanographic situation was analyzed through the study of data from CTD, Nν-Shuttle and ADCP surveys. LFA were determined by ordination analyses based on larval abundance, and the relationships between larval assemblages and environmental variables were investigated through canonical correspondence analysis. The importance of some hydrographic variables (temperature, salinity and stability of the water column), current fields (along-shelf and across-shelf transport) and the abundance of zooplankton are discussed as important factors shaping the structure of larval assemblages. In early summer, LFA were mainly structured by a combination of bathymetry and trophic components, although sea surface temperature also played a role in shaping the horizontal larval distributions. In autumn, trophic variables were the main factors influencing the shelf-dwelling species assemblage. Larvae of oceanic species, on the other hand, were not related to trophic variables but were more affected by current fields.     

东海区小黄鱼海底水温分布特征及分析方法比较

水温是影响鱼类活动的重要因素,鱼类繁殖、生长、发育等阶段都对水温有着不同需求。本研究基于2014—2015年4个季节的大面积调查数据,结合R语言绘图技术,采用频率和生物量两种确定温度分布范围的方法,对小黄鱼(Larimichthys polyactis)的水温分布特征进行了分析和比较。研究结果发现:依据频率和生物量分别确定的温度分布范围差别较大;前者覆盖站位比例相对较高,后者覆盖站位累积生物量比例较高;后者在确保生物量累积比例的条件下,某些季节也能有较高的站位覆盖比例,如夏季和秋季;基于生物量确定的温度范围可以更好地反映目标对象的分布特征。与历史研究结果比较,本文的水温分布范围基本保持一致,推断小黄鱼水温分布范围相对稳定,本文研究结果可以反映小黄鱼的一般特征。研究过程中发现小黄鱼有两个群体,水温和空间分布有明显差异;一个群体分布偏北,分布最南水域不超过舟山渔场水域;另一个群体分布偏南,一般分布水域处于长江口渔场以南海域。群体分布环境特征的分离可以作为种群划分的一个依据,本文观察结果可以作为小黄鱼种群划分的一个侧面依据。研究还发现,春、冬冷季小黄鱼资源聚集程度高,而夏、秋暖季资源分布相对分散;冷季群体单一,主要是越冬索饵群体;而暖季群体复杂,包括产卵亲体、补充群体等;暖季出现多个高生物量水温范围,可能与不同群体不同的温度取向相关。     

Dietary lipid levels affect lipoprotein clearance,fatty acid transport,lipogenesis and lipolysis at the transcriptional level in muscle and adipose tissue of large yellow croaker (Larimichthys crocea)

To investigate the effect of dietary lipid levels on lipid metabolism in muscle and adipose tissue of large yellow croaker (Larimichthys crocea), fish (average body weight 150.0 g) were fed to one of three diets with a low (6%), moderate (12%, the control diet) or high (18%) crude lipid level for 10 weeks. The expression of genes related to lipoprotein clearance (lpl, ldlr, lrp1 and srbi), fatty acid transport (cd36, fatp1, fabp3 and fabp11), lipogenesis (fas and dgat2) and lipolysis (atgl, cpt1 and aco) in muscle and adipose tissue was analysed. In muscle of fish fed the low‐lipid diet, the expression of lpl, lrp1, fas, dgat2 and atgl was significantly downregulated, whereas fabp3 and fabp11 were significantly upregulated compared with those fed the control diet. In muscle of fish fed the high‐lipid diet, the expression of lpl, ldlr, lrp1, cd36, fabp3, fas and dgat2 was significantly decreased, while fabp11 expression was significantly increased compared with those fed the control diet. Compared with the control group, only a significant downregulation of fabp3 expression in adipose tissue was observed in the low‐lipid group. In the high‐lipid group compared with the control group, the expression of ldlr, fabp11, dgat2, atgl and aco in adipose tissue was significantly upregulated, whereas cpt1 expression in adipose tissue was significantly downregulated. These results indicated that the expression of the selected key genes related to lipid metabolism was tissue‐specific regulation by dietary lipid levels in large yellow croaker.     

Vertical distribution and migration of fish larvae in the NW Iberian upwelling system during the winter mixing period: implications for cross‐shelf distribution

The vertical distribution and vertical migrations of fish larvae and implications for their cross‐shelf distribution were investigated in the northern limit of the NE Atlantic upwelling region during the late winter mixing period of 2012. The average positive values of the upwelling index for February and March of this year were far from normal, although the average hydrographic conditions during the period of study were of downwelling and the water column was completely mixed. Fish larvae, most in the preflexion stage, were concentrated in the upper layers of the water column and their distribution was depth stratified, both day and night. However, the larval fish community was not structured in the vertical plane and fish larvae did not show significant diel vertical migration (DVM), although five species showed ontogenetic vertical migration. In regions of coastal upwelling and in the absence of DVM, the location of fish larvae in the water column is crucial for their cross‐shelf distribution. Thus, the cross‐shelf distribution of the six most abundant species collected in this study can be explained by the surface onshore flow associated with coastal downwelling, retaining larvae of the coastal spawning species with a relatively shallow distribution in the shelf region and transporting larvae of slope spawning species onto the shelf. The wide vertical distribution shown by larvae of the offshore spawning species could be an adaptation of these species to ensure that some larvae reach the inshore nursery areas.     

Understanding patterns of distribution shifts and range expansion/contraction for small yellow croaker (Larimichthys polyactis) in the Yellow Sea

Detecting and analyzing patterns of distribution shifts and range expansion/contraction is important to understand population dynamics and changes in stock status. Here, we develop a spatio‐temporal model for yellow croaker (Larimichthys polyactis), which was fitted to bottom trawl survey biomass data collected in the Yellow Sea in the winter of 2001–2017. The model accounts for both spatial and spatio‐temporal structure and can potentially include the effects of surface temperature and of an annual index, the Pacific Decadal Oscillation, represented using a recently developed spatially varying coefficient model. We selected a spatio‐temporal model with no covariates based on Akaike's information criterion. The center of gravity for yellow croaker was found to move northwest between 2001 and 2010, and then southwest over the period 2010–2017. These results reflected the predicted progressive disappearance of yellow croaker density hotspots (i.e., highest density areas) in the north and southeast areas of the Yellow Sea, which resulted in the central area of the Yellow Sea becoming the only yellow croaker density hotspot in 2017. This finding has important implications for fisheries management in the context of the China–South Korea fisheries agreement, as it indicates a measurable displacement of yellow croaker biomass toward China. The exclusion of covariates from the spatio‐temporal model was not expected a priori and may be due to the facts that environmental variations are not pronounced in winter in the Yellow Sea and that the representation of spatial and spatio‐temporal structure in spatio‐temporal models accounts for a large proportion of the variability in the data.     

Eulerian views of layered water currents, vertical distribution of some larval fishes, and inferred advective transport over the continental shelf off North Carolina, USA, in winter

Currents that effect the shoreward transport of the larvae of estuarine-dependent fishes spawned in winter in Onslow Bay, North Carolina, USA, were driven by winds and pressure gradients, and influenced by the Gulf Stream. Aside from storms, winds over the continental shelf in Onslow Bay blew predominantly alongshore with velocities approaching 14 m s^-1 during February and March 1986, and January and February 1989. Water masses and currents observed at two current-meter moorings, one at mid-shelf and the other on the outer shelf, reflected the onshore (or offshore) advection of interior water in compensation for the offshore (or onshore) advection of wind-driven surface water. Winds and currents reversed direction approximately every 4 to 6 days. The larvae of Atlantic menhaden, Brevoortia tyrannus , spot, Leiostomus xan- thurus, and Atlantic croaker, Micropogonias undulatus , were most abundant in 17–19^oC and 20–21^oC water of the outer shelf and Gulf Stream fronts. There was little indication of diel vertical migration; larval Atlantic menhaden were most abundant in mid- and surface water, while spot and Atlantic croaker were most abundant in mid- and deep water. Given this distribution, the inferred advective transport of larvae was at times onshore, but at other times it was offshore. Within a spawning season, the prevalence of either reciprocation could determine the number of larvae that reach coastal inlets.     

Behavioural changes of Atlantic cod (Gadus morhua) after marine boulder reef restoration: Implications for coastal habitat management and Natura 2000 areas

While marine reefs are degraded globally, the responses of fish to marine reef restoration remain uncertain, particularly in temperate waters. This study measured the effect of marine boulder reef restoration on the behaviour of Atlantic cod, Gadus morhua L., in a Natura 2000 area using acoustic telemetry. Cod were tagged and released in the study area before and after the restoration and tracked continuously for six months. A larger fraction of the released fish remained in the study area after restoration (94%) than before (53%). Moreover, throughout the study period, cod spent significantly more hours per day and prolonged their residence time in the study area after the restoration. The study indicates that marine reefs subjected to boulder extraction can be restored and function as favourable cod habitats. Temperate marine boulder reef restoration represents a valuable management tool to improve habitats for temperate fish species.     

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.     

Distribution of jack mackerel (Trachurus japonicus) larvae and juveniles in the East China Sea, with special reference to the larval transport by the Kuroshio Current

Horizontal distribution patterns of jack mackerel (Trachurus japonicus) larvae and juveniles were investigated in the East China Sea between 4 February and 30 April 2001. A total of 1549 larvae and juveniles were collected by bongo and neuston nets at 357 stations. The larvae were concentrated in the frontal area between the Kuroshio Current and shelf waters in the upstream region of the Kuroshio. The abundance of small larvae (<3 mm notochord length) was highest in the southern East China Sea (SECS) south of 28°N, suggesting that the principal spawning ground is formed in the SECS from late winter to spring. Jack mackerel also spawned in the northern and central East China Sea (NECS and CECS, respectively), as some small larvae were also collected in these areas. In the SECS, the abundance of small larvae was highest in February and gradually decreased from March to April. The habitat temperature of small larvae in the SECS and CECS (20–26°C) was higher than that in the NECS (15–21°C), suggesting higher growth rates in the SECS and CECS than in the NECS. The juveniles (10‐ to 30‐mm standard length) became abundant in the NECS off the west coast of Kyushu Island and CECS in April and were collected in association with scyphozoans typical of the Kuroshio waters. However, juveniles were rarely collected in the SECS, where the small larvae were concentrated. Considering the current systems in the study area, a large number of the eggs and larvae spawned and hatched in the SECS would be transported northeastward by the Kuroshio and its branches into the jack mackerels’ nursery grounds, such as the shallow waters off the west coast of Kyushu and the Pacific coast of southern Japan.