The spatial distribution of cod (Gadus morhua L.) spawning grounds in the Kattegat, eastern North Sea

Similar to many other commercial marine fish species, Atlantic cod (Gadus morhua L.) migrate towards specific sites at spawning. These temporal aggregations are generally the most targeted by the commercial fishery. The Kattegat cod has undergone a substantial reduction over the past 25 years and both stock size and spawning stock biomass have remained at very low levels since the end of the 1990s. There is, therefore, an urgent need to map and document spawning grounds still in use. In the present study, spawning sites of Atlantic cod were identified in the Kattegat using a combination of commercial and fishery independent data from 1996 to 2004. Moreover, putative spawning and non-spawning areas were also sampled before and during the reproductive season between 2002 and 2006, and the proportion of mature females together with the individual physiological status were used to validate and strengthen the spatial analyses.

The spatial analyses identified several spawning areas in the region and two areas in the southeastern part of the Kattegat which appeared to be the most important. The results showed the presence of cod spawning aggregations, although reduced in size, in areas that have been utilized for more than 25 years according to historical information. Some local spawning grounds may have also disappeared. The proportion of mature females was higher in putative spawning than in non-spawning areas (p < 0.001) and females from spawning areas had higher gonadosomatic (p < 0.05) and hepatosomatic (p < 0.001) indices than those from non-spawning areas.

Knowledge of stock spatial and temporal distribution is essential in designing recovery strategies for depleted fish populations. The unambiguous stability of the locations of spawning aggregations over time, as shown in this study, represents a useful aid in order to efficiently implement a recovery plan for the collapsing cod stock in this area.     

The influence of temperature on advective loss of Atlantic cod (Gadus morhua) eggs from the inshore environment

We use a simple model of temperature-dependent egg development and mortality to develop several hypotheses concerning the effect of temperature on the occurrence of eggs of Atlantic cod, Gadus morhua , in Placentia Bay, Newfoundland. Predictions from this exploratory model were tested through a series of synoptic ichthyoplankton surveys throughout spawning and postspawning periods during 1997 and 1998. Although several egg mortality relationships were explored through the simulation, a constant mortality rate best represented the pattern observed in the two years of data. Peaks in late stage egg densities occurred in August of both 1997 and 1998 and were apparently decoupled from egg production peaks in April. We observed a decrease in mortality and the distance dispersed during egg development with increases in water temperature. We suggest that the effects of predation are small relative to the advective effects within this system, and that the interaction between advection and temperature-dependent vital rates of eggs may have dramatic consequences for coastal retention of propagules produced by inshore spawning events.     

Modelling advection of cod eggs and larvae on the Newfoundland Shelf

We investigate interannual variations in dispersion for the drift and retention of cod eggs and larvae on the Newfoundland Shelf using a two-dimensional Lagrangian tracking model. The velocity field for the drift model is obtained from a diagnostic calculation of objectively analysed density data. Time-dependent currents are generated using an inertial-current slab model driven by observed winds. Eggs and larvae are treated as passive drifters seeded in a dispersion model of the Newfoundland Shelf region. We identify favourable and unfavourable zones of retention on the Newfoundland Shelf. We show that northerly, shelf-break spawning locations are more favourable than southerly shelf-break spawning locations for northern cod, ( Gudus morhua , in NAFO divisions 2J3KL).     

Residency of adult Atlantic cod (Gadus morhua) in the western Gulf of Maine

A mark and recapture study of cod in the western Gulf of Maine was conducted to study the seasonal movements of fish, particularly as they related to areas closed to commercial fishing. A total of 27,772 cod were tagged, and 1334 (4.8%) were recaptured with sufficiently detailed recapture location and date to be included in the study. Results indicated that the group is resident to the area and sedentary. Although there were a small percentage of fish (2.5%) that traveled long distances (>100 km), most were recaptured near their release location. There was no linear relationship between fish length and linear distances traveled, and linear distance traveled was only weakly related to days-at-large. For all groups of cod tagged and released in particular areas and months, mean distances traveled were small (<65 km), rates of travel were slow (<0.2 km/day), and rates of group dispersion were <50 km²/day. There were no recognizable spatial or temporal patterns in the mean angles of travel for groups released in various month/area combinations, and the angular deviations associated with the mean angles were quite large. Although there seems to be little pattern in the movement of cod in our study area, temporal and spatial changes in abundance indicate that movements are occurring.

The general pattern was a concentration of large cod into one small, inshore area (Area 133) in both the spring and winter, and dispersion from this area in the ensuing months. Monthly percentages of ripe cod in Area 133 provide evidence for two spawning groups; a winter group that spawns from November through January, and a spring group that spawns from April through July. Thus it is likely that the observed spring and winter concentrations of fish in Area 133 were associated with spawning. Fish in the spring group were the largest encountered in the study. The study also provided some evidence of natal homing, i.e. a return to the same spawning grounds year after year, for the spring spawning group. Although there is some exchange of fish between most of the closed areas, the timing of the closures appears to protect the largest aggregations of cod.     

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.     

Temperature influences on growth of unfished juvenile Northern cod (Gadus morhua) during stock collapse

This study examined growth of unfished juvenile Northern cod (Gadus morhua) off Newfoundland concomitant with stock collapse in the cold early 1990s. Two unpublished data sets were examined, one from collapse‐period trapping sites along the northeast coast of Newfoundland and one from a post‐collapse inshore trawl survey. Cumulative surface and bottom temperatures were significant predictors of growth rates of the young fish with year‐classes born during collapse experiencing slower growth than those born during subsequent warming. Relationships between accrued temperature and growth were consistent across periods, with slow growth of collapse‐period fish reflecting slower accumulation of temperature‐at‐age. Temperature influences were spatially broad‐based with no significant differences in growth rates for fish captured along the entire northeast coast of Newfoundland. Predicted differences in growth rates for collapse versus recovery year‐classes were proportional to cumulative surface temperatures but not cumulative bottom temperatures. Although significant, temperature effects on growth were relatively unimportant at youngest ages. Overall, growth differences between periods were small but large differences occurred between slowest and fastest growing year‐classes. The results suggest initial responses to increasing temperatures were delayed following collapse. We conclude that although temperature was an important determinant of dampened productivity that it alone cannot account for the collapse and slow recovery of the stock. This is the first known study to directly quantitatively link temperature impacts to an unfished component of the Northern cod stock complex during collapse, removing need for implicit assumptions about whether or not cold conditions contributed to the collapse of this iconic fish stock.     

Movements and spatiotemporal distribution of escaped farmed and local wild Atlantic cod (Gadus morhua L.)

Commercial farming of Atlantic cod (Gadus morhua L.) is now being developed in several countries. The ecological consequences of cod culture are poorly understood, but recent research suggests that Atlantic cod are more prone to escape from net pens than Atlantic salmon. Here, we describe the movements and the spatiotemporal distribution of farmed cod after escape relative to wild cod, both during and outside the natural spawning season. The experimental design included simulating escape incidents of farmed cod tagged with acoustic transmitters and using an array of automatic listening stations to monitor their dispersal and distribution. For comparison, local wild cod were monitored using the same array of receivers. The farmed cod dispersed rapidly after a simulated escape, they randomly distributed over large areas and their distribution overlapped with local wild cod. Moreover, escaped farmed fish were found at local cod spawning areas during the spawning season. The study also indicated that the recapture rate of escaped farmed cod was high compared with that of escaped farmed salmon. Thus, while our results showed that there is a considerable potential for ecosystem effects caused by escaped farmed cod, mitigating actions such as an efficient recapture fishery for escapees may be possible.     

A model of meta-population dynamics for North Sea and West of Scotland cod—The dynamic consequences of natal fidelity

It is clear from a variety of data that cod (Gadus morhua) in the North Sea do not constitute a homogeneous population that will rapidly redistribute in response to local variability in exploitation. Hence, local exploitation has the potential to deplete local populations, perhaps to the extent that depensation occurs and recovery is impossible without recolonisation from other areas, with consequent loss of genetic diversity. The oceanographic, biological and behavioural processes which maintain the spatial population structures are only partly understood, and one of the key unknown factors is the extent to which cod exhibit homing migrations to natal spawning areas. Here, we describe a model comprising 10 interlinked demes of cod in European waters, each representing groups of fish with a common natal origin. The spawning locations of fish in each deme are governed by a variety of rules concerning oceanographic dispersal, migration behaviour and straying. We describe numerical experiments with the model and comparisons with observations, which lead us to conclude that active homing is probably not necessary to explain some of the population structures of European cod. Separation of some sub-populations is possible through distance and oceanographic processes affecting the dispersal of eggs and larvae. However, other evidence suggests that homing may be a necessary behaviour to explain the structure of other sub-populations. The consequences for fisheries management of taking into account spatial population structuring are complicated. For example, recovery or recolonisation strategies require consideration not only of mortality rates in the target area for restoration, but also in the source areas for the recruits which may be far removed depending on the oceanography. The model has an inbuilt capability to address issues concerning the effects of climate change, including temperature change, on spatial patterns of recruitment, development and population structure in cod.     

Relationships between temperature and landing patterns in the southern Labrador and eastern Newfoundland inshore cod fishery

The temporal pattern of cod landings in the inshore trap fishery off the east coast of Newfoundland and Labrador is examined in this study. Unlike previous studies which have been broad scale or of a localized nature, this analysis looks at variations within fishing seasons and among fishing areas in trap landings. A significant relationship between water temperature and the weekly pattern is demonstrated. Significant spatial patterns in weekly landings are described and related to longshore differences in the seasonal variation in temperature.     

Oceanographic influences on the distribution of Baltic cod, Gadus morhua, during spawning in the Bornholm Basin of the Baltic Sea

The Baltic Sea is a stratified, semienclosed sea typified by a low-salinity surface layer and a deep saline layer of varying volume, salinity, temperature and oxygen concentration. The relationships between these oceanographic factors and the distribution of Baltic cod are presented, utilizing results from a survey carried out during the 1995 spawning period in the Bornholm Basin, at present the main spawning area of this stock. Cod distribution, abundance and population structure were estimated from hydroacoustic and trawl data and related to hydrographic parameters as well as to bottom depth. In the central basin, cod were aggregated in an intermediate layer about 15 m thick. This area of peak abundance was defined at its upper limit by the halocline and at the lower limit by oxygen content. The majority of individuals caught in the basin centre were in spawning or pre-spawning condition with a high proportion of males to females. On the basin slopes, aggregations of cod were found near the bottom. These individuals were mainly immature and maturing stages with an increasing proportion of females to males with size. Salinity and oxygen conditions were found to be the major factors influencing the vertical and horizontal distribution of adult cod. Abundance of immature cod was also positively related to decreasing bottom depths. The effect of temperature was minor. The observed size- and sex-dependent spawning aggregation patterns, in association with habitat volume and stock size, may influence cod catchability and thereby the assessment and exploitation patterns of this stock.     

Tests of larval retention in a tidally energetic environment reveal the complexity of the spatial structure in herring populations

Dispersion during the larval phase is of central importance in the dynamics of marine fish and invertebrate populations. Rapid transport or dispersion of larvae may contribute to connectivity and mixing, whereas spatial persistence (retention) is hypothesized to favour stock complexity and local subpopulations. Larval retention, while rarely quantified, may be defined in species with protracted spawning by the spatial co‐occurrence of larvae of different sizes or ages. The spatial distributions of larval Atlantic herring (Clupea harengus) were examined from 22 annual autumn surveys (1975–1998) and 9 spring surveys (1975–1984) from the Bay of Fundy, a region with large tides and residual flow. Larvae of all sizes (3–27 mm in length, from hatch to nearly 4 months post‐hatch) were observed each year in two major aggregations; one off southwestern Nova Scotia, and the other in the mid‐inner Bay of Fundy off the northwestern shore of Nova Scotia. Two similar aggregations were evident over 5 months later from 9 spring surveys (1975–1984), despite the residual flow that would have swept the larvae from the region within 1 month. Larval retention was apparent from overlapping centres of mass of different size (=age) classes of larvae, and tested using a size diversity index, based on the co‐occurrence of 1‐mm‐size categories, derived from protracted spawning of several weeks. Geospatial ‘hot spots’ (G_i* statistic) of four size (age) classes were evident at specific stations in the 50–100 m bathymetric zone and not elsewhere. These metrics provide quantitative measures of retention that may be applied to many ichthyoplankton data sets. One of the three main spawning areas collapsed during the study period after a period of intense fishing and failed to rebuild, but there was no substantial change in the location of larval hotspots in subsequent years. While larval retention does not directly relate to each spawning location, larval retention in the Bay of Fundy contributes to the complex ‘metapopulation’ structure of herring stocks in the western 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.     

Integration of ecology,demography and genetics to reveal population structure and persistence: a mini review and case study of stream‐dwelling Dolly Varden

Abstract – Spatial structures can drive population dynamics and persistence. The roles of space and dispersal on dynamics, however, remain largely unknown in wild populations, mainly because of the difficulty in extensive/intensive field surveys at multiple locations. Here, I integrate results of ecological, demographic and genetic studies to elucidate detailed population structure and to identify the mechanisms of population persistence in a stream fish. Spatial structures, such as habitat size and connectivity, are particularly important in river and freshwater networks because such ecosystems are highly vulnerable to human activities. These issues are explored by examining results from a decadal research on the stream‐dwelling Dolly Varden charr in the Sorachi River basin, Japan. More than 100 local habitats (i.e. tributaries) were examined for spatial structure of populations by, such as, species presence, population census, redd counts, analysis of population synchrony and microsatellite DNA analysis. The results indicate that (i) population demography is largely independent in each tributary, (ii) some large tributaries support self‐sustaining populations, (iii) despite small population sizes in most tributaries (<30 spawning females), local extinction is rare, except in human‐influenced areas and (iv) probabilities of extinction are likely reduced by immigration of fish from neighbouring tributaries. The evidence suggests that Dolly Varden in this river system function as a source‐sink or mainland‐island metapopulation structure and that processes influencing these structures vary within the stream network. Overall, local populations may be highly persistent as long as local populations are well connected. I also discuss the vulnerability of stream fishes to habitat alterations.     

Spatial assessment of predator–prey relationships in the North Sea: the influence of abiotic habitat properties on the spatial overlap between 0‐group cod and grey gurnard

The understanding of spatio‐temporal dynamics of marine ecosystems is crucial for ecosystem‐based fisheries management and climate change impact assessments. We quantified temporal changes in the distribution of 0‐group cod (Gadus morhua) and grey gurnard (Eutriglia gurnardus), a primary predator of 0‐group cod, with the help of regression kriging and assessed the temporal dynamics of the related spatial predator–prey overlap of these two species at different spatial scales. We analysed the robustness of relationships among abiotic habitat properties (temperature, salinity and depth) and abundance. Small cod was mainly found in low salinity areas of the Skagerrak but larger year classes were able to expand their distribution area towards the central and northern North Sea. In contrast, grey gurnard was mainly found in waters with salinities above 33 and temperatures above 14°C. This species has expanded its high density areas in the central North Sea northward over the last two decades. Recruitment success of cod was negatively correlated to a Moran's I cross‐correlation index, a proxy for the degree of spatial overlap between both species. Strong cod year classes overlapped less with grey gurnard at the large and medium spatial scale. In general, the relationships between abiotic habitat properties and abundance showed an increased inter‐annual variability, which was likely caused by underlying factors not taken into account in the distribution models. Thus assemblage modeling approaches combining the strength of different model types should be considered in the future to predict potential distribution patterns under climate change scenarios.     

Quantifying the influence of geography and environment on the northeast Atlantic mackerel spawning distribution

Mackerel (Scomber scombrus) in the northeast Atlantic have shown changes in distribution at certain times of the year, which have impacted their exploitation and management. In this study, mackerel spawning habitat over 21 recent years was characterised using generalised additive modelling, based on spatial egg density data collected every third year during targeted ichthyoplankton surveys. Mackerel spawning distribution was found to depend primarily on geographical variables (coordinates and bottom depth), with preferred spawning locations on the shelf‐edge from the north of the Iberian peninsula to the west of Scotland, with a maximum west of Ireland. Environmental drivers had a lesser influence on egg distribution. Dome shaped relationships were found with temperature and mixed layer depth, with respective optimum at 13°C and around –300 m. The model was used to reconstruct maps of the potential habitat (areas where conditions were suitable, but not necessarily used, for spawning). Little changes were observed over the years in the potential habitat, suggesting that the expansion of the egg distribution (realised habitat) was not triggered by changes in the environmental variables investigated. Little evidence was found for density‐dependent habitat selection. There was a tendency for mackerel to make more use of areas of lesser suitability in years with large stock size (1992 and 2010). This pattern, however, broke down in 2013, when stock size was the highest, as spawning occurred very south and concentrated in the most suitable habitat.     

Seasonal spawning and wind‐regulated retention‐dispersal of early life stage Atlantic cod (Gadus morhua) in a Newfoundland fjord

Sequential ichthyoplankton surveys were used to determine the spatial and temporal distribution of eggs and larvae over coastal spawning grounds of Atlantic cod (Gadus morhua) in Smith Sound, Trinity Bay, Newfoundland, during the spring and summer of 2006 and 2007. Egg densities showed similar patterns for both years with two peaks in abundance in spring (March–April) and late summer (late July). A clear progression of development stages (1–4) was observed in spring and summer in 2006 and summer in 2007, suggesting retention of eggs within the Sound during these periods. Modelled predictions of vertical egg distributions indicated eggs were broadly distributed near the surface during spring (March–April), but were concentrated below the pycnocline (>10 m) within the inner portions of the Sound during the summer months (July–August). Back‐calculated peaks in spawning based on water temperatures were estimated at 11 and 4 April for 2006 and 2007, respectfully, with late season peaks centred on 21–24 July for both years. Environmental data indicated cooler water temperatures and periods of high wind stress in spring, and warmer, calmer periods late summer, consistent with higher retention and faster development times on the spawning grounds later in the season. We conclude that spring and summer spawning events result in different distributions of early life stages and may lead to different distributions of juvenile and adult fish.     

Pacific cod in the Anthropocene: An early life history perspective under changing thermal habitats

The rapid decline in Pacific cod (Gadus macrocephalus, Gadidae) biomass following multiple Gulf of Alaska marine heatwaves (2014–2016 and 2019) may be one of the most dramatic documented changes in a sustainably managed marine fishery. As such, fisheries managers are exploring new recruitment paradigms for Pacific cod under novel environmental conditions. In this review, we address the challenges of managing and forecasting Pacific cod populations in the Eastern Pacific where thermal habitats for early life stages are undergoing varying rates of change across space and time. We use observational data to examine changes in distribution, abundance and demographics of the population from 1993 to 2020, and model contemporary and future changes of thermal habitat for both spawning success and age-0 juvenile growth potential. Results indicate that reduced spawning habitat and early life stage abundance may be a precursor to regional population decline, but the recent apparent increases in size-at-age of pre-recruits will have unknown impacts on future recruitment in these regions. We contend that continued monitoring of early life stages will be necessary to track changes in phenology and growth that likely determine size-at-age and the survival trajectories of year classes into the adult population. These include complex size- and temperature-dependent energetics spanning seasonal habitats through the first winter. Climate-ready management of Pacific cod will, therefore, require new process investigations beyond single-season surveys focused on one-life stage.     

A metapopulation perspective for salmon and other anadromous fish

Salmonids are an important component of biodiversity, culture and economy in several regions, particularly the North Pacific Rim. Given this importance, they have been intensively studied for about a century, and the pioneering scientists recognized the critical link between population structure and conservation. Spatial structure is indeed of prime importance for salmon conservation and management. At first glance, the essence of the metapopulation concept, i.e. a population of populations, widely used on other organisms like butterflies, seems to be particularly relevant to salmon, and more generally to anadromous fish. Nevertheless, the concept is rarely used, and barely tested. Here, we present a metapopulation perspective for anadromous fish, assessing in terms of processes rather than of patterns the set of necessary conditions for metapopulation dynamics to exist. Salmon, and particularly sockeye salmon in Alaska, are used as an illustrative case study. A review of life history traits indicates that the three basic conditions are likely to be fulfilled by anadromous salmon: (i) the spawning habitat is discrete and populations are spatially separated by unsuitable habitat; (ii) some asynchrony is present in the dynamics of more or less distant populations and (iii) dispersal links populations because some salmon stray from their natal population. The implications of some peculiarities of salmon life history traits, unusual in classical metapopulations, are also discussed. Deeper understanding of the population structure of anadromous fish will be advanced by future studies on specific topics: (i) criteria must be defined for the delineation of suitable habitats that are based on features of the biotope and not on the presence of fish; (ii) the collection of long‐term data and the development of improved methods to determine age structure are essential for correctly estimating levels of asynchrony between populations and (iii) several key aspects of dispersal are still poorly understood and need to be examined in detail: the spatial and temporal scales of dispersal movements, the origin and destination populations instead of simple straying rates, and the relative reproductive success of immigrants and residents.     

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.     

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.