Spatio-temporal variability in fatty acid trophic biomarkers in stomach contents and muscle of Iberian sardine (<Emphasis Type="Italic">Sardina pilchardus</Emphasis>) and its relationship with spawning

Temporal variation in the fatty acid (FA) composition of stomach contents of Iberian sardines was compared to the relative contribution to dietary carbon made by different prey types for fish from two areas off Portugal. The effect of the FA content of the diet on sardine muscle FA composition was also studied, aiming at (1) analysing if FA biomarkers can be used as a complementary technique for the study of sardine diet and (2) to relate spatial and temporal variations of prey FA content with sardine condition and reproduction. Significant spatial differences in the FA composition of sardine diet occurred with concentrations of n-3 polyunsaturated FA, namely eicosapentaenoic acid [EPA, 20:5n-3] and linolenic acid 18:3n-3, being significantly higher in the diet of sardines from the west coast, whilst the diet of sardines from the south coast was richer in monounsaturated fatty acids (MUFA), namely the carnivory biomarker oleic acid 18:1n-9. These results are in agreement with the higher contribution made by diatoms and dinoflagellates to the diet of sardines off the west coast. Spatial variation in sardine dietary FA was also detected in their muscle composition, specifically for EPA, and the eicosapentaenoic/docosahexaenoic acid and (n-3)/(n-6) ratios, which were higher in sardines from the west coast. No difference in FA composition was detected between sexes, and the seasonal variability in sardine total FA concentration was primarily related to the seasonality of spawning. Sardines accumulate high concentrations of FAs during the resting stage of reproduction when the feeding intensity is similar or lower to that observed during the spawning season. Additionally, sardines show a high selective retention of MUFA and polyunsaturated FA (PUFA) throughout the year except at the beginning of the spawning season, when these FAs are largely invested in the formation of the gonads. Therefore, temporal and regional differences of prey environments are strong enough to be reflected in fish body composition, namely on the accumulation of essential FAs, which can have a strong impact on reproduction success for this species.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Evaluation of spawning frequency in a Mediterranean sardine population (<Emphasis Type="Italic">Sardina pilchardus sardina</Emphasis>)

The postovulatory follicle method was used to assess the frequency of spawning of sardine (Sardina pilchardus sardina) in the Central Aegean Sea (eastern Mediterranean) in November–December 1999 and 2000. A field-based aging key for postovulatory follicles (POFs) was developed, and preliminary evidence is presented of a temperature influence on the degeneration of POFs. Bias in the fraction of day-0 spawners (females that had, were, or would spawn the night of sampling) with respect to sampling time was identified and attributed to spawning behavior. An appraisal of consistency in spawners' fractions and sex ratio, based on four sets of sequential trawl hauls, showed a considerably high small-scale variability, which suggested that spawning schools are structures limited in space and/or ephemeral in time. Sampling gear, i.e. commercial purse seines and research pelagic trawls, did not differ significantly with respect to spawners' fractions. The fractions of day-1 and day-2 females were the same during both years, and their combination resulted in more precise, composite estimates of spawning frequency. In both years, the Mediterranean sardine spawned every 11–12 days. The incidence of spawning increased significantly with fish size. When compared to all estimates available for other sardine genera, species and subspecies, the spawning frequency of the Mediterranean sardine was lower; this was attributed to the synergetic effect of oligotrophy and smaller body sizes in the Mediterranean.     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Factors affecting the spawning period of sardine in two highly oligotrophic Seas

Ovarian histology and weight dynamics of reproductive and somatic tissues were used to investigate seasonality of spawning and factors affecting reproductive period in two E Mediterranean sardine, Sardina pilchardus, populations (central Aegean and Ionian Seas), between September 1999 and May 2001. Despite similarities in the general spawning pattern of the two populations, i.e. protracted spawning season during the colder months of the year, reproductive period in the Ionian was shorter and lagged behind the Aegean by 2–3 months. In both Seas, reproductive period was influenced by body size as smaller females tended to have both shorter and delayed spawning season. Whereas hepatosomatic index (HSI) was consistently higher in reproductively active females during the whole spawning period, seasonality of spawning did not match variations of HSI. In both Seas fluctuations of HSI matched the spring burst of primary productivity and were followed by fluctuations in somatic condition and mesenteric fat by a 2 months’ lag. Somatic condition and reproductive activity displayed inverse seasonal patterns, implying that egg production in sardine basically relies on visceral and muscular fat deposits, accumulated during the summer growing season. Reliance of egg production on past energetic reserves suggests that sardine is a capital breeder, which is a rather efficient strategy in such oligotrophic habitats as the E Mediterranean, where winter spawning is accompanied by a particular decline in adult preying fields.     

Asymptotic growth trajectories of larval sardine (Sardinops melanostictus ) in the coastal waters off western Japan

Growth trajectories of individual larvae of Japanese sardine, Sardinops melanostictus, caught in the coastal waters off western Japan were back-calculated from the first feeding stage up to date of capture (approximate size of 20 to 35 mm total length; TL) based on individually determined allometric relationships between otolith daily ring radii and fish total lengths. The larvae in January-, February-, and March-hatched cohorts in the coastal waters grew faster and more uniformly than those in the oceanic waters offshore of the Kuroshio current. Growth trajectories of the three hatch-month cohorts were similar and could be expressed by the Gompertz model. The inflection points of the growth curves were reached at 9 to 11 d after hatching, when larvae were 10.8 to 11.8 mm TL. Maximum growth rates at these points were 0.80 to 0.85 mm d⁻¹. Growth rates gradually declined after the inflection points, and larval TLs converged into the infinite length of 29 to 32 mm, the sizes at which metamorphosis from larvae to juveniles is initiated. This asymptotic growth pattern in the larval stage resulted in the narrow ranges in TLs in spite of the wide range of ages of the larvae caught by boat seiners in the coastal waters. Slow growth and therefore long duration of the metamorphosing stage could be influential in determining the cumulative total mortality in the early life stages of the Japanese sardine. Received: 14 July 1996 / Accepted: 20 August 1996     

Negative health effects in rats fed on sardine fishes

An adequate human diet should satisfy the requirements for energy and nutritive components including essential polyunsaturated fatty acids, essential amino acids, mineral components, vitamins, and fat. The benefits of the fish consumption are, however, often counteracted by toxic metals such as lead present in fish meat. Adverse effects of toxic metals on the aquatic environment and human health have aroused increasing interest in recent years. The achievement of an appropriate balance between the risks and benefits associated with fish consumption has therefore become a key health issue in current research. Therefore, we studied health impacts associated with the consumption of sardine fish, one of the most commonly consumed fish species throughout the world. Hepatic damages associated with the consumption of white or red muscle of sardine fish were explored and evaluated using a rat model. Rats were fed for 60 days with white or red sardine meat. Findings revealed (1) an elevated level of uric acid in the blood, (2) an accumulation of lead in the liver, (3) an atrophy of the liver, (4) an increase in plasma aspartate aminotransferase and alanine aminotransferase activities, and (5) an oxidative stress in the liver, including increased levels of lipid peroxidation and enhanced activities of superoxide dismutase, catalase, and glutathione peroxidase. Several abnormalities were also observed in liver histology. The alterations observed in the animals can be attributed to both the accumulation of lead and the high levels of purine in the sardine meat. The findings show that special attention should be given to the health effects associated with high intakes of sardine meat, particularly dark-meat sardine.     

Density-dependent migratory waves in the marine pelagic ecosystem

The migration of large aggregations of animals that sweep through the landscape is a phenomenon with large consequences in many ecosystems. It has been suggested that such migrations are mediated by resource depletion. Under this hypothesis it has been shown that simple foraging rules may generate density-dependent migratory waves (DDMW) in which the speed and amplitude increase with animal abundance. We tested these predictions on a 32-year data set of the spatial distribution of the two youngest age groups of a small pelagic schooling fish, the capelin (Mallotus villosus), by the end of their annual feeding migration in the Barents Sea. Our data suggest that the two age groups divided the Barents Sea by forming migratory waves that moved in opposite directions. The aggregation and spatial displacement of these waves increased with increasing age-specific abundance. However, possibly through social interactions, migratory pattern was modified by the abundance of the other age group.     

Seasonal migrations of plaice (Pleuronectes platessa) through the Dover Strait

Analysis of paired-haul mid-water trawling experiments conducted in the five months November to March during the years 1985 and 1988 revealed the seasonal pattern of migration of plaice (Pleuronectes platessa L.) moving through the Dover Strait between the Southern Bight of the North Sea and the English Channel. Mid-water catches of maturing fish were highest on south-going tidesprior to peak spawning (late January), while catches of spent fish, were highest on north-going tidesafter peak spawning. Detailed analysis of the catch data provided evidence that males complete their pre-spawning migration earlier, and their post-spawning migration later, than female fish and consequently spend more time on the spawning ground. Catches of immature female fish indicated a migration south following that of the mature fish. This dummy run may be a method by which immature plaice use the presence of high densities of mature and spawning fish to learn the location of the spawning ground. The analysis also indicated that differences in catch rates between day and night are unlikely to be the result of gear avoidance by day. The results confirm that selective tidal-stream transport is a key migratory mechanism for plaice in areas of fast tidal streams on the European continental shelf.     

Stratification and Tidal Current Effects on Larval Transport in the Eastern English Channel: Observations and 3D Modeling

We study how the combination of tides and freshwater buoyancy affects the marine organisms accumulation and horizontal transport in the ROFI system of the eastern English Channel. The Princeton Ocean Model coupled with a particle-tracking module is used to study the migration of fish eggs and larvae under different forcing conditions. Results of modeling are validated against observed concentrations of Flounder (Pleuronectes flesus) larvae. Numerical Lagrangian tracking experiments are performed with passive and active particles, representing sea-water organisms. Passive particles are neutrally buoyant whereas active particles are able to exercise light dependent vertical migrations equating to the swimming behavior of larvae. The experiments reveal that the strongest accumulation of particles occurs along the French coast on the margin of the ROFI. This happens because the interaction between the turbulence, the freshwater buoyancy input, and tidal dynamics, produces particle trapping and vertical spreading within the frontal convergence zone. Tides and freshwater input induce net alongshore horizontal transport toward the North. Tidal currents modulate the magnitude of horizontal transport whereas the fresh water input controls more the location of accumulation zones. Tracking experiments with active particles indicate that the vertical migration leads to a significant departure from the passive particle transport pattern. Differences lie in the shape of the particle transport pattern and the rate of the northward migration. In particular, vertically migrating particles travel slower. To find possible Flounder migration pathways, particles are released within the assumed spawning area of Flounder. The model predicts larvae drift routes and demonstrates that throughout the entire particle-tracking period the horizontal structure of the particle distribution is consistent with the larvae concentrations observed during the field experiments.     

Growth and ovarian development of Maurolicus muelleri during spring

The size, fecundity and gonad development of Maurplicus muelleri (Gmelin) were studied in Herdlefjorden, western Norway, from January to June 1994. In January, the two age groups formed separate sound-scattering layers (SSL). Juvenile 1-group fish were found to form the upper SSL, while adult 2+ individuals inhabited the lower, these distributions reflecting the different feeding to fitness functions of these two age groups. In June there was only one SSL, and in March and May some 2+ were also found in the upper SSL. Growth of the younger fish to 2+ fish size was observed over the period, and it was found that M. muelleri mature after 1 yr, although size at maturity was lower than found in previous studies. Growth of the 2+ fish changed as the period progressed, to allow gonadal development after a winter of low feeding. Increased investment in development of the gonads was also observed in the 1-group fish over the study period, but substantial maturation only occurred from May to June. Absolute fecundity was size dependent, although high variability was found in the larger size ranges. Oocyte development occurred in discrete batches, although lack of gonadal synchrony between individuals means that spawning is continuous in the population as a whole, once the spawning period begins. Batch fecundity was found to be similar to previous studies, and 2+ fish were found to start spawning earlier, and with an increased intensity, than the 1-group fish.     

The energetics and evolution of intraspecific predation (egg cannibalism) in the anchovy Engraulis capensis

Anchovy feeding activity (based on samples collected around the South African coast between 1986 and 1988) seems to peak twice daily, with the major peak between 17:00 and 19:00 hrs. Mean daily consumption in the field was estimated at 3.45% of body wt d^-1. Since the various food types available to anchovy differ in their caloric values, this figure might be expected to vary according to the type of diet available to the fish. To obtain a diet energetically equivalent to that measured in the field anchovy feeding exclusively on phytoplankton would need to consume the equivalent of 5.8% of their body wt d^-1 and those feeding on zooplankton 3.5 to 4.5% of the body wt, while fish feeding on their own eggs would require only 2.5% of their body wt daily. Feeding on eggs at peak spawning on the spawning grounds —where there is limited alternative food (i.e., zooplankton) available — seems adequate to support reproduction. In selection at the population level, cannibalistic fish benefit from homeostatic self-regulation.     

Concentration of ascorbic acid and innate immune effectors in <Emphasis Type="Italic">Engraulis ringens</Emphasis> and <Emphasis Type="Italic">Strangomera bentincki</Emphasis> during their main spawning period (2007–2008) in the Humboldt current system off Chile

In this study, we determined variations in ascorbic acid (AA) concentrations in the gonads and liver, and of the innate immune response (lysozyme activity and antiprotease) in the plasma of Engraulis ringens (anchoveta) and Strangomera bentincki (common sardine) during their reproductive seasons of 2007 and 2008 in the central area of the Humboldt Current. During the main spawning season in 2007, colder environmental conditions, higher phytoplankton biomasses, low dinoflagellate abundance and the dominance of large copepods in winter-early spring and of small copepods in late spring contributed to explain higher AA concentrations in anchoveta tissues. During the warmer year of 2008, lower phytoplankton biomasses, a notable increase of dinoflagellate abundance and the dominance of small-size copepods in winter and of larger copepods in spring occurred along with a drastic drop in AA concentrations in anchoveta tissues. These results contrasted with those in common sardine, a species in which AA did not vary as much as in anchoveta. An inverse trend between AA concentration in the liver and the gonadosomatic index suggested the AA utilization during gonadic maturation. Innate immune parameters were determined for the first time in small pelagic fish and constitute a new baseline data to evaluate their natural response to environmental changes. Changes in the distribution and abundance of edible taxa containing AA and edible for fish along with variations in AA in fish tissues during their reproductive season could become good indicators of the overall fish physiological condition resulting from variations in the trophic web structure.     

Ephemeral spawning aggregations in the Mediterranean sardine, <Emphasis Type="Italic">Sardina pilchardus</Emphasis>: a comparison with other multiple-spawning clupeoids

Using previously published histological data on multiple, monthly samples of Sardina pilchardus collected in the central Aegean and Ionian Seas (September 1999–August 2000, and November 2000–February 2001), the Mediterranean sardine was treated as a case study to investigate the biological characteristics of ephemeral spawning aggregations in multiple-spawning clupeoids. Actively spawning (Day₀) females in the Mediterranean sardine, i.e., the daily class of spawners caught a few hours prior, during, or after the spawning act, were shown to separate spatially from late (Day₁₊) spawners and non-spawning females, taking with them a large proportion of conspecific males which were also in advanced spawning condition and in better somatic condition compared to the remaining population. In addition, information from 28 stocks of multiple-spawning clupeoids from a wide range of geographic locations, belonging to 14 species and 2 families (Engraulidae and Clupeidae), was reviewed and analyzed pertinent to the formation of ephemeral spawning aggregations. Results from the latter analysis indicated similar patterns of spatial segregation of Day₀ spawners in the reviewed clupeoid stocks as in the Mediterranean Sardine, which strongly suggested that the formation of ephemeral spawning aggregations is a common behavioral trait among multiple-spawning clupeoids. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Calculation of search volume on cruise-searching planktivorous fish in foraging model

Search volume of cruising planktivorous fish was calculated based on its detailed behavior Th examine the factors influencing search volume, a series of experiments were conducted by varying ambient conditions, such as structural complexity light intensity and turbidity Pseudorasbora parva were used in experiment as predator and Daphnia pulex was selected as prey The shape of scanning area of P parva showed elliptic and the search volume changed drastically depending on ambient conditions. Compared with the results of previous foraging model, the search volumes of the fish under previous study were larger (1.2 to 2.4 times) than those from our study These results on the changes in feeding rate can be useful in determining microhabitat requirement of P parva and othercyprinids with a similar foraging behavior The calculated search volume is compared with other foraging model andthe effect of zooplankton-planktivore interactions on aquatic ecosystem is discussed.     

Using multistate mark-recapture methods to model adult salmonid migration in an industrialized river

A multistate mark-recapture (MSMR) model of the adult salmonid migration through the lower Columbia River and into the Snake River was developed, designed for radiotelemetry detections at dams and tributary mouths. The model focuses on upstream-directed travel, with states determined from observed fish movement patterns indicating directed upstream travel, downstream travel (fallback), and use of non-natal tributaries. The model was used to analyze telemetry data from 846 migrating adult spring-summer Chinook salmon (Oncorhynchus tshawytscha) tagged in 1996 at Bonneville Dam on the Columbia River. We used the model to test competing hypotheses regarding delayed effects of fallback at dams and visits to tributaries, and to define and estimate migration summary measures. Tagged fish had an average probability of 0.755 () of ending migration at a tributary or upstream of Lower Granite Dam on the Snake River, and a probability of 0.245 () of unaccountable loss (i.e., mortality or mainstem spawning) between the release site downstream of Bonneville Dam and Lower Granite Dam. The highest probability of unaccountable loss (0.092; ) was in the reach between Bonneville Dam and The Dalles Dam. Study fish used the tributaries primarily as exits from the hydrosystem, and visits to non-natal tributaries had no significant effect on subsequent movement upriver (P = 0.4245). However, fallback behavior had a small effect on subsequent tributary entry and exit (P = 0.0530), with fish using tributaries as resting areas after reascending Bonneville Dam after fallback. The spatial MSMR model developed here can be adapted to address additional questions about the interaction of migrating organisms with their environment, or for the study of migrations in other river systems.     

Spawning migrations and local movements of a tropical sparid fish

Between June 1994 and September 1996, a mark–recapture study was conducted in the Hinchinbrook Channel area of tropical northeastern Australia to investigate the movements of the sparid fish Acanthopagrus berda. A total of 962 A. berda were tagged, 922 within a single estuary system, Deluge Inlet. No movements among estuaries were detected. Of the fish tagged in Deluge Inlet, 12.9% were recaptured. Two movement regimes were detected: local movements (mostly <500 m) during non-spawning times, and longer migrations (up to 3.12 km) during spawning seasons, to and from a spawning site at the estuary mouth. Although there appeared to be a major spawning aggregation in the mouth of Deluge Inlet, ripe female A. berda were present in upstream sites, indicating that spawning probably occurs there also. This contrasts to the situation in South Africa, where A. berda is reported to spawn only close to estuary mouths. Differences such as this highlight our limited knowledge of variations in the reproductive biology of widespread species such as A. berda. Received: 25 November 1997 / Accepted: 10 September 1998     

Annual reproductive cycle of the captive female Japanese sardineSardinops melanostictus: Relationship to ovarian development and serum levels of gonadal steroid hormones

Gonad and blood samples were taken from the captive female Japanese sardineSardinops melanostictus between 1988 and 1989, and changes in serum levels of gonadal steroids were correlated with the annual gonadal cycle. Under captive conditions, female fish did not mature and spawn spontaneously, although oocytes developed up to the end of vitellogenic growth. Based on evidence from ovarian histology, the annual gonadal cycle of the Japanese sardine was divisible into four periods, i.e., immature (June to October), vitellogenesis (November to December), spawning (January to March), and post-spawning (April to May). The pattern of seasonal change in the gonadosomatic index (GSI) showed an inverse correlation to change in water temperature and reflected the degree of ovarian maturity. The serum estradiol-17 level increased from its lowest concentration (0.12 ng ml^–1) in September to a peak (1.14 ng ml^–1) in March. Serum 17,20-dihydroxy-4-pregnen-3-one (17,20-P) was detectable at low levels (<0.3 ng ml^–1) between October and February, but was below the assay detection limit (0.06 ng ml^–1) at all other times. Testosterone was not detectable (<0.06 ng ml^–1) in the serum of any fish throughout the year. The effects of several steroids on the maturation of follicle-enclosed oocytes of sardine were examined in vitro, and 17,20-P was found to be the most potent inducer of maturation. This suggests that post-vitellogenic oocytes of the Japanese sardine in captivity have an ability to respond to an appropriate hormonal effector and subsequently to resume meiotic maturation.