Study of sardine (Sardina pilchardus) regime shifts in the Iberian Atlantic shelf waters

Sardine fisheries in the Iberian Atlantic shelf (36°N–44.5°N) show decadal‐scale cycles. In the late 1990s, a positive phase in sardine stock was expected; on the contrary, catches have declined until now. Regime shifts in climatic and oceanographic variables on different scales (as forcing factor) and shifts in sardine stock (as result) have been used with the aim of identifying the physical variables that explain most of the sardine population variance in the region. Circa 1998, when last sardine regime shift was detected, the main patterns of large‐scale atmospheric circulation in the Northern Hemisphere with influence in the study area namely Northern Atlantic Oscillation (NAO) and East Atlantic (EA) pattern changed and coupled in a combination that led to a rise in sea surface temperature and a decline in the coastal upwelling intensity. Several years with a downwelling situation in average in the main spawning and feeding Iberian sardine areas would have affected the stock abundance, averting the return to the projected positive regime. The sardine negative regime shift was detected first in the regions of the study located further north. The regional variable latent heat flux that groups a set of environmental processes related to the ocean–atmosphere heat exchanges and so with the turbulence manages to explain the 72% of sardine recruitment.     

Impact of freshwater input and wind on landings of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in shelf waters surrounding the Ebre (Ebro) River delta (north-western Mediterranean)

Time series analyses (Box–Jenkins models) were used to study the influence of river runoff and wind mixing index on the productivity of the two most abundant species of small pelagic fish exploited in waters surrounding the Ebre (Ebro) River continental shelf (north‐western Mediterranean): anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus). River flow and wind were selected because they are known to enhance fertilization and local planktonic production, thus being crucial for the survival of fish larvae. Time series of the two environmental variables and landings of the two species were analysed to extract the trend and seasonality. All series displayed important seasonal and interannual fluctuations. In the long term, landings of anchovy declined while those of sardine increased. At the seasonal scale, landings of anchovy peaked during spring/summer while those of sardine peaked during spring and autumn. Seasonality in landings of anchovy was stronger than in sardine. Concerning the environmental series, monthly average Ebre runoff showed a progressive decline from 1960 until the late 1980s, and the wind mixing index was highest during 1994–96. Within the annual cycle, the minimum river flow occurs from July to October and the wind mixing peaks in winter (December–April, excluding January). The results of the analyses showed a significant correlation between monthly landings of anchovy and freshwater input of the Ebre River during the spawning season of this species (April–August), with a time lag of 12 months. In contrast, monthly landings of sardine were significantly positively correlated with the wind mixing index during the spawning season of this species (November–March), with a lag of 18 months. The results provide evidence of the influence of riverine inputs and wind mixing on the productivity of small pelagic fish in the north‐western Mediterranean. The time lags obtained in the relationships stress the importance of river runoff and wind mixing for the early stages of anchovy and sardine, respectively, and their impact on recruitment.     

Climate impacts on the landings of Indian oil sardine over the south‐eastern Arabian Sea

The landings of Indian oil sardine (Sardinella longiceps, Clupeidae) along the south‐eastern Arabian Sea are about 43.8% of total Indian oil sardine production. The annual landings of this species exhibit large‐scale variability with prolonged years of surplus or deficit landings without identified reason. Evaluating Indian oil sardine landings along the Kerala coast during 1961–2017 in relation to environmental variations, we have elucidated a putative link between variability in landings versus environmental parameters and climate indices. The variables examined in this study, such as salinity and temperature along with physical indices such as upwelling and mixed layer depth (MLD) of the ocean help to propose a mechanism to temporal variability in the landings of Indian oil sardine. Colder temperature and timely intense upwelling lead to nutrient enrichment in the surface water, which promotes the growth of phytoplankton (chl‐a) and thereby food availability to Indian oil sardine are found during years with surplus catch. Less saline surface waters and shoaling of MLD at these times could lead to the aggregation of fish at particular depths and thereby a good catches. The reverse mechanism, such as more surface saline water, warm temperature, downwelling or weak upwelling, and less nutrient enrichment, leads to deficit landings. Further, it was noticed that the Pacific decadal oscillation and Atlantic multidecadal oscillation have a more pronounced impact on Indian oil sardine landings over the coast of south‐eastern Arabian Sea than previously reported ENSO associated impacts. All these point towards climate change implications for the Indian oil sardine fishery.     

Seasonal size distribution,condition status and muscle yield of Mediterranean horse mackerel <Emphasis Type="Italic">Trachurus mediterraneus</Emphasis> from the North Aegean Sea,Greece

ABSTRACT:   The seasonal intrinsic variability of the North Aegean Sea Mediterranean horse mackerel Trachurus mediterraneus population has been studied from monthly samples taken from the Greek purse-seine fishery over a 4-year period (2001–2005). The length and weight frequency distributions differed significantly by season, the highest mean lengths and weights attained in spring and the lowest in autumn. Length-weight relationships were determined on a monthly and annual basis, and monthly variations on the allometry coefficient were found. The differences between the coefficient obtained for annual data and the coefficients for each month were not significant in 50% of the cases. The b -values obtained both in this study and some of the previously reported in the Mediterranean Sea and other Greek waters were compared, showing basic similarities and minor spatial variation. The seasonal cycle in Mediterranean horse mackerel condition appears to be related to the reproductive cycle of the fish. Values of muscle yield ranged between 23.33% and 33.07% (mean value 28.48 ± 2.49%), but variations within seasons were not significant.     

Habitat suitability modelling for sardine juveniles (Sardina pilchardus) in the Mediterranean Sea

Identification of potential juvenile grounds of short‐lived species such as European sardine (Sardina pilchardus) in relation to the environment is a crucial issue for effective management. In the current work, habitat suitability modelling was applied to acoustic data derived from both the western and eastern part of the Mediterranean Sea. Early summer acoustic data of sardine juveniles were modelled using generalized additive models along with satellite environmental and bathymetry data. Selected models were used to construct maps that exhibit the probability of presence in the study areas, as well as throughout the entire Mediterranean basin, as a measure of habitat adequacy. Areas with high probability of supporting sardine juvenile presence persistently within the study period were identified throughout the Mediterranean Sea. Furthermore, within the study period, a positive relationship was found between suitable habitat extent and the changes in abundance of sardine juveniles in each study area.     

The selection of spawning location of sardine (Sardinops sagax) in the northern Benguela after changes in stock structure and environmental conditions

Most reports on the distribution of spawning areas of sardine (Sardinops sagax) in the northern Benguela originate from the 1970s and 1980s. The northern Benguela system was in a high upwelling regime during those decades. Since the early 1990s upwelling favourable winds have decreased and a trend of increasing sea surface temperature (SST) has been observed. Changes in the structure of sardine stock in the northern Benguela have been observed and it has been suggested that a reduced biomass and changes in stock structure has led to decreased spawning in the favourable southern locations, thus preventing a recovery of the sardine stock. The present paper on the contrary shows that there has been a shift in spawning location from the less favourable northern areas in the early 1980s to spawning areas further south in the 2000s. Thus, the failure of the northern Benguela sardine stock to recover since its collapse in the late 1960s cannot be explained by spawning in less favourable areas. The shift in preferred spawning location to more southern areas since the 1980s was to be expected with a general warming of the northern Benguela system. Alternative explanations for the failure of the sardine stock to recover such as a reduction in average length as well as length at 50% maturity, leading to a reduction in reproductive output, increased predation pressure, and increased low oxygen waters are proposed.     

Variability of oceanographic and meteorological conditions in the northern Alboran Sea at seasonal,inter‐annual and long‐term time scales and their influence on sardine (Sardina pilchardus Walbaum 1792) landings

Time series of European sardine (Sardina pilchardus) landings from 1962 and environmental variables from 1978 in the northern Alboran Sea are analysed. European sardine spawns in the northern Alboran Sea from mid‐autumn to late winter at a temperature range slightly higher than the one observed in the nearby Eastern North Atlantic and the North Western Mediterranean. Individuals hatched during autumn and winter are incorporated to the fishery during the following summer and autumn producing the maximum annual landings. These landings show both a decreasing long‐term trend and a strong inter‐annual variability. Although further research is needed, the warming trend of sea surface temperature and the decrease in upwelling intensity inferred from empirical orthogonal function (EOF) analyses could have some influence on the negative trends of sardine landings. The inter‐annual variability of sardine abundance seems to be related to the wind intensity at a local scale, the second principal component of the chlorophyll concentration and the sardine abundance during the preceding year. If the inter‐annual variability is considered, a linear model including these three variables with a one‐year time lag allows to explain 79% of the sardine landings variance. If the negative linear trend is also considered, the model explains 86% of the variance. These results indicate that the body condition of spawners, linked to the food availability during the preceding year, is the main factor controlling the recruitment success. The possibility of predicting sardine landings 1 year in advance could have important implications for fishery management.     

Environmental effects on the spatio‐temporal patterns of abundance and distribution of Sardina pilchardus and sardinella off the Mauritanian coast (North‐West Africa)

From 1998 to 2011, the effects of environmental conditions on the spatial and temporal trends of sardine and sardinella catch rates in the Mauritanian waters were investigated using generalized additive models. Two models were used: a global model and an oceanographic model. The global models explained more of the variability in catch rates (60.4% for sardine and 40% for sardinella) than the oceanographic models (42% for sardine and 32.4% for sardinella). Both species showed clear and inverse seasonal variations in abundances corresponding to their main spawning activities and the hydrologic seasons off the Mauritanian waters. Sardine prefer colder waters and seem to occupy the ‘gap’ in the northern part of the Mauritanian waters as soon as sardinella has left the area because of to lower water temperatures. Unlike sardinella, sardine showed a gradual southward extension between 2002 and 2009. The oceanographic model revealed that a high proportion of catch variability for the two species could be explained by environmental variables. The main environmental parameters explaining the variability are sea surface temperature (SST) and the upwelling index for sardinella, and the chlorophyll‐a (Chl‐a) concentration, the upwelling index and SST for sardine. The sardinella spatio‐temporal variations off Mauritania seem to be more controlled by thermal than productivity gradients, probably linked to the species physiological constraints (thermal tolerance) whereas sardine seems to be more controlled by an ‘optimal upwelling and temperature’ windows. The results presented herein may be useful for understanding the movement of these species along the Mauritanian coast and hence their management under a changing climate.     

Effects of seasonal variations in meteorology and oceanography on the Brazilian sardine fishery

Seasonal variations in meteorological and oceanographic processes in the South Brazil Bight were studied to assess their influence on the Brazilian sardine Sardinella brasiliensis fishery. According to its environmental characteristics, the South Brazil Bight was divided into three sectors. The northern and southern sectors were similar during the spring and summer, showing areas of coastal upwelling and low rainfall rates. The middle sector was characterized by the presence of low-salinity coastal waters in the inner shelf and by high rainfall. During autumn and winter, the middle and southern sectors were similar, both being under the influence of sub-Antarctic waters. Multiple regression analyses showed significant correlation between environmental data and catches, although at a lower level in the northern sector ( r ² = 0.57). In the middle and southern sectors, environmental data were highly correlated with the catches ( r ² = 0.90 and 0.84). The sardine landings in the middle sector are principally affected by meteorological parameters, and in the south by oceanographic ones. In the middle sector, an inverse correlation between landings and rainfall is shown, while in the south, landings are positively correlated with salinity and sea temperature. The conclusion is that the seasonal landings of the Brazilian sardine within the South Brazil Bight are influenced by the distribution of less saline waters, the coastal waters and sub-Antarctic waters.     

Importance of the Shatsky Rise Area in the Kuroshio Extension as an offshore nursery ground for Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus)

Recent findings suggest that recruitment of Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus) depends on survival during not only the first feeding larval stage in the Japanese coastal waters and the Kuroshio front but also during the post‐larval and juvenile stages in the Kuroshio Extension. Spatial distributions of juvenile anchovy and sardine around the Shatsky Rise area in the Kuroshio Extension region and the Kuroshio–Oyashio transition region are described, based on a field survey in the late spring using a newly developed mid‐water trawl for sampling juveniles. All stages of anchovy from post‐larvae to juveniles were obtained in the northern Shatsky Rise area. The Kuroshio Extension bifurcates west of the Shatsky Rise area and eddies are generated, leading to higher chlorophyll concentrations than in the surrounding regions in April and May. When Japanese anchovy and sardine spawn near the Kuroshio front or the coastal waters south‐east of Japan, their larvae are transported by the Kuroshio Extension and are retained in the Shatsky Rise area, which forms an important offshore nursery ground, especially during periods of high stock abundance.     

Evidence of density-dependent effects on population variation of Japanese sardine (Sardinops melanosticta) off Korea

Interannual variation of some biological parameters for the Japanese sardine, Sardinops melanosticta, population in Korean waters was investigated using scientific surveys and fisheries information since the late 1970s. The abundance and geographical coverage of sardine eggs were high (peak in 1986) and broad when spawning biomasses were high in the mid 1980s, and vice versa in the late 1970s and the early 1990s. Also, feeding and spawning areas based on fisheries information exhibited the same pattern of expansion/contraction as seen in ichthyoplankton surveys. Annual Gonadal Somatic Index (GSI) in spawning season (February to April) and the size at age 1 of sardine were reduced during the high abundance period. It is suggested that density‐dependent effects on the reproduction and growth of the sardine population in Korean waters existed.     

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

A 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.     

Linking small pelagic fish and cetacean distribution to model suitable habitat for coastal dolphin species,Delphinus delphis and Tursiops truncatus,in the Greek Seas (Eastern Mediterranean)

1. A large‐scale assessment of the summertime suitable habitat for Delphinus delphis (short‐beaked common dolphin) and Tursiops truncatus (common bottlenose dolphin) in Greek Seas (Eastern Mediterranean) was conducted using data from dedicated and opportunistic cetacean surveys and published data records.
2. Using a presence/absence approach, generalized additive models were applied to define a suite of environmental, bathymetric and biotic factors that best describe common and bottlenose dolphin spatial distribution, during early (May, June, July) and late (August, September) summer.
3. A geographic information system (GIS) was used to integrate sightings data with environmental characteristics, distance from the coast and sardine probability of presence. These variables were considered as good proxies for defining species‐suitable habitat within the study area's coastal environment.
4. The final selected models were used to produce annual probability maps of the presence of the species in the entire Greek Seas, as a measure of habitat suitability. Based on the mean probability and standard deviation maps for the study period GIS techniques were subsequently used to determine the persistent (areas with high mean and low variation) and occasional (high mean and high variation) habitat of each species.
5. Results showed that there was a high probability of common dolphin presence in areas with a high probability of sardine presence. For bottlenose dolphin, higher probability of the presence of species occurred in areas closer to the shore, with a high probability of sardine presence and with high concentrations of chlorophyll‐a.
6. In both seasons, the North Aegean Sea and the Inner Ionian Sea Archipelago were indicated as the most suitable areas for common dolphin distribution. Persistent habitat areas of the bottlenose dolphin included enclosed seas, continental shelf waters, and waters surrounding islands. The indicated suitable areas are discussed along with deficiencies of the models and future implications for conservation.

Copyright © 2016 John Wiley & Sons, Ltd.     

Success of the Introduction of the Fresh Water Clupeid Limnothrissa miodon (Boulenger) in Lake Kivu

The freshwater clupeid Limnothrissa miodon was transplanted with success from Lake Tanganyika to Lake Kivu during 1958–1960. An artisanal fishery to exploit the clupeid stock started in the beginning of 1980. Based on the results of this artisanal fishery a minimum sustained yield of 4,000 tons/year was estimated for the Rwandese territorial waters of Lake Kivu. Biological research revealed information about growth rate, condition factor K, fat content, migration, distribution and reproduction of Limnothrissa. An obvious relation was found between fat content and sexual activity with the seasonal fluctuations of plankton production. The main spawning peak coincided with a plankton bloom caused by the turnover of the lake at the end of the dry season in September. At that time a small percentage of the clupeids had fat in their body cavity. It is stated that horizontal migration is mainly induced by reproduction, while the vertical migration is determined by feeding habits. Socio-economic benefits of the artisanal fishery using trimarans and lift nets were compared with those of the semi-industrial purse seining.     

A continuous, underway fish egg sampler

We describe a method to sample the highly contagious distribution of pelagic fish eggs. CUFES, the continuous, underway fish egg sampler, consists of a submersible pump, concentrator, electronics and sample collector. This system operates continuously and under nearly all sea conditions, providing a real-time estimate of the volumetric abundance of pelagic fish eggs at pump depth, usually 3 m. CUFES-derived estimates of volumetric abundance agree well with those from nets towed at pump depth and with areal abundance estimated from vertically integrated plankton tows. CUFES has been used successfully to sample the eggs of menhaden, pinfish, sardine, and anchovy off the coasts of the eastern and western United States and South Africa. Two large patches of eggs of the Atlantic menhaden were sampled off North Carolina in winter 1993–94, had a linear scale of 5–10 km, and were found in waters between the Gulf Stream and mid-shelf front. Spawning location may be related to bathymetry. CUFES is now being used to estimate spawner biomass by the daily egg production method. An optical plankton counter provided accurate estimates of the number of Atlantic menhaden eggs sample by CUFES.     

Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.     

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

The 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.     

Growth and age composition of northern shrimp Pandalus eous estimated by multiple length frequency analysis

We examined growth of northern shrimp Pandalus eous in the Sea of Japan, off western Hokkaido, to improve estimations of catch-at-age for stock assessment. Multiple length frequency analysis based on length frequency data collected by a scientific research vessel was conducted to examine length-at-age in the shrimp population. Multi-normal distributions estimated using maximum likelihood indicated a good fit to length distributions. AIC values and regression analyses revealed annual growth variation and a decreasing trend in the length at several age classes in the shrimp population. We revised the method for estimating catch-at-age from the age-conversion table (ACT), which is a simple method for age determination, to age–length keys (ALK) calculated from the results of multiple length frequency analysis. Abundant year classes caught successively year after year could be more easily identified from the catch-at-age data computed using ALK than by using ACT. Our results suggested not only that the mean size of commercial landings fluctuated based on changes in age composition but also that a decrease in the length-at-age in the population influenced the consistent size decrease of commercial landings.     

Larval growth rates differ in response to seasonal temperature variations among clupeoid species inhabiting the Pacific coastal waters of Japan

Growth rates of round herring Etrumeus teres, Japanese anchovy Engraulis japonicus, and Japanese sardine Sardinops melanostictus larvae respond differently to seasonal temperature variations in Sagami Bay on the Pacific coast of central Japan. Larvae of these three clupeoid species were collected from small seine net catches between February 2008 and April 2009. Correlations between otolith growth increment widths and ambient temperatures were examined at five body sizes and indexed to otolith radii of 50–175 μm, which corresponded to notochord lengths of 15–24 mm. Round herring larvae showed positive correlations between growth rate and habitat temperature at all five body sizes. In contrast, Japanese sardine larvae did not show significant correlations between growth rate and ambient temperature at any of the five body sizes. Japanese anchovy larvae showed significant but weaker correlations between growth rate and temperature than round herring. Although these three species share habitats with some differences in seasonal occurrence in the bay, they showed different species-specific responses to seasonal habitat temperature variations. These differences in response to temperature variations may constitute an ecological basis for different migration patterns among the three species.