Fine scale spatial pattern of Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs

Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs exhibited different spatial structure on the scale of 0.75–2.5 km in two egg patches sampled in the Southern California Bight in April 2000. Plankton samples were collected at 4‐min intervals with a Continuous Underway Fish Egg Sampler (CUFES) on 5 × 5 km grids centered on surface drifters. Variograms were calculated for sardine and anchovy eggs in Lagrangian coordinates, using abundances of individual developmental stages grouped into daily cohorts. Model variograms for sardine eggs have a low nugget effect, about 10% of the total variance, indicating high autocorrelation between adjacent samples. In contrast, model variograms for anchovy eggs have a high nugget effect of 50–100%, indicating that most of the variance at the scales sampled is spatially unstructured. The difference between observed spatial patterns of sardine and anchovy eggs on this scale may reflect the behavior of the spawning adults: larger, faster, more abundant fish may organize into larger schools with greater structure and mobility that create smoother egg distributions. Size and mobility vary with population size in clupeoids. The current high abundance of sardines and low abundance of anchovy off California agree with the greater autocorrelation of sardine egg samples and the observed tendency for locations of anchovy spawning to be more persistent on the temporal scale of days to weeks. Thus the spatial pattern of eggs and the persistence of spawning areas are suggested to depend on species, population size and age structure, spawning intensity and characteristic physical scales of the spawning habitat.     

Anchovy (Engraulis ringens) and sardine (Sardinops sagax) spatial dynamics and aggregation patterns in the Humboldt Current ecosystem, Peru, from 1983–2003

Three indexes of spatial aggregation are developed and used to examine the aggregation pattern of sardine (Sardinops sagax) and anchovy (Engraulis ringens) in the Peruvian Humboldt Current System, determined from 36 acoustic surveys conducted from 1983 through 2003 by the Peruvian Marine Institute (IMARPE). Each index assesses a different aspect of aggregation: the concentration, the percent occupancy of space and the clustering of high‐fish abundance. Both time‐series correlation and tree‐based clustering‐regression method, classification and regression trees (CART), were used to relate each of the indexes to environmental variables (season, temperature anomaly and year). Additionally, a measure of onshore–offshore distribution, the average distance from the coast, and abundance variables (the average acoustic backscatter per occupied sampling unit, and the acoustically estimated total abundance of sardine and anchovy from IMARPE) were related to environmental factors by using CART. We show that the 1983–2003 time series can be divided into three different periods: with shifts in 1992 and in 1997–98. Sardine and anchovy showed large differences in both abundance and aggregation among these periods. Although upwelling ecosystems support dramatic and sudden changes in environmental conditions, fish responses are sometimes smoother than usually suggested and there are transition periods with concomitant high biomasses of anchovy and sardine, but with different spatial aggregation patterns. Observed relationships between environmental proxies and aggregation patterns support the habitat‐based hypothesis that environmentally mediated alterations in range lead to population changes.     

Historical fluctuations in spawning location of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during 1967–73 and 2000–2004

The spatial extent of small pelagic fish spawning habitat is influenced by environmental factors and by the state of the adult population. In return, the configuration of spawning habitat affects recruitment and therefore the future structure of the adult population. Interannual changes in spatial patterns of spawning reflect variations in adult population structures and their environment. The present study describes the historical changes in the spatial distribution of spawning of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during two periods: 1967–72 and 2000–2004. Using data from egg surveys conducted in spring, the spatial distributions of anchovy and sardine eggs are characterized by means of geostatistics. For each survey, a map of probability of egg presence is constructed. The maps are then compared to define (1) recurrent spawning areas, (2) occasional spawning areas and (3) unfavourable spawning areas during each period. Sardine spawning habitat is generally fragmented and appears spatially limited by the presence of cold bottom water. It is confined to coastal or shelf break refuge areas in years of restricted spawning extent. For anchovy, recurrent spawning sites are found in Gironde and Adour estuaries whilst spawning can extend further offshore in years of more intense spawning. For both species, the mean pattern of spawning has changed between 1967–72 and 2000–2004. Noticeably, the spatial distribution of anchovy eggs in spring has expanded northward. This trend possibly results from changes in environmental conditions during the last four decades.     

Environmental processes driving anchovy and sardine distribution in a highly variable environment: the role of the coastal structure and riverine input

Engraulis encrasicolus and Sardina pilchardus stocks are highly variable in terms of recruitment, biomass and spatial distribution. Changes in habitat conditions may influence both the survival of the early life stages and the adult stages. Detailed studies on the spatial distribution and habitat selection of such species have been performed in different areas of the world, highlighting the importance of environmental processes. The present study analyzes the spatial distribution of anchovy and sardine in the Tyrrhenian Sea in relation to environmental heterogeneity. Four acoustic surveys were carried out in this area in the period 2009–2014. Analysis of the environmental dataset permitted identification, in two specific areas, of a pattern of variables driving enrichment processes and impacting on the habitat suitability of the two species. In the northern and central parts of the study area, both anchovy and sardine showed a marked preference for shallower areas characterized by lower salinity. In these areas, PCA results on an environmental dataset highlighted a strong link between primary production, particulate organic carbon, distance from the mouth of the river, salinity and depth. A less clear picture was obtained for the southern part of the Tyrrhenian sea, characterized by a narrow continental shelf, moderately complex coastline morphology and the presence of very small rivers. Most of the anchovy biomass was found to be located in enclosed areas (gulfs) under the influence of relatively small rivers. This finding, taking into account that the surveys were carried out during the anchovy spawning period, highlights for such species a positive effect of the interaction between coastal morphology and riverine input, probably favoring food supply and retention of spawning products.     

From small-scale habitat loopholes to decadal cycles: a habitat-based hypothesis explaining fluctuation in pelagic fish populations off Peru

The Peru‐Humboldt Current system (HCS) supports the world's largest pelagic fisheries. Among the world's eastern boundary current systems, it is the most exposed to high climatic stress and is directly affected by El Niño and La Niña events. In this volatile ecosystem, fish have been led to develop adaptive strategies in space and time. In this paper, we attempt to understand the mechanisms underlying such strategies, focusing on the El Niño 1997–98 in Peru from which an extensive set of hydrographic, capture and acoustic survey data are available. An integrated analysis of the data is crucial, as each has substantial shortcomings individually; for example, both catch data and acoustic surveys may easily lead to wrong conclusions. Existing hypotheses on anchovy and sardine alternations lead us to a ‘habitat‐based’ synthetic hypothesis. Using our data, an integrated approach evaluated how fish responded to habitat variation, and determined the consequences in terms of fish‐population variability. Various factors occurring at a range of different spatio‐temporal scales were considered: interdecadal regime (warm ‘El Viejo’/cool ‘La Vieja’ decadal scale); strength and the duration of the El Niño Southern Oscillation event (interannual scale); population condition before the event (interannual scale); fishing pressure and other predation (annual scale); changes in reproductive behaviour (intra‐annual scale); presence of local upwelling (local scale). During El Niño 1997–98, anchovy was able to exploit a small‐scale temporal and spatial ‘loophole’ inside the general unfavourable conditions. Moreover, sardine did not do better than anchovy during this El Niño and was not able to take advantage of the ‘loophole’ opened by this short‐term event. Our results question the traditional view that El Niño is bad for anchovy and good for sardine.     

Variability in the spawning habitat of Pacific sardine (Sardinops sagax) off southern and central California

The spatial pattern of sardine spawning as revealed by the presence of sardine eggs is examined in relation to sea surface temperature (SST) and mean volume backscatter strength (MVBS) measured by a 150 kHz acoustic Doppler current profiler (ADCP) during four spring surveys off central and southern California in 1996–99. Studies in other regions have shown that MVBS provides an excellent measure of zooplankton distribution and density. Zooplankton biomass as measured by survey net tows correlates well with concurrently measured MVBS. The high along‐track resolution of egg counts provided by the Continuous Underway Fish Egg Sampler (CUFES) is a good match to the ADCP‐based data. Large interannual differences in the pattern and density of sardine eggs are clearly related to the concurrently observed patterns of surface temperature and MVBS. The strong spatial relationship between sardine eggs and MVBS is particularly evident because of the large contrast in zooplankton biomass between the 1998 El Niño and 1999 La Niña. The inshore distribution of sardine spawning appears to be limited by the low temperatures of freshly upwelled waters, although the value of the limiting temperature varies between years. Often there is an abrupt offshore decrease in MVBS that is coincident with the offshore boundary of sardine eggs. Possible reasons for this association of sardine eggs and high zooplankton biomass include an evolved strategy that promotes improved opportunity of an adequate food supply for subsequent larval development, and/or adult nutrient requirements for serial spawning. Hence, the distribution of these parameters can be used as an aid for delineating the boundaries of sardine spawning habitat.     

1999年2月黄海声学调查鳀鱼种群群体结构评估

IntroductionEstimation of the size structure of a fish population is often one of the primary objectives of an acoustic/trawl survey (Gunderson 1993;Honkalehto and Williamson 1996 ) .The size structure contains much informa-tion enabling a quick estimate of the fishable size of a stock in a regulated fishery or the recruitment status ofa stock.It is also a common practice to use the number at length of a population together with the age/lengthkey to produce the age structure of the fish popul…     

Alternating dominance of postlarval sardine and anchovy caught by coastal fishery in relation to the Kuroshio meander in the Enshu-nada Sea

In the mid 1970s, the fishery catch of postlarval Japanese anchovy (Engraulis japonica) in a shelf region of the Enshu‐nada Sea, off the central Pacific coast of Japan, started to decline corresponding to a rapid increase of postlarval sardine (Sardinops melanostictus). In late 1980s, sardine started to decline, and it was replaced by anchovy in the 1990s. This alternating dominance of postlarval sardine and anchovy corresponded to the alternation in egg abundance of these two species in the spawning habitat of this sea. It was also noteworthy that during the period of sardine decline, sardine spawning occurred in April–May, a delay of two months compared with spawning in the late 1970s. The implication of oceanographic changes in the spawning habitat for the alternating dominance of sardine and anchovy eggs was explored using time‐series data obtained in 1975–1998, focusing on the effect of the Kuroshio meander. Large meanders of the Kuroshio may have enhanced the onshore intrusion of the warm water into the shelf region and contributed to an increase in temperature in the spawning habitat. This might favour sardine, because its egg abundance in the shelf region was more dependent on the temperature in early spring than was that of anchovy. In addition, enhanced onshore intrusion could contribute to transport of sardine larvae from upstream spawning grounds of the Kuroshio region. On the other hand, anchovy egg abundance was more closely related to lower transparency at the shelf edge, which may indicate the prevalence and prolonged residence of the coastal water, and therefore higher food availability, frequently accompanying non‐meandering Kuroshio. The expansion/shrinkage of the spawning habitat of sardine and anchovy in the shelf region, apparently responding to the change in the Kuroshio, possibly makes the alternation in dominance of postlarval sardine and anchovy most prominent in the Enshu‐nada Sea, in combination with changes in the abundance of spawning adults, which occurred almost simultaneously in the overall Kuroshio region. The implication of this rather regional feature for the alternating dominance of sardine and anchovy populations on a larger spatial scale is also discussed.     

Modelling potential spawning habitat of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) in the Bay of Biscay

Large amplitude variations in recruitment of small pelagic fish result from interactions between a fluctuating environment and population dynamics processes such as spawning. The spatial extent and location of spawning, which is critical to the fate of eggs and larvae, can vary strongly from year to year, as a result of changing population structure and environmental conditions. Spawning habitat can be divided into ‘potential spawning habitat’, defined as habitat where the hydrographic conditions are suitable for spawning, ‘realized spawning habitat’, defined as habitat where spawning actually occurs, and ‘successful spawning habitat’, defined as habitat from where successful recruitment has resulted. Using biological data collected during the period 2000–2004, as well as hydrographic data, we investigate the role of environmental parameters in controlling the potential spawning habitat of anchovy and sardine in the Bay of Biscay. Anchovy potential spawning habitat appears to be primarily related to bottom temperature followed by surface temperature and mixed‐layer depth, whilst surface and bottom salinity appear to play a lesser role. The possible influence of hydrographic factors on the spawning habitat of sardine seems less clear than for anchovy. Modelled relationships between anchovy and sardine spawning are used to predict potential spawning habitat from hydrodynamical simulations. The results show that the seasonal patterns in spawning are well reproduced by the model, indicating that hydrographic changes may explain a large fraction of spawning spatial dynamics. Such models may prove useful in the context of forecasting potential impacts of future environmental changes on sardine and anchovy reproductive strategy in the north‐east Atlantic.     

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.     

Characterizing the potential habitat of European anchovy Engraulis encrasicolus in the Mediterranean Sea,at different life stages

Identification of the potential habitat of European anchovy (Engraulis encrasicolus) at different life stages in relation to environmental conditions is an interesting subject from both ecological and management points of view. For this purpose, acoustic data from different seasons and different parts of the Mediterranean Sea along with satellite environmental and bathymetry data were modelled using generalized additive models. Similarly, egg distribution data from summer ichthyoplankton surveys were used to model potential spawning habitat. Selected models were used to produce maps presenting the probability of anchovy presence (adults, juveniles and eggs) in the entire Mediterranean basin, as a measure of habitat adequacy. Bottom depth and sea surface chlorophyll concentration were the variables found important in all models. Potential anchovy habitats were located over the continental shelf for all life stages examined. An expansion of the potential habitat from the peak spawning (early summer) to the late spawning season (early autumn) was observed. However, the most suitable areas for the presence of anchovy spawners seem to maintain the same size between seasons. Potential juvenile habitats were associated with highly productive inshore waters, being less extended and closer to coast during winter than late autumn. Potential spawning habitat in June and July based on ichthyoplankton surveys overlapped but were wider in extent compared with adult potential habitat from acoustics in the same season. Similarities and dissimilarities between the anchovy habitats as well as comparisons with sardine habitats in the oligotrophic Mediterranean Sea and other ecosystems with higher productivity are discussed.     

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

ABSTRACT:   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 back-calculation 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.     

Size,permeability and buoyancy of anchovy (Engraulis Encrasicolus) and sardine (Sardina Pilchardus) eggs in relation to their physical environment in the Bay of Biscay

The size and specific gravity of eggs of marine pelagic fish partly determine their dispersal and survival. Using an original dataset of anchovy and sardine eggs, sampled in spring over the last decade in the Bay of Biscay, we provide a parameterization of these properties on ambient water temperature and salinity. We used the density gradient column for measurement of egg specific gravity. The column was also filled with homogeneous water for sinking velocity experiments. For anchovy, these experiments confirm that the effect of egg permeability through the chorion could be neglected when modelling sinking, while it has to be considered for sardine, its perivitelline space representing 78.6% (±6.2%) of the total egg volume, as opposed to 5–10% for most teleosts species. We estimated a coefficient of permeability of the chorion of 0.0038 mm s^?1. However, permeability should not affect the measurement of sardine egg specific gravity in a gradient column, provided a minimum duration before reading is respected for equilibrium to be reached. In relation to their environment, we found that the egg specific gravity is largely determined by sea surface salinity for both species, whereas egg size is weakly but significantly impacted by temperature, for sardine only. On average, the estimated difference in specific gravity between egg and surface water is ?0.92 σ_T for anchovy and ?1.06 σ_T for sardine. The detailed parameterization of the relationship between eggs and water properties should prove useful, in particular to modellers dealing with the dispersal of fish early life stages.     

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.     

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.     

The spatial structure of the oceanic spawning of jack mackerel (Trachurus murphyi) off central Chile (1998–2001)

The jack mackerel population has a widespread oceanic spawning habitat off central Chile, extending more than one thousand nautical miles offshore. In this paper, the spatial structure of jack mackerel eggs density is analyzed on the basis of four surveys carried out in oceanic waters (32°S–39°S, 75°W–92°W), from 1998 to 2001. In each survey, a grid of plankton stations was sampled through vertical hauls with WP2 plankton nets by using several purse-seine fishing ships sampling simultaneously along the E–W transects. With the aim of finding the bulk of the egg distribution within the surveyed area, an exploratory analysis between jack mackerel egg densities, latitude, longitude, and sea surface temperature (SST) was carried out. The spatial structure of the egg distribution was studied using geostatistical techniques. The bulk of the jack mackerel spawning tends to occur offshore between 80°W and 92°W, is maximal at 35°S and associated to SST warmer than 15–16 °C. All of the variograms showed clear spatial autocorrelations without anisotropy, with the range fluctuating between 125 and 252 nautical miles. The range of variograms suggests that the spawning of jack mackerel is a large scale process, probably reflected in the adult behavior of the spawning by favoring a high dispersion of eggs and/or associated with sea surface temperature characterizing the subtropical frontal zone (16–18 °C) off central Chile.     

Density dependence in total egg production per spawner for marine fish

A paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.     

Recruitment of common sardine (Strangomera bentincki) and anchovy (Engraulis ringens) off central-south Chile in the 1990s and the impact of the 1997–1998 El Niño

The recruitment of Strangomera bentincki (common sardine) and Engraulis ringens (anchovy) and the relationships with oceanographic conditions in the upwelling ecosystem of central-south Chile were investigated from 1990 to 1998, with emphasis on the 1997–1998 El Niño. Time series of recruitment, biomass, local sea surface temperature, and a coastal upwelling index were used to explore relationships during the spawning (July–August) and pre-recruitment (August–December) periods. The 1997–1998 El Niño caused physical changes in the small pelagic fish habitat off central-south Chile. Anomalous sea surface temperatures (SST) and upwelling indexes began to be detected from May 1997 and persisted into 1998. Recruitment of common sardine showed significantly negative relationship with SST anomalies during the pre-recruitment period, as well as with the upwelling index during the peak of spawning. However, the recruitment of anchovy did not seem to be affected by the environmental changes observed in the 1990s. Instead, the recruitment rate of anchovy showed negative relationship with the recruitment rate of common sardine. We conclude that the conditions of the 1997–1998 El Niño off central-south Chile affected the survival of common sardine offspring, and that the recruitment success of anchovy could be determined by less-abundant cohorts of common sardine through a biological mechanism of interaction.