Characteristics of a Pacific herring Clupea pallasii spawning bed off Minedomari, Hokkaido, Japan

ABSTRACT:   The spawning bed selection of herring Clupea pallasii off Minedomari, Atsuta, on the west coast of Hokkaido, Japan, is discussed in relation to the topography of the rocky shore and wave conditions during the spawning season. From 1998 to 2003, herring have spawned their eggs mainly on the leaves of the seagrass Phyllospadix iwatensis Makino in almost the same site on the rocky shore off Minedomari in the Atsuta area. The site is connected to a valley-like feature offshore. Wave conditions were surveyed at Minedomari during the spawning season in 2000 and the wave height was estimated from 1998 to 2003, except for in 2000, using the correlation of wave height between Minedomari and Ishikari Bay New Port, approximately 18 km south-west of Minedomari. Herring spawned under calm conditions, during which the wave height was approximately 0.5 m off Atusta and Aoshima and was 0.18–1.28 m off Minedomari. The distribution of water particle velocity on the sea bottom surface as a result of ocean waves off the Minedomari area, which was estimated based on the wave height and the topography of the coast, suggested herring could swim easily into the shallower area along the valley-like feature off Minedomari. Therefore, topographical features are thought to be one of the reasons why herring have used Minedomari as a spawning bed. Additionally, seepage of freshwater from the bottom, which was observed in this area, could also be the reason why herring spawn in Minedomari repeatedly.     

Ecosystem controls of juvenile pink salmon (Onchorynchus gorbuscha) and Pacific herring (Clupea pallasi) populations in Prince William Sound,Alaska

Five years of field, laboratory, and numerical modelling studies demonstrated ecosystem‐level mechanisms influencing the mortality of juvenile pink salmon and Pacific herring. Both species are prey for other fishes, seabirds, and marine mammals in Prince William Sound. We identified critical time‐space linkages between the juvenile stages of pink salmon and herring rearing in shallow‐water nursery areas and seasonally varying ocean state, the availability of appropriate zooplankton forage, and the kinds and numbers of predators. These relationships defined unique habitat dependencies for juveniles whose survivals were strongly linked to growth rates, energy reserves, and seasonal trophic sheltering from predators. We found that juvenile herring were subject to substantial starvation losses during a winter period of plankton diminishment, and that predation on juvenile pink salmon was closely linked to the availability of alternative prey for fish and bird predators. Our collaborative study further revealed that juvenile pink salmon and age‐0 herring exploit very different portions of the annual production cycle. Juvenile pink salmon targeted the cool‐water, early spring plankton bloom dominated by diatoms and large calanoid copepods, whereas young‐of‐the‐year juvenile herring were dependent on warmer conditions occurring later in the postbloom summer and fall when zooplankton was composed of smaller calanoids and a diversity of other taxa. The synopsis of our studies presented in this volume speaks to contemporary issues facing investigators of fish ecosystems, including juvenile fishes, and offers new insight into problems of bottom‐up and top‐down control. In aggregate, our results point to the importance of seeking mechanistic rather than correlative understandings of complex natural systems.     

Pacific herring (Clupea pallasii) consumption by marine birds during winter in Prince William Sound,Alaska

Following the 1989 MV Exxon Valdez oil spill (EVOS) and subsequent herring population collapse in Alaska's Prince William Sound (PWS), the Pacific herring (Clupea pallasii) fishery was closed. In the 25 yr since EVOS, herring and several herring‐dependent marine bird species have failed to reach pre‐spill population levels. One hypothesis is that intense predation pressure may be inhibiting herring recovery. To inform herring modeling efforts, this study estimated marine bird consumption of juvenile and adult herring in PWS for 10 winters over an 18‐yr period (1989–90 through 2006–2007). Total estimated herring consumption by wintering marine birds averaged 2409 ± 950 t, indicating that avian consumption represents a substantial and inter‐annually variable source of herring mortality. Common murre (Uria aalge) consumed the greatest portion (16–80%) of herring in all years among marine bird species. Juvenile herring biomass consumed annually by common murre was greater than murre consumption of adult herring biomass. Time lag analyses showed that marine bird consumption of adult herring is negatively correlated with the amount of herring spawn observed in subsequent years, but such effects were not observed more than 2 yr. Our models indicate that during years in which herring recruitment is low or bird populations are particularly large, marine birds can consume up to 10% of the annual adult herring biomass. Our results highlight the importance of herring to wintering PWS birds. We propose that future management of herring stocks seeks to reduce negative impacts on marine birds that prey on herring.     

Larval Pacific herring, Clupea pallasii (Valenciennes), are highly susceptible to viral haemorrhagic septicaemia and survivors are partially protected after their metamorphosis to juveniles

Pacific herring were susceptible to waterborne challenge with viral haemorrhagic septicaemia virus (VHSV) throughout their early life history stages, with significantly greater cumulative mortalities occurring among VHSV‐exposed groups of 9‐, 44‐, 54‐ and 76‐day‐old larvae than among respective control groups. Similarly, among 89‐day–1‐year‐old and 1+year old post‐metamorphosed juveniles, cumulative mortality was significantly greater in VHSV‐challenged groups than in respective control groups. Larval exposure to VHSV conferred partial protection to the survivors after their metamorphosis to juveniles as shown by significantly less cumulative mortalities among juvenile groups that survived a VHS epidemic as larvae than among groups that were previously naïve to VHSV. Magnitude of the protection, measured as relative per cent survival, was a direct function of larval age at first exposure and was probably a reflection of gradual developmental onset of immunocompetence. These results indicate the potential for easily overlooked VHS epizootics among wild larvae in regions where the virus is endemic and emphasize the importance of early life history stages of marine fish in influencing the ecological disease processes.     

Testicular development and serum profiles of steroid hormone levels in captive male Pacific herring Clupea pallasii during their first maturational cycle

The present study investigates the relationship between testicular development and serum steroid hormone levels in captive Pacific herring Clupea pallasii during the first reproductive cycle. The maturity of the testis was divided into five periods based on histological observation. These are early spermatogenic stage (April to July), mid-spermatogenic stage (August to November), late spermatogenic stage (December to March), functional maturation stage (early April) and spent stage (late April). The pattern of seasonal change in gonadosomatic index (GSI) clearly reflected testicular maturity. 11-Ketotestosterone (11-KT) levels increased from October to a peak level (6.58 ± 1.87 ng/mL) in January, and were maintained at this level until March. In contrast, testosterone levels were consistently low, less than 1 ng/mL, at all times. These results suggest that 11-KT is the predominant androgen that controls spermatogenesis in this species. 17,20β-Dihydroxy-4-pregnen-3-one (DHP) showed a single sharp peak (3.38 ± 0.35 ng/mL) in early April of the second year, suggesting that milt production is induced by DHP as in some other teleost species.     

Viral replication in excised fin tissues (VREFT) corresponds with prior exposure of Pacific herring, Clupea pallasii (Valenciennes), to viral haemorrhagic septicaemia virus (VHSV)

Procedures for a viral replication in excised fin tissue (VREFT) assay were adapted to Pacific herring, Clupea pallasii, and optimized both to reduce processing time and to provide the greatest resolution between naïve herring and those previously exposed to viral haemorrhagic septicaemia virus (VHSV), Genogroup IVa. The optimized procedures included removal of the left pectoral fin from a euthanized fish, inoculation of the fin with >10⁵ plaque‐forming units (PFU) mL^?1 VHSV for 1 h, rinsing the fin in fresh medium six times to remove unadsorbed virions, incubation of the fin in fresh medium for 4 days and enumeration of the viral titre in a sample of the incubation medium by plaque assay. The optimized VREFT assay was effective at identifying the prior exposure history of laboratory‐reared Pacific herring to VHSV. The geometric mean VREFT value was significantly greater (P < 0.01) among naïve herring (1.2 × 10³ PFU mL^?1) than among groups that survived exposure to VHSV (1.0–2.9 × 10² PFU mL^?1); additionally, the proportion of cultures with no detectable virus was significantly greater (P = 0.0002) among fish that survived exposure to VHSV (39–47%) than among naïve fish (3.3%). The optimized VREFT assay demonstrates promise for identifying VHSV exposure history and forecasting disease potential in populations of wild Pacific herring.     

A spatially explicit, individual-based model to assess the role of estuarine nurseries in the early life history of North Sea herring, Clupea harengus

Herring (Clupea harengus) enter and remain within North Sea estuaries during well‐defined periods of their early life history. The costs and benefits of the migrations between offshore spawning grounds and upper, low‐salinity zones of estuarine nurseries are identified using a dynamic state‐variable model, in which the fitness of an individual is maximized by selecting the most profitable habitat. Spatio‐temporal gradients in temperature, turbidity, food availability and predation risk simulate the environment. We modeled predation as a function of temperature, the optical properties of the ambient water, the time allocation of feeding and the abundance of whiting (Merlangius merlangus). Growth and metabolic costs were assessed using a bioenergetic model. Model runs using real input data for the Scheldt estuary (Belgium, The Netherlands) and the southern North Sea show that estuarine residence results in fitter individuals through a considerable increase in survival probability of age‐0 fish. Young herring pay for their migration into safer estuarine water by foregoing growth opportunities at sea. We suggest that temperature and, in particular, the time lag between estuarine and seawater temperatures, acts as a basic cue for herring to navigate in the heterogeneous space between the offshore spawning grounds at sea and the oligohaline nursery zone in estuaries.     

Long‐term variation in the abundance of Pacific herring (Clupea pallasii) from the Yellow Sea in the western North Pacific and its relation to climate over the past 590 years

A qualitative understanding of the long‐term variation in the population dynamics of Yellow Sea (YS) herring is particularly important for clarifying the evolutionary processes and driving mechanisms of the YS large marine ecosystem. Unfortunately, because of a lack of long‐term, continuous, and simultaneous monitoring data, the specific driving processes and mechanisms of climate effects on the population dynamics of YS herring remain largely unknown. In response to this scientific issue, we preliminarily propose the idea of reconstructing long‐term changes in YS herring abundance over the past 590 years (AD 1417–2004) based on historical documents and attempt to explore the impacts of climate on the population. Our results show that YS herring abundance maintained at a relatively high level from AD 1417 to 1870 (during the Little Ice Age); in contrast, the population declined significantly from AD 1870 to 2004 at different rates. In addition, we also found that there were strong relationships between the population abundance of YS herring and the Pacific decadal oscillation (PDO) and drought/flood cycles. We suggest that the fluctuations in YS herring abundance may be influenced by ocean–climatic circulation shifts throughout the North Pacific, especially the PDO.     

A synthesis of the life history and ecology of juvenile Pacific herring in Prince William Sound,Alaska

Physical and biological variables affecting juvenile Pacific herring (Clupea pallasi) in Prince William Sound (PWS) from 1995 to 1998 were investigated as part of a multifaceted study of recruitment, the Sound Ecosystem Assessment (SEA) program. Though more herring larvae were retained in eastern PWS bays, ages‐0 and ‐1 herring used bays throughout PWS as nursery areas. Water transported into PWS from the Gulf of Alaska (GOA) contributed oceanic prey species to neritic habitats. Consequently, variations in local food availability resulted in different diets and growth rates of herring among bays. Summer food availability and possible interspecific competition for food in nursery areas affected the autumn nutritional status and juvenile whole body energy content (WBEC), which differed among bays. The WBEC of age‐0 herring in autumn was related to over‐winter survival. The limited amount of food consumption in winter was not sufficient to meet metabolic needs. The smallest age‐0 fish were most at risk of starvation in winter. Autumn WBEC of herring and winter water temperature were used to model over‐winter mortality of age‐0 herring. Differences in feeding and energetics among nursery areas indicated that habitat quality and age‐0 survival were varied among areas and years. These conditions were measured by temperature, zooplankton abundance, size of juvenile herring, diet energy, energy source (GOA vs. neritic zooplankton), WBEC, and within‐bay competition.     

The influence of life history dynamics and environment on the determination of year class strength in North Sea herring (Clupea harengus L.)

The inter-annual variability in year class strength (1976–2000) of North Sea herring (Clupea harengus) was investigated using Paulik diagrams based on survey data and Virtual Population Analysis. The herring life cycle was split into five stages: spawning stock biomass (SSB), egg production, larvae, fish with 0 winter rings on the otolith (0-wr), 1-wr and 2-wr. Surveys were used as indices and Paulik analysis revealed relationships between stages. In 80% of the years, year class strength reflected SSB. Poorer than expected year classes were determined during the larva to 0-wr phase, whilst stronger than expected year classes were apparently determined during the 0-wr to 1-wr stage. There was no clear relationship between survival of young stages of herring and the abundance of Calanus finmarchicus but the year class strength of 0-wr and 1-wr had a negative relationship to bottom water temperature. Lower sea water temperatures in the North Sea are associated with higher Calanus abundance. The analysis shows that the strength of aberrant year classes of North Sea herring is determined between the pelagic larval and the juvenile stages.     

An investigation of the biological basis of recruitment,growth and adult survival rate variability of Pacific herring (Clupea pallasi) from British Columbia: a synthesis

I explored the biological basis of variation in recruitment (age 3 abundance), growth and age‐specific adult survival rate for the major populations [West Coast Vancouver Island (WCVI), Strait of Georgia, Central Coast, North Coast and Haida Gwaii] of Pacific herring (Clupea pallasi) that inhabit British Columbian waters. The analyses were based on a synthesis of time series of empirical observations of herring population characteristics (egg deposition, age‐specific abundance and size) and prey, competitor and predator biomass/abundance. Recruitment was not correlated among populations. Recruitment variability was explained for WCVI herring only, as a consequence of prey (the euphausiid Thysanoessa spinifera) biomass during August in each of the first 3 years of life, and the biomass of piscivorous Pacific hake (Merluccius productus) during the first year of life. Recruit mass and adult mass‐at‐age were correlated among populations and over ages within populations. Recruit mass was affected by T. spinifera biomass in August of the first and third years of life. Adult mass‐at‐age variability was determined mainly by size at the beginning of the growth season, but also by T. spinifera biomass in August. Age‐specific adult survival rates were not correlated among the five populations. Survival rates decreased with age; there were additional population‐specific effects of somatic mass and T. spinifera biomass in August. The analyses were repeated using physical oceanographic explanatory variables. Only recruit mass variation was explained significantly by physical oceanographic variables, and the biological‐based explanation of recruit mass variability accounted for more of the variation.     

The effects of survey design and circulation pattern on the perceived abundance of herring larvae: a case study for Norwegian spring spawning herring (Clupea harengus)

A larval survey is used in the annual assessment as an index of the spawning stock size of Norwegian spring spawning herring (Clupea harengus). To test how inter‐annual fluctuations in circulation pattern, survey design and execution of the survey affected the larval abundance estimate we conducted simulated surveys using a model framework with idealized assumptions to model larval drift and sampled larvae using several realistic survey scenarios. The results suggest that inter‐annual variations in circulation pattern alone can have a profound effect on the perception of larvae abundance and that the direction of the survey (north to south versus south to north) can have a significant effect on the estimated abundance, particularly if hatching occurs over a short period of time. Additionally, disruptions to a continuous survey schedule also have an effect and, as such, sampling strategies in case of disruption to the survey are proposed.     

Habitat use of cownose rays (Rhinoptera bonasus) in a highly productive,hypoxic continental shelf ecosystem

Highly productive surface waters and hypoxic (dissolved oxygen, DO ≤ 2.0 mg L^?1) bottom waters develop seasonally on the northwestern Gulf of Mexico continental shelf due to nutrient and freshwater inputs from the Mississippi‐Atchafalaya River system. We investigated the spatial distribution of the cownose ray (Rhinoptera bonasus), a highly mobile, bentho‐pelagic species that is a seasonal resident of the shelf, in relation to surface chlorophyll, bottom‐water hypoxia, and other environmental variables (salinity, temperature, depth). We used synoptic trawl and aerial surveys to investigate ray distributions at both shelfwide (100–1000s km) and local (5–50 km) spatial scales. Shelfwide sampling indicated that rays were associated with regions of high surface chlorophyll and low bottom salinity and DO, conditions characterizing the Mississippi‐Atchafalaya plume region. Local sampling in and around the hypoxic zone indicated that rays preferred habitats where bottom waters were hypoxic but they primarily occupied normoxic (DO > 2.0 mg L^?1) waters above the bottom hypoxic layer. Stomach fullness and diet composition were similar between rays sampled in habitats with hypoxic versus normoxic bottom waters. These results indicate that cownose rays are strongly associated with riverine‐influenced regions of the shelf and preferentially use habitats with hypoxic bottom waters, perhaps for benthic foraging. Collectively, our results highlight the importance of considering the responses of mobile species to enhanced productivity and to hypoxia‐induced habitat degradation, which are both the products of coastal eutrophication.     

Spatial distribution and feeding habits of Pacific cod (Gadus macrocephalus) larvae in Mutsu Bay, Japan

The spatial distributions and feeding habits of Pacific cod ( Gadus macrocephalus ) larvae, and the spatial distributions of copepod nauplii and copepodites, their main prey, were examined in Mutsu Bay from February to March during 1989–92. Yolk-sac larvae were caught at 30–45 m depth at the bay mouth. Larvae without yolk were collected at 8–45 m depth at the bay mouth and the inner part of the bay, and large larvae were chiefly found in the bay. This geographical pattern in larval size may have been because of transport to the inner part of Mutsu Bay by the Tsugaru Warm Current. The dominant taxa of copepod nauplii and copepodites in the diet and the environment changed each year. Larvae fed mainly on abundant taxa in the environment, suggesting that larvae are opportunistic feeders. Nauplii and copepodites were abundant in the bay, especially in 1992. Copepodites were slightly more abundant in the diet of cod larvae in 1992 than in 1991, but this difference was smaller than in the environment. In addition, larvae with empty digestive tracts were scarce in 1991 and 1992. Prey concentrations in the bay in 1991 and 1992 seem to have been high enough to sustain most Pacific cod larvae.     

Application of a sensitive,specific and controlled real‐time PCR assay to surveillance indicates a low prevalence of viral haemorrhagic septicaemia virus (VHSV) in wild herring,Clupea harengus L., in Scottish waters

Surveillance data on the distribution of viral haemorrhagic septicaemia virus (VHSV) in the North Sea (UK), targeting Atlantic herring in areas with previous virus detection, were obtained from research cruises conducted during 2005. The sensitive molecular approach of real‐time RT‐PCR (qRT‐PCR) was applied alongside a newly developed endogenous positive control assay specific for herring (elongation factor 1α) to ensure integrity of template. Three hundred and five pools from 1937 individual herring were tested, and no evidence of VHSV in association with wild Atlantic herring was detected. Samples were obtained from Scottish waters where marine aquaculture is conducted. The results confirm that previous tissue culture studies have most likely not significantly underestimated the prevalence of carrier herring in this area. The significance of migratory species such as herring as a reservoir species for VHSV, with the potential to translocate virus genotypes between geographical areas, is discussed.     

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.     

Heritability for body weight at harvest size in the Pacific white shrimp, Penaeus (Litopenaeus) vannamei, from a multi-environment experiment using univariate and multivariate animal models

To estimate family BLUP breeding values and the heritability of body weight at harvest size (BW) in the Pacific white shrimp, Penaeus (Litopenaeus) vannamei, an experiment was conduced using information from two farm units of a Mexican hatchery and two shrimp population densities at each location. Data consisted of 12,658 shrimps that were siblings from 48 sires and 77 dams with a nested dam-sire structure. Shrimps were individually weighed at an average age of 130 days post-hatching. BW phenotypic mean (S.D.) was 18.2 (2.4) g, with values ranging from 8.4 to 30.0 g. Data were analyzed using univariate and multivariate models that considered BW within location by density pond environment as a different trait and included or not a common full-sib effect (c). The multivariate animal model included fixed effects of days from hatching and sex. For univariate models that included c effects, BW heritability (S.E.) estimates ranged from 0.24 (0.14) to 0.35 (0.18) across environments (heritability was zero in one environment). For multivariate models (excluding the environment with zero heritability) the heritabilities increased and ranged from 0.37 (0.06) to 0.45 (0.09). Standard errors of heritabilities and c effects were both drastically reduced in the multivariate analysis. Pairwise genetic correlations between environments were from 0.80 (0.08) to 0.86 (0.04). These differences may be indicative of genotype-environment interaction for BW at 130 days post-hatching. Statistical problems found to separate c from additive genetic effects both in univariate models were reduced using multivariate models. Correlation between family raw phenotypic means and family BV means from the multivariate analysis was 0.93 indicating a rather low risk of miss selecting superior families if BLUP solutions were neglected using replicated environment data. It is also concluded that use of incorrect statistical models or unreplicated data may lead to biased or inaccurate estimates of genetic parameters in shrimp breeding programs.     

Changes to vertical thermoregulatory movements of juvenile bigeye tuna (Thunnus obesus) in the northwestern Pacific Ocean with time of day,seasonal ocean vertical thermal structure,and body size

Vertical movements related to the thermoregulation were investigated in 12 juvenile bigeye tuna (Thunnus obesus) in Japanese waters using archival tag data. Movements changed with time of day, season, and body size. During daytime, bigeye tuna descended to greater depths, presumably to feed in the deep scattering layer (DSL). Thereafter, they repeatedly ascended to shallower layers, suggesting attempts at behavioral thermoregulation, although the beginning of vertical thermoregulatory ascents might reflect a shift in DSL depth. By the end of such movement, the whole‐body heat‐transfer coefficient might decrease because, although the depth and ambient temperature of the upper layers did not change, the body temperature gradually decreased significantly just after ascent for thermoregulation. Seasonal patterns indicated that the vertical thermal structure of the ocean might influence this ascent behavior. For example, from January to May, bigeye tuna made fewer ascents to less shallow waters, suggesting that they respond to increasing depths of the mixed surface layer by reducing energy expenditure during vertical migration. In addition, as body size increased, fewer thermoregulatory ascents were required to maintain body temperature, and fish remained deeper for longer periods. Thus, vertical thermoregulatory movements might change with body size as bigeye tuna develop better endothermic and thermoregulatory abilities. We hypothesize that bigeye might also increase cold tolerance as they grow, possibly due to ontogenetic shifts in cardiac function.