Sardine Larvae Research Articles

Feeding ecology of sprat (Sprattus sprattus L.) and sardine (Sardina pilchardus W.) larvae in the German Bight, North Sea

The abundance of the sardine in the North Sea suddenly increased after 1995. Since 2002, the sardine has been spawning regularly in the German Bight, and all its life stages can be found in the area. The larval feeding ecology of two small pelagic clupeiform species with very similar life histories was investigated, the particular aim being to determine signs of food overlap. The distribution and feeding of sprat and sardine larvae were investigated during late spring 2003 on two transects * This work was conducted within the framework of the GLOBEC Germany project, with financial support from BMBF (03F0320C). Thecomplete textof thepaperis available athttp://www.iopan.gda.pl/oceanologia/

Larval fish abundance and distribution during the late winter bloom off Gran Canaria Island, Canary Islands

AbstractThe species composition, distribution and abundance of fish larvae off Gran Canaria (Canary Islands), an oceanic island in the NE Atlantic, were studied from January to August 2005. Weekly samplings were carried out at six stations in the eastern and southern flanks of the island, at the edge of the island shelf, in order to analyze the fluctuations of the planktonic community. The highest chlorophyll a values were recorded in March, coinciding with the lowest values of temperature in the mixed layer, which gave rise to the ‘late winter bloom’. Mesozooplankton biomass peaked with only a week‐lag to phytoplankton, and its average value (13.5 ± 10.9 SD mg dry weight m−3) was typical for the area. A clear windward‐leeward distribution pattern was found for small mesozooplankton and neritic larvae, showing higher values in two stagnation points, upstream and downstream of the island. A total of 128 taxa of fish larvae were identified. Neritic and oceanic larvae appeared in quite similar proportions. Only three families accounted for half of the total larval fish collected: Myctophidae (24.9%), Sparidae (12.7%) and Clupeidae (11.9%). Sardinella aurita (8% of total larvae collected) was the most abundant species, appearing during the whole period of study and at each of the six sampled stations. Sardina pilchardus larvae were rarely captured but were always encountered with the arrival of NW African upwelling filaments to the island coast, suggesting that these larvae were transported in those mesoscale structures.

Temporal patterns of growth in larval cohorts of the Japanese sardine Sardinops melanostictus in a coastal nursery area

Growth patterns of larval sardine Sardinops melanostictus were studied in a coastal nursery area, in southern Japan for four monthly hatch cohorts of larvae (November, December, January and February) for the 2003–2004 and 2004–2005 seasons. Laird–Gompertz models were fitted to each cohort using both total length (LT)‐at‐age at capture and mean LT‐at‐age data derived from backcalculations. In both approaches, the absolute daily growth rates (GR) and absolute daily growth rates at the inflection point (GXO) were estimated. In parallel, individual growth rates (GI) were derived from backcalculated LT (LB). Growth showed the following general common patterns irrespective of hatch month, season and methods: (1) significant Laird–Gompertz fits, (2) asymptotic growth, (3) a decrease in GR after the inflexion point, except for February for the 2003–2004 season that showed an apparent constant growth pattern, (4) six in eight cohorts showed GXO ranging from 0·8 to 1·2 mm day−1 and (5) a decreasing tendency of GI from 1·75 to 0·24 mm day−1, from first feeding through the first month of larval life. The contrasting pattern between the 2003–2004 and the 2004–2005 seasons were: (1) allometric v. logarithmic (ln) LT and otolith radius relationships, (2) low GXOv. high GXO, (3) high GRv. low GR when growth turned asymptotic, (4) low GXOv. high GXO when monthly hatch cohorts were combined and (5) LB and GI not differing among monthly hatch cohorts. The differences in growth patterns and growth rates between seasons seemed to be linked to the influx of warmer and oligotrophic waters of the Kuroshio Current that triggered an increase of 3° C in the coastal area for the 2003–2004 seasons. In the overall context, however, the high GXO, within cohorts and seasons reported in the current study, suggests that either sea surface temperature (SST) or food availability, or both are in the optimal range of preferences for S. melanostictus larvae. Consequently, nearshore coastal areas seem to be playing an important role as a nursery area for the larval stage of this species.

Japanese sardine (Sardinops melanostictus) mortality in relation to the winter mixed layer depth in the Kuroshio Extension region

AbstractA drastic population change in Japanese sardine (Sardinops melanostictus) has been noted as being related to winter sea surface temperature (SST) in the Kuroshio Extension region. The former studies suggest two possible explanations. One is that temperature itself affects sardine. The other is that SST represents the environmental change of the Kuroshio Extension region and other causes directly affecting sardine. In this study, we found that sardine mortality from post‐larva to age 1 negatively correlated with the winter mixed layer depth (MLD) in the Kuroshio Extension region from 1979 to 1993. During the period of a deep winter mixed layer (during the early 1980s), sardine mortality was low, whereas mortality was high when the winter mixed layer was shallow (during the late 1980s to early 1990s). By using a lower trophic‐level ecosystem model forced by the observed time series of MLD, SST, light intensity and nutrient data, we found that the estimated spring zooplankton density drastically varies from year to year and has a significant negative correlation with sardine mortality. The inter‐annual variation of spring zooplankton density is caused by the winter MLD variation. During the deep winter mixed layer years, a phytoplankton bloom occurs in spring, whereas during the shallow winter mixed layer years, the bloom occurs in winter. The results of our study suggest that the decline in the Japanese sardine population during the late 1980s to early 1990s was due to an insufficient spring food supply in the Kuroshio Extension region where sardine larvae and juvenile are transported.

Spawning, larval abundance and growth rate of Sardinops sagax off southwestern Australia: influence of an anomalous eastern boundary current

The temporal and spatial distributions of sardine Sardinops sagax eggs and larvae off the oligotrophic southwestern coast of Australia were examined and related to gonadosomatic index, daily growth rates of larvae and regional biological oceanography. Seasonal environmental cycles were established from remotely sensed sea surface temperature and chlorophyll concentration, wind and sea surface height data. Sardine egg and larval distributions were determined from regular transect surveys and annual grid surveys. Sardine eggs and larvae were common across the continental shelf throughout the year between Two Rocks and Cape Naturaliste (~32 to 34° S), and gonadosomatic index data suggested a distinct winter peak in spawning activity. Surface chlorophyll concentrations were highest during winter, coincident with the seasonal peak in the southward flow of the Leeuwin Current along the continental shelf break. Retention conditions on the mid-outer shelf for pelagic eggs and larvae were therefore poor during this time. Egg and larval concentrations were lower than expected in winter and higher in summer when retention conditions were more favourable. Larval sardine growth rates were unexpectedly high, averaging 0.82 mm d -1 . Fisheries for clupeiod species off southwestern Australia are insignificant compared to other eastern boundary current systems. Our data suggest that this may be due to a combination of low primary productivity caused by suppression of large-scale upwelling by the Leeuwin Current and the modest seasonal maximum in primary productivity occurring during the time least favourable for pelagic larval retention.

Growth effect on the otolith and somatic size relationship in Japanese anchovy and sardine larvae

Relationships between otolith and somatic sizes were examined for Japanese anchovy Engraulis japonicus and sardine Sardinops melanostictus larvae collected broadly in the western North Pacific, based on a substantial data set derived from a previous paper. Allometric formulae showed close fits to the relationships between otolith radius and standard length, and the formulae differed between anchovy and sardine larvae. Despite the high correlations, the effect of somatic growth rate on the otolith and somatic size relationship (the ‘growth effect’) was significantly detected for both anchovy and sardine larvae. Slower growing larvae tended to have larger otoliths than faster growing conspecifics at the same somatic size. This growth effect was more obvious for sardine larvae than for anchovy larvae, probably because of their differential responses of somatic growth to temperature shifts. The growth effect could lead to the possibility of biases in the backcalculation and size estimation processes. As the growth effect is considered to be a general phenomenon and its extent to be species-specific, the relationship between otolith and somatic size and its uncoupling should be scrutinized before application of techniques based on the otolith and somatic size correlation.

Transport and survival of larvae of pelagic fishes in Kuroshio system region estimated with Lagrangian drifters

Transport and survival of larvae of pelagic fishes in the Kuroshio system region were studied using Lagrangian drifter data recorded from 1990 to 2003. A large portion of the drifters from the Kuroshio area south of Japan spread around the Kuroshio Extension up to 170°E, while some moved south to the offshore area of the Kuroshio because of a recirculation gyre. The monthly mean eastward movement from areas south of Japan was approximately 800-900 km, which was smaller than previous numerical estimates. The results of a survival model assuming optimal temperatures for larvae suggest that surface waters during the observation period were too warm for larval Japanese sardine, which has an optimal temperature of 16°C, and the adult abundances did not increase during the observation period. In contrast, the spawning ground temperatures and transport conditions from an area south-west of Japan in April-June matched the requirement for the larval Japanese anchovy, which has an optimal temperature of 22°C. The combined effects of temperature variations due to seasonality and water mass mixing are suggested to play an important role in determining the environmental temperatures that occur in an area.

Importance of scale to the relationship between abundance of sardine larvae, stability, and food

We used a spatially explicit regression model to relate sardine (Sardinops sagax) larval abundance to water column stability, phytoplankton, and zooplankton off South Australia. The distribution of sardine larvae was significantly associated with stability (p < 0.05) and phytoplankton fluorescence or zooplankton displacement volume (p < 0.001) at broad scales. In contrast there was no relationship between sardine larvae, stability, phytoplankton, or zooplankton at medium or fine scales. The relationships are shown to be scale‐dependent. Although the results generally support Lasker’s (1978) “stable ocean” hypothesis, mid‐range rather than high stability appears to be favored, in accord with theoretical expectations at both large and small scales.

Optimal growth temperature hypothesis: Why do anchovy flourish and sardine collapse or vice versa under the same ocean regime?

The out-of-phase population oscillations between anchovy and sardine have been attributed to climate changes. However, the biological processes causing these species alternations have remained unresolved. Here we propose a simple "optimal growth temperature" hypothesis, in which anchovy and sardine regime shifts are caused by differential optimal temperatures for growth rates during the early life stages. Dome-shaped relationships between growth rate and sea temperature were detected for both Japanese anchovy (Engraulis japonicus) and Japanese sardine (Sardinops melanostictus) larvae based on otolith microstructure analysis. The optimal growth rate for anchovy larvae occurred at 22.0 °C, whereas that for sardine larvae occurred at 16.2 °C. Ambient temperatures have historically fluctuated between these optima, which could lead to contrasting fluctuations in larval growth rates between the two species. This simple mechanism could potentially cause the shifts between the warm anchovy regime and the cool sardine regime in the western North Pacific. Although retrospective analysis suggested synergistic effects of other factors (e.g., trophic interactions and fishing), the optimal growth temperature concept would provide a possible biological mechanism of anchovy and sardine regime shifts.

Seasonal and inter-annual changes in the planktonic communities of the northwest Alboran Sea (Mediterranean Sea)

The response of the northwestern Alboran Sea pelagic ecosystem to temporal changes in hydrological conditions has been examined for the time period of 1992–2002. In addition, the bottom–up linkages between the lower trophic levels and the growth and nutritional status of sardine larvae were examined using quarterly data from 1992 to 2002 within the frame of the monitoring Project ECOMALAGA. The study area was characterised by the almost permanent presence of an upwelling which was intensified in the spring period. Consequently, an annual peak of nutrients was usually found during this season when the nitrate concentration averaged 1.35 μM. Accordingly, chlorophyll- a concentration and cell abundance of micro- plus nano-phytoplankton increased in that season (1.51 μg L −1 and 446 cell mL −1 compared to 0.85 μg L −1 and 225 cell mL −1 obtained from summer to fall). Despite these seasonal changes, the analysis of the taxonomic composition of the phytoplankton communities did not reveal a clear annual succession pattern. Contrastingly, peaks of zooplankton abundance were obtained in summer (1964 ind m −3) due to the increased presence of brachiopods with respect to copepods (which dominated from fall to spring). Significant inter-annual changes were obtained in the phytoplankton and zooplankton communities. Thus, dinoflagellate and coccolitophorid abundances relative to diatom abundances tended to increase from 1997 to 2002. This trend matched the progressive reduction of the upwelling intensity. These inter-annual changes significantly affected the larval growth of Sardine pilchardus and their nutritional condition, as higher growth rates in terms of body length coupled to higher somatic mass increases (expressed by DNA content) occurred in spring, matching with the higher chlorophyll- a concentration. Furthermore, the highest larval growth was obtained in 2001, coinciding with the change observed in the composition of phytoplankton community.

Egg and larval distributions of seven fish species in north‐east Atlantic waters

ABSTRACTThe distribution of egg and larvae of mackerel, horse mackerel, sardine, hake, megrim, blue whiting and anchovy along the European Atlantic waters (south Portugal to Scotland) during 1998 is described. Time of the year, sea surface temperature and bottom depth are used to define the spawning habitat of the different species. Mackerel, horse mackerel, and sardine eggs and larvae presented the widest distribution, whereas megrim and anchovy showed a limited distribution, restricted to the Celtic Sea and the Bay of Biscay respectively. Correspondingly mackerel, horse mackerel and sardine showed the highest aggregation indices. Blue whiting larvae were found at the lowest temperatures, whereas anchovy eggs and larvae were found in the warmest waters. The analysis is a basis for evaluation of ongoing changes in the pelagic ecosystem of the north‐east Atlantic.

Physical–biological interactions in the life history of small pelagic fish in the Western Iberia Upwelling Ecosystem

This paper reviews relevant investigations conducted in the Western Iberia Upwelling Ecosystem (WIUE) on the dynamics of small pelagic fish (SPF), its relationship to climate changes and processes related to the ‘ocean triad’ (enrichment, concentration, transport/retention). In the last decades, a decline in the productivity of several SPF species (e.g., sardine and horse mackerel) was observed in the WIUE, which is partially explained by environmental variability (e.g., changes in coastal upwelling). The main mechanism proposed to explain this decline is the increased frequency and intensity of upwelling events during the spawning season (winter) of these species, which is typically a period when convergence conditions prevail. Thus, this ‘anomalous’ situation promotes egg and larval dispersal away from the favourable coastal habitat (the shelf) and consequently has a negative impact on their survival and recruitment. However, the variability of local features like the Western Iberia Buoyant Plume (WIBP) and the Iberian Poleward Current (IPC) introduce important fluctuations in the transport patterns of the region, and could modulate the impact of these winter upwelling events in the survival of larvae. The retention on the shelf of larval sardine along convergence zones formed by the interaction of these two features plays an important role in their survival. Furthermore, the WIBP is a suitable environment for the growth of phytoplankton and for larval fish survival. From these findings it is clear that simplified Ekman transport models could not explain larval fish transport/retention patterns in this region and more realistic biophysical models should be used to simulate the local oceanographic conditions to understand larval dynamics and the success of recruitment of SPF in the Western Iberia.

Infection by a Protozoan Endoparasite of the genus Ichthyodinium in Embryos and Yolk-sac Larvae of Yellowfin Tuna Thunnus albacares

In October 2005, a mass mortality of yolk-sac larvae of yellowfin tuna Thunnus albacares due to an endoparasite infection occurred during the seed production in a semi-closed system tank in Bali, Indonesia. Prevalence of the infection reached 100% maximally. The affected embryo or yolk sac of hatched larvae was filled with numerous parasites by 24 h post spawning. The larvae died due to a burst of the yolk sac. Phylogenetic analysis with 18S rRNA sequence and morphological characteristics of the parasite indicated that the present parasite was identified as a protozoan endoparasite Ichthyodinium chabelardi infecting the yolk-sac larvae of Atlantic sardine Sardina pilchardus or as its closely related species. PCR with a primer set designed from the 18S rRNA sequence detected the genome of the parasite from infected fertilized eggs, yolk-sac larvae and rearing water but not from the gonads of broodstock or feed fish for broodstock. The infection was not observed when the spawned eggs were immediately transferred into sterilized seawater, but the eggs kept in the rearing water for more than 1 h were infected by the parasite. These results suggest that the parasite infects fertilized eggs horizontally through the rearing water.

The application of adaptive cluster sampling for rare subtidal macroalgae

Adaptive cluster sampling (ACS) is a targeting sampling method that provides unbiased abundance estimators for populations of rare species that may be inadequately sampled with simple random sampling (SRS). ACS has been used successfully to estimate abundances of rockfish and sardine larvae from shipboard surveys. In this study, we describe the application of ACS for subtidal macroalgae. Using SCUBA, we measured abundances of Codium mamillosum, C. pomoides, and Halimeda cuneata at three islands and two levels of wave exposure. The three species were relatively patchy and could be sampled with ACS at one site per dive. Their distributions differed among islands and with exposure to wave energy, with H. cuneata found at only one island. ACS is a useful tool for understanding the spatial distribution and abundance of populations of rare benthic species, but, as was the case in this study, may not be as efficient as sampling with SRS with comparable replication.

Field comparison of sardine post-flexion larval growth and biochemical composition from three sites in the W Mediterranean (Ebro river coast, bays of Almería and Málaga)

Late larval stages of sardine (16-23 mm) were sampled during the 2003 spawning season in their nursery grounds located off the Ebro river mouth, on the Catalan coast, and in two bays of the Alboran Sea coasts, the Bay of Almeria and the Bay of Malaga. The daily growth analysis of each sampled population revealed faster growth in the Ebro sardine larvae than in both of the Alboran Sea larval populations. This fact is supported by their greater content with age of DNA, RNA and protein. However, the significantly higher carbohydrate content of the Bay of Almeria sardine larvae and a higher Fulton’s index, indicative of energy storage of individuals, in both of the sardine populations sampled in the Bays of Almeria and Malaga show evident differences in the daily growth of the Alboran Sea larvae from those originating in the Ebro region. Late larval growth in the Alboran Sea sardine tends to favour an increase in body mass rather than in body length. This study hypothesises that the productivity pulses off the Alboran Sea coasts induced by the north and northwestern wind regimes may be responsible for the growth pulses observed in the otolith microstructure.

Postflexion larvae and juvenile daily growth patterns of the Alborán Sea sardine (<em>Sardina pilchardus</em> Walb.): influence of wind

The Alboran Sea is a hydrographically complex and variable area in which wind-driven coastal upwelling occurs. Its most abundant small pelagic resource is the sardine ( Sardina pilchardus ), which is subject to major interannual oscillations. Postflexion stages of sardine larvae were sampled in their main nursery grounds in the north Alboran Sea, and sardine juveniles were sampled periodically throughout the recruitment season. Daily growth analysis was used to identify periods favourable for larval survival and to assess the evolution of daily increment widths during the first months of life. The results showed that juveniles born later in the spawning season grew relatively faster than those born earlier. Two main growth phases were observed in juveniles: an initial one in which daily increment widths increased progressively, and a second one in which widths fluctuated, showing a decreasing trend. The beginning of the second phase was almost synchronous among different sub-cohorts, suggesting that it was triggered by environmental factors. The estimated mean daily growth rates during the larval phase were higher in surviving juveniles than in postflexion larvae born in the same month, supporting the “bigger is better” hypothesis in relation to larval survival. The influence of the environmental regime on growth was explored, and the evolution of a growth-related index, derived from otolith increment width variability, was compared with a single environmental descriptor, a “wind index” based on wind stress and direction. This analysis suggested that larval survival and larval and juvenile growth rates showed a positive correlation with NW component winds, associated with upwelling events on the continental shelf and calm sea weather conditions in the inshore nursery grounds. Conversely, Levantine and southern wind periods, which interrupt the upwelling and create rough seas in the inshore bays, lead to a decrease in growth rates in juveniles and low larval survival.

Analysis of feeding of <i>Sardina pilchardus</i> (Walbaum, 1792) larval stages in the central Cantabrian Sea

SUMMARY: Diet analysis of larval fish may clarify the role that zooplankton—not only its abundance but also its size structure and taxonomic composition—may have on larval growth and subsequent recruitment levels. Taking into account the increase in size during larval stages, we followed a larval size-dependent approach for the analysis of prey size in the diet of larval sardine (Sardina pilchardus). Studies on feeding patterns of clupeid larvae typically involve a problem due to regurgitation and defecation of gut contents during capture. Therefore, an alternative sampling method was tested in this study, but no significant differences from conventional methods were found. Despite the low feeding incidence observed (23%), we found a circadian feeding pattern with the highest mean gut contents after dawn, decreasing during the day and with the lowest values at night. Diet was mostly composed of copepod developmental stages, mainly nauplii, and prey size increased with larval size following a power function. Maximum and mean prey size were related to larval mouth gape, though other factors may be restricting the maximum prey size ingested, since most prey width values were between 20 and 40% of larval mandible width.

Vertical distribution of the European sardine (Sardina pilchardus) larvae and its implications for their survival

This study presents results of the vertical behaviour of the European sardine (Sardina pilchardus) larvae as observed at sea off the NW Iberian coast during an oceanographic cruise conducted in May 2002. Samples were taken in a grid of 38 stations (conductivity-temperature-depth [CTD] measurements and Longhurst Hardy Plankton Recorder [LHPR] plankton hauls); a 69-h fixed station study was also performed (hourly CTD measurements and LHPR/neuston hauls every 2 h). The horizontal distribution of larvae is closely related to the circulation patterns measured by a current metre-mooring array deployed during the cruise. Larvae were mainly distributed in the upper 20-25 m of the water column, in evident association with the waters of the Western Iberia Buoyant Plume (WIBP). Large (older) larvae are found mainly in the surface layers, and larval size decreases with depth. A diel rhythm of migration to the neuston layer was observed, correlated with the inflation/deflation activity of the swim bladder. Larvae with lengths greater than 12.5 mm and inflated swim bladders were only found in this layer. Considering the near surface stratification conditions for food availability and Ekman transport in the upper few metres, even small vertical migrations of larvae can be very important for their survival and subsequent recruitment success.

Annual cycle of clupeiform larvae around Gran Canaria Island, Canary Islands

AbstractThe distribution and abundance of fish larvae was studied along the eastern and southern shelf of Gran Canaria Island (Canary Islands) from July 2000 to June 2001. Oblique bongo hauls were carried out fortnightly during the daytime, coinciding with days of full and new moon. During February, the area was sampled every 2–5 days. About 17.3% of the ichthyoplanktonic community was composed of clupeiform larvae: 92.9% of these larvae were Sardinella aurita, whereas 4.7% and 2.4% were respectively Engraulis encrasicolus and Sardina pilchardus. Sardinella aurita larvae appeared during the whole year with two periods of maximum abundance: June to September and December to February. During the full moon their abundance was on average 38.5% (±6.8%) of their numbers during the new moon, showing a clear lunar periodicity. Engraulis encrasicolus larvae appeared from November to March, also coinciding with the new moon. Sardina pilchardus larvae only appeared during two short periods, both coinciding with filaments shed from the African coastal upwelling which reached the island. This fact confirms the transport of fish larvae from the upwelling area off northwest Africa to the Canary Islands, promoting a genetic flow among both sites.

Link between environmental anomalies, growth and condition of pilchard Sardina pilchardus larvae in the northwestern Mediterranean

Relationships between environmental factors and the growth and condition of pilchard Sardina pilchardus (Walbaum) larvae were studied in November 1998 in the northwestern Mediterr- anean (NWM). Long-term growth of fish was assessed using otolith analysis and condition was assessed through histological and biochemical (RNA/DNA) indices. These measurements were con- ducted on larvae belonging to 2 station groups (A and B) previously defined through principal com- ponent analysis (PCA) on 7 environmental variables. Correlations between individual condition indices and environmental values at place of capture were also performed. Condition in environmen- tal group A was significantly higher than in group B as assessed by both histological indices and RNA/DNA. Long-term growth, however, did not differ significantly between groups except for the otolith diameter vs. standard length (SL) relationship. The stations conforming to environmental group A were characterised by environmental ranges that would form a favorable habitat for larval success, including surface temperatures 4500 nauplii (N ) m -3 or >5500 of other microzooplankters (T-N ) m -3 . Apart from the good agreement found between histological and RNA/DNA results, each approach offered different information: histological indices detected the largest portion of unfit larvae at size-ranges ≤8 mm, whereas RNA/DNA best discriminated large lar- vae. The results from this study support the view that links exist between mesoscale anomalies, spawning intensity and growth or condition of pilchard larvae in the NWM.