Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela

A regime shift is considered to be a sudden shift in structure and functioning of a marine ecosystem, affecting several living components and resulting in an alternate state. According to this definition, regime shifts differ from species replacement or alternation of species at similar trophic levels, whereby the ecosystem is not necessarily significantly altered in terms of its structure and function; only its species composition changes. This paper provides an overview of regime shifts, species replacements and alternations that have been observed in the northern and southern Benguela ecosystems over the past few decades. Bottom-up control, initiating and sustaining regime shifts or species replacements via environmental forcing, is documented for both the southern and the northern Benguela ecosystems. Fishing (a case of top-down control) appears to have played an important role in regime shift processes in the Namibian ecosystem. Very low biomass levels of exploited fish stocks associated with less efficient energy transfer in the northern Benguela are indicative of a regime shift. Very high biomass levels have been reached in the southern Benguela in the 2000s. However the alternation between sardine and anchovy that has been observed in the southern Benguela over the last two decades appears not to have had major effects on the overall functioning of the ecosystem. The consequences of regime shifts for exploitation are highlighted, suggesting that fisheries managers should move towards a more effective ecosystem approach to fisheries.     

Regime shifts: Can ecological theory illuminate the mechanisms?

“Regime shifts” are considered here to be low-frequency, high-amplitude changes in oceanic conditions that may be especially pronounced in biological variables and propagate through several trophic levels. Three different types of regime shift (smooth, abrupt and discontinuous) are identified on the basis of different patterns in the relationship between the response of an ecosystem variable (usually biotic) and some external forcing or condition (control variable). The smooth regime shift is represented by a quasi-linear relationship between the response and control variables. The abrupt regime shift exhibits a nonlinear relationship between the response and control variables, and the discontinuous regime shift is characterized by the trajectory of the response variable differing when the forcing variable increases compared to when it decreases (i.e., the occurrence of alternative “stable” states). Most often, oceanic regime shifts are identified from time series of biotic variables (often commercial fish), but this approach does not allow the identification of discontinuous regime shifts. Recognizing discontinuous regime shifts is, however, particularly important as evidence from terrestrial and freshwater ecosystems suggests that such regime shifts may not be immediately reversible. Based on a review of various generic classes of mathematical models, we conclude that regime shifts arise from the interaction between population processes and external forcing variables. The shift between ecosystem states can be caused by gradual, cumulative changes in the forcing variable(s) or it can be triggered by acute disturbances, either anthropogenic or natural. A protocol for diagnosing the type of regime shift encountered is described and applied to a data set on Georges Bank haddock, from which it is concluded that a discontinuous regime shift in the abundance of haddock may have occurred. It is acknowledged that few, if any, marine data are available to confirm the occurrence of discontinuous regime shifts in the ocean. Nevertheless, we argue that there is good theoretical evidence for their occurrence as well as some anecdotal evidence from data collection campaigns and that the possibility of their occurrence should be recognized in the development of natural resource management strategies.     

Covariance among North Sea ecosystem state indicators during the past50 years — Contrasts between coastal and open waters

Regime shift and principal component analysis of a spatially disaggregated database capturing time-series of climatic, nutrient and plankton variables in the North Sea revealed considerable covariance between groups of ecosystem indicators. Plankton and climate time-series span the period 1958–2003, those of nutrients start in 1980. In both regions, the period from 1989 to 2001 identified in principal component 1 had warmer surface waters, higher Atlantic inflow and stronger winds, than the periods before or after. However, it was preceded by a regime shift in both open (PC2) and coastal (PC3) waters during 1977 towards more hours of sunlight and higher water temperature, which lasted until 1997. The relative influence of nutrient availability and climatic forcing differed between open and coastal North Sea regions. Inter-annual variability in phytoplankton dynamics of the open North Sea was primarily regulated by climatic forcing, specifically by sea surface temperature, Atlantic inflow and co-varying wind stress and NAO. Coastal phytoplankton variability, however, was regulated by insolation and sea surface temperature, as well as Si availability, but not by N or P. Regime shifts in principal components of hydrographic and climatic variables (explaining 55 and 61% of the variance in coastal and open water variables) were detected using Rodionov's sequential t-test. These shifts in hydroclimatic variables which occurred around 1977, 1989, 1997 and 2001, were synchronized in open and coastal waters, and were tracked by open water chlorophyll and copepods, but not by coastal plankton. North–central–south or open-coastal spatial breakdowns of the North Sea explained similar amounts of variability in most ecosystem indicators with the exception of diatom abundance and chlorophyll concentration, which were clearly better explained using the open-coastal configuration.     

Abrupt environmental shift associated with changes in the distribution of Cape anchovy Engraulis encrasicolus spawners in the southern Benguela

Cape anchovy Engraulis encrasicolus spawners in the southern Benguela showed an eastward shift in their distribution on the Agulhas Bank that occurred abruptly in 1996 and has since persisted. We assessed whether this shift was environmentally mediated by examining sea surface temperature data from different regions of the Agulhas Bank, which showed that in 1996 the inner shelf of the Agulhas Bank to the east of Cape Agulhas abruptly became 0.5°C colder than in previous years and has since remained that way. In addition, signals, coherent with the 1996 shift recorded in sea surface temperatures, were also found in atmospheric surface pressure and zonal wind data for that region; interannual coastal SST variability is also shown to be correlated with zonal wind-stress forcing. As a result, increased wind-induced coastal upwelling east of Cape Agulhas is proposed as the main driver of the observed cooling in the coastal region. The synchrony between the environmental and biological signals suggests that the eastward shift in anchovy spawner distribution was environmentally mediated and arose from a change in environmental forcing that altered the relative favourability for spawning between regions to the west and east of Cape Agulhas. The results highlight how a relatively minor change in environmental conditions can lead to a drastic spatial reorganisation of the life history of one species in an ecosystem.     

An integrated approach for assessing the relative significance of human pressures and environmental forcing on the status of Large Marine Ecosystems

An ecosystem approach to the management of the marine environment has received considerable attention over recent years. However, there are few examples which demonstrate its practical implementation. Much of this relates to the history of existing marine monitoring and assessment programmes which (for many countries) are sectoral, making it difficult to integrate monitoring data and knowledge across programmes at the operational level.To address this, a scientific expert group, under the auspices of the International Council for the Exploration of the Sea (ICES), prepared a plan for how ICES could contribute to the development of an Integrated Ecosystem Assessment (IEA) for the North Sea by undertaking a pilot study utilising marine monitoring data. This paper presents the main findings arising from the expert group and in particular it sets out one possible integrated approach for assessing the relative significance of environmental forcing and fishing pressure on the ecological status of the North Sea, it then compares the findings with assessments made of other Large Marine Ecosystems (LMEs).We define the North Sea ecosystem on the basis of 114 state and pressure variables resolved as annual averages between 1983 and 2003 and at the spatial scale of ICES rectangles. The paper presents results of integrated time-series and spatial analysis which identifies and explains significant spatial and temporal gradients in the data. For example, a significant shift in the status of the North Sea ecosystem (based upon 114 state-pressure variables) is identified to have occurred around 1993. This corresponds to previously documented shifts in the environmental conditions (particularly sea surface temperature) and changes in the distribution of key species of plankton (Calanus sp.), both reported to have occurred in 1989. The difference in specific timing between reported regime shifts for the North Sea may be explained, in part, by time-lag dependencies in the trophic structure of the ecosystem with shifts in higher trophic levels occurring later than 1989.By examining the connection (or relatedness) between ecosystem components (e.g. environment, plankton, fish, fishery and seabirds) for the identified regime states (1983–1993; 1993–2003) we conclude that both the North Sea pelagic and benthic parts of the ecosystem were predominantly top-down (fishery) controlled between 1983 and 1993, whereas between 1993 and 2003 the pelagic stocks shifted to a state responding mainly to bottom-up (environment) influences. However, for the demersal fish stocks between 1993 and 2003 top-down (fishery) pressure dominated even though over this period significant reductions in fishing pressure occurred. The present analysis, therefore, provides further evidence in support of the need for precautionary management measures taken in relation to setting fishery quotas.     

Variations in the abundance of fisheries resources and ecosystem structure in the Japan/East Sea

Evidence supports the hypothesis that two climatic regime shifts in the North Pacific and the Japan/East Sea, have affected the dynamics of the marine ecosystem and fisheries resources from 1960 to 2000. Changes in both mixed layer depth (MLD) and primary production were detected in the Japan/East Sea after 1976. The 1976 regime shift appears to have caused the biomass replacement with changes in catch production of major exploited fisheries resources, including Pacific saury, Pacific sardine and filefish. Both fisheries yield and fish distribution are reflected in these decadal fluctuations. In the 1960s and 1990s, common squid dominated the catches whereas in the 1970s and 1980s, it was replaced by walleye pollock. In the post-1988 regime shift, the distribution of horse mackerel shifted westward and southward and its distributional overlap with common mackerel decreased. The habitat of Pacific sardine also shifted away from mackerel habitats during this period. To evaluate changes in the organization and structure of the ecosystem in the Japan/East Sea, a mass-balanced model, Ecopath, was employed. Based on two mass-balanced models, representing before (1970–75) and after (1978–84) the 1976 regime shift, the weighted mean trophic level of catch increased from 3.09 before to 3.28 after. Total biomass of species groups in the Japan/East Sea ecosystem increased by 15% and total catch production increased by 48% due to the 1976 regime shift. The largest changes occurred at mid-trophic levels, occupied by fishes and cephalopods. The dominant predatory species shifted from cephalopods to walleye pollock due to the 1976 regime shift. It is concluded that the climatic regime shifts caused changes in the structure of the ecosystem and the roles of major species, as well as, large variations in biomass and production of fisheries resources.     

Detecting regime shifts in the ocean: Data considerations

We review observational data sets that have been used to detect regime shifts in the ocean. Through exploration of data time series we develop a definition of a regime shift from a pragmatic perspective, in which a shift is considered as an abrupt change from a quantifiable ecosystem state. We conclude that such changes represent a restructuring of the ecosystem state in some substantial sense that persists for long enough that a new quasi-equilibrium state can be observed. The abruptness of the shift is relative to the life-scale or the reproductive time-scale of the higher predators that are influenced by the shift. In general, the event-forcing is external to the biological ecosystem, usually the physical climate system, but we also identify shifts that can be ascribed to anthropogenic forcing, in our examples fishing. This pragmatic definition allows for several different types of regime shift ranging from simple biogeographic shifts to non-linear state changes. In practice it is quite difficult to determine whether observed changes in an oceanic ecosystem are primarily spatial or temporally regulated. The determination of ecosystem state remains an unresolved, and imprecise, oceanographic problem.We review observations and interpretation from several different oceanic regions as examples to illustrate this pragmatic definition of a regime shift: the Northeast Pacific, the Northwest and Northeast Atlantic, and Eastern Boundary Currents. For each region, different types of data (biological and physical) are available for differing periods of time, and we conclude, with varying degrees of certainty, whether a regime shift is in fact detectable in the data.     

Changes in the trophic structure of the southern Benguela before and after the onset of industrial fishing

Despite a human presence in the Benguela region for at least one million years, exploitation of marine resources by European seafarers only began in earnest in the 1400s. Ecopath with Ecosim was used to construct and compare mass-balanced foodweb models of the southern Benguela ecosystem, representing the following eras of human influence: aboriginal (10 000 BP–1651), pre-industrial (1652–1909), industrial (1910–1974) and post-industrial (1975–present). Biomass at higher trophic levels (TLs) decreased over the periods examined, whereas that of sardine and anchovy increased in the early 2000s, reflected by the decline in weighted TL of the community (excluding plankton). Fishing became an important predatory impact, taking over consumption of small pelagics and horse mackerel from declined natural predators such as hake. Harvesting of apex predators such as seals and seabirds during the pre-industrial era meant that the mean TL of the catch declined markedly between the pre-industrial (1900) and industrial (1960) models. Biomass removals by fishing have increased substantially over time. Total biomass, consumption, respiration, production and throughput decreased from the pristine model to 1960 and then increased again in the 2000s, probably influenced by the abnormally high small pelagic biomass in the early 2000s. Three additional alternate scenarios were examined for each of the retrospective models, in particular to explore the effects of removing large fish and forage fish from the system. Although biomasses and consumption of various groups in these scenarios differed from base models, indicators such as TL of the community and piscivore groups, and the diversity indices, were not altered much, suggesting that outputs from such retrospective models in the form of derived, relative indicators, may be more robust than comparisons of absolute flows, although the latter provide supplementary inferences. Although South African fisheries have certainly impacted ecosystem structure since their commencement, these effects are in addition to natural (specifically environmental) forcing that has always been influencing the system. Fishing stress at the ecosystem level and the collapse of small pelagic stocks may lead to a shift toward a bottom-up trophic control mechanism becoming the dominant driver of ecosystem dynamics, increasing the impact of environmental events including climate change. It is thus possible that pristine systems were not as severely affected by environmental anomalies as are modern systems.     

东北大西洋北海渔场鱼类群落结构年际变化研究

根据2001-2015年东北大西洋北海渔场进行的国际底拖网调查渔获数据,采用生物多样性指数和多元统计分析研究该海域群落结构的年际变化,并利用格局转变贯序t检验的方法研究鱼类种群的转变规律,结合环境因素与捕捞因素分析群落结构变化的原因。结果显示:2001-2015年北海渔场共出现280种渔业资源,其中鱼类有222种,资源丰度波动较大;物种多样性整体呈上升趋势。聚类分析和非度量多维标度排序分析表明,研究期间大致分为2001-2003年、2004-2011年和2012-2015年3个阶段。大西洋鲱分别在2004年和2014年种群结构发生格局转变,格局转变指数（RSI）分别为-0.45和0.41;黑线鳕在2003年和2012年格局发生转变,RSI值分别为-0.58和-0.66;黍鲱在2014年格局发生转变,RSI值为2。通过对环境因素与捕捞因素的分析发现,北海渔场群落格局第一次发生转变主要受捕捞因素影响,第二次发生转变主要受环境因素影响。     

The North Sea regime shift: Evidence, causes, mechanisms and consequences

This paper focuses on the ecosystem regime shift in the North Sea that occurred during the period 1982–1988. The evidence for the change is seen from individual species to key ecosystem parameters such as diversity and from phytoplankton to fish. Although many biological/ecosystem parameters and individual species exhibited a stepwise change during the period 1983–1988, some indicators show no evidence of change. The cause of the regime shift is likely to be related to pronounced changes in large-scale hydro-meteorological forcing. This involved activating of complex intermediate physical mechanisms which explains why the exact timing of the shift can vary from 1982 to 1988 (centred around two periods: 1982–1985 and 1987–1988) according to the species or taxonomic group. Increased sea surface temperature and possibly change in wind intensity and direction at the end of the 1970s in the west European basin triggered a change in the location of an oceanic biogeographical boundary along the European continental shelf. This affected both the stable and substrate biotope components of North Sea marine ecosystems (i.e. components related to the water masses and components which are geographically stable) circa 1984. Large-scale hydro-climatic forcing also modified local hydro-meteorological parameters around the North Sea after 1987 affecting the stable biotope components of North Sea ecosystems. Problems related to the detection and quantification of an ecosystem regime shift are discussed.     

Accumulation of organic and inorganic carbon in Pliocene–Pleistocene sediments along the SW African margin

The Ocean Drilling Program Leg 175 recovered a unique series of stratigraphically continuous sedimentary sections along the SW African margin, an area which is presently affected by active coastal upwelling. The accumulation rates of organic and inorganic carbon are a major component of this record. Four Leg 175 sites (1082, 1084, 1085, 1087) are chosen as part of a latitudinal transect from the present northern to southern boundaries of the Benguela Current upwelling system, to decipher the Pliocene–Pleistocene history of biogenic production and its relationship with global and local changes in oceanic circulation and climate. The pattern of CaCO₃ and C_org mass accumulation rates (MARs) over 0.25-Myr intervals indicates that the evolution of carbon burial is highly variable between the northern and the southern Benguela regions, as well as between sites that have similar hydrological conditions. This, as well as the presence over most locations of high-amplitude, rapid changes of carbon burial, reflect the partitioning of biogenic production and patterns of sedimentation into local compartments over the Benguela margin. The combined mapping of CaCO₃ and C_org MARs at the study locations suggests four distinct evolutionary periods, which are essentially linked with major steps in global climate change: the early Pliocene, the mid-Pliocene warm event, a late Pliocene intensification of northern hemisphere glaciation and the Pleistocene. The early Pliocene spatially heterogeneous patterns of carbon burial are thought to reflect the occurrence of mass-gravitational movements over the Benguela slope which resulted in disruption of the recorded biogenic production. This was followed (3.5–3 Ma) by an episode of peak carbonate accumulation over the whole margin and, subsequently, by the onset of Benguela provincialism into a northern and a southern sedimentary regime near 2 Ma. This mid and late Pliocene evolution is interpreted as a direct response to changes in the ventilation of bottom and intermediate waters, as well as to dynamics of the subtropical gyral circulation and associated wind stress.     

A comparison of condition factor and gonadosomatic index of sardine Sardinops sagax stocks in the northern and southern Benguela upwelling ecosystems, 1984–1999

Time-series of condition factor (CF) and gonadosomatic index (GSI) were generated using general linear models (GLM) for sardine Sardinops sagax stocks in the northern and southern Benguela ecosystems over the period 1984–1999. During this period the biomass of sardine in the northern Benguela remained at relatively low levels of <500 000 tons, whereas that of southern Benguela sardine increased 40-fold to 1.3 million tons. The GLMs explained 27 and 45% of the observed variation in CF, and 32 and 28% of the observed variation in GSI, for sardine in the northern and southern Benguela subsystems respectively. Whereas the sardine CF in the northern Benguela remained stable over time, that for the southern Benguela stock declined steadily during the study period. Sardine CF showed a seasonal cycle in the southern but not in the northern Benguela. Time-series of GSI showed high interannual variability but no trends in either subsystem, and the seasonal pattern was similar for both stocks. The lack of coherence between the CF time-series for sardine in the two subsystems further suggests that sardine stocks in the northern and southern Benguela subsystems are independent.     

Exploring the spatial distribution patterns of South African Cape hakes using generalised additive models

We developed delta generalised additive models (GAMs) to predict the spatial distribution of different size classes of South African hakes, Merluccius capensis and M. paradoxus, using demersal trawl survey data and geographical (latitude and longitude) and environmental features (depth, temperature, bottom dissolved oxygen and sediment type). Our approach consists of fitting, for each hake size class, two independent models, a binomial GAM and a quasi-Poisson GAM, whose predictions are then combined using the delta method. Delta GAMs were validated using an iterative cross-validation procedure, and their predictions were then employed to produce distribution maps for the southern Benguela. Delta GAM predictions confirmed existing knowledge about the spatial distribution patterns of South African hakes, and brought new insights into the factors influencing the presence/absence and abundance of these species. Our GAM approach can be used to produce distribution maps for spatially explicit ecosystem models of the southern Benguela in a rigorous and objective way. Ecosystem models are critical features of the ecosystem approach to fisheries, and distribution maps constructed using our GAM approach will enable a reliable allocation of species biomasses in spatially explicit ecosystem models, which will increase trust in the spatial overlaps and, therefore, the trophic interactions predicted by these models.     

Density-dependent changes in reproductive parameters and condition of southern Benguela sardine Sardinops sagax

The sardine Sardinops sagax population in the southern Benguela has undergone substantial fluctuations in size over the past 50 years, collapsing from an apparently large population in the 1950s to low levels in the mid-1960s, remaining low for the next two decades, and recovering from the late 1980s to a population size that is now similar to or larger than that which occurred during the 1950s. Marked changes in condition and reproductive parameters of sardine have also occurred during this period; condition and standardised gonad mass are higher and length-at-maturity is lower at low population size compared with high population size. The correspondence between the temporal patterns in condition, reproductive parameters and population size are strongly suggestive of density-dependence, and indicate a compensatory response arising from reduced intra-specific competition. This is likely to have resulted from greater per capita food intake, improved body condition and hence faster growth, thus enabling fish to achieve maturation at a presumably younger age and smaller size. Biological parameters did not vary in or out of phase with time-series of sea surface temperature in the southern Benguela, weakening the hypothesis of environmentally mediated changes in these parameters and hence providing support for the hypothesis of a direct density-dependent response by sardine.     

Methods for detecting regime shifts in large marine ecosystems: a review with approaches applied to North Pacific data

This note provides a brief review of five analytical methods previously used or promoted for diagnosing regime shifts in marine ecosystems. The methods discussed are: (i) principal components analysis, (ii) compositing average standard deviates, (iii) autoregressive moving average and intervention analysis modeling, (iv) vector autoregressive process modeling, and (v) Fisher information. Assessments of the relative strengths and weaknesses for the different analytical approaches are also offered. Some of these methods are applied to a collection of fishery oceanographic time series for the N. Pacific to illustrate aspects of their relative utility and limitations for diagnosing regime shift behavior. One recommendation for future studies is to analyze biotic and abiotic time series separately in order to identify ecosystem state variables of interest and to better isolate ecosystem behaviors from other influences like environmental change. Methods that allow for quantitative assessments of the statistical significance of hypothesized regime shifts should be favored over those that do not. Analyses of especially large collections of time series may benefit from first using a data compression technique and then applying one of the methods that are more appropriate for just one or a small number of time series. Because of the difficulties in observing and adequately documenting many aspects of marine ecosystem variability, it is crucial that future research attempt to combine empirical studies of large marine ecosystems with theoretical and modeling studies of other systems for which the dynamics and predictability are better understood. With such a comparative approach it should be possible to refine conceptual and simulation models, while also identifying crucial gaps in existing observations.     

Downscaling biogeochemistry in the Benguela eastern boundary current

Dynamical downscaling is developed to better predict the regional impact of global changes in the framework of scenarios. As an intermediary step towards this objective we used the Regional Ocean Modeling System (ROMS) to downscale a low resolution coupled atmosphere–ocean global circulation model (AOGCM; IPSL-CM4) for simulating the recent-past dynamics and biogeochemistry of the Benguela eastern boundary current. Both physical and biogeochemical improvements are discussed over the present climate scenario (1980–1999) under the light of downscaling.Despite biases introduced through boundary conditions (atmospheric and oceanic), the physical and biogeochemical processes in the Benguela Upwelling System (BUS) have been improved by the ROMS model, relative to the IPSL-CM4 simulation. Nevertheless, using coarse-resolution AOGCM daily atmospheric forcing interpolated on ROMS grids resulted in a shifted SST seasonality in the southern BUS, a deterioration of the northern Benguela region and a very shallow mixed layer depth over the whole regional domain. We then investigated the effect of wind downscaling on ROMS solution. Together with a finer resolution of dynamical processes and of bathymetric features (continental shelf and Walvis Ridge), wind downscaling allowed correction of the seasonality, the mixed layer depth, and provided a better circulation over the domain and substantial modifications of subsurface biogeochemical properties. It has also changed the structure of the lower trophic levels by shifting large offshore areas from autotrophic to heterotrophic regimes with potential important consequences on ecosystem functioning. The regional downscaling also improved the phytoplankton distribution and the southward extension of low oxygen waters in the Northern Benguela. It allowed simulating low oxygen events in the northern BUS and highlighted a potential upscaling effect related to the nitrogen irrigation from the productive BUS towards the tropical/subtropical South Atlantic basin. This study shows that forcing a downscaled ocean model with higher resolution winds than those issued from an AOGCM, results in improved representation of physical and biogeochemical processes.     

Remotely sensed variability of temperature and chlorophyll in the southern Benguela: upwelling frequency and phytoplankton response

High-resolution (1km) satellite data from the NOAA AVHRR (Advanced Very High Resolution Radiometer) and OrbView-2 SeaWiFS (Sea-viewing Wide Field-of-view Sensor) are used to investigate the upper layer dynamics of the southern Benguela ecosystem in more detailed space and time scales than previously undertaken. A consistent time-series of daily sea surface temperature (SST) and chlorophyll a concentration images is generated for the period July 1998–June 2003, and a quantitative analysis undertaken. The variability in SST, upwelling and phytoplankton biomass is explored for selected biogeographic regions, with particular focus on intra-seasonal time scales. The location and emergence of upwelling cells are clearly identified along the length of the southern Benguela, being distinct on the narrow inner and the mid-continental shelves. Most notable is the rapidly pulsating nature of the upwelling, with intense warm/cold events clearly distinguished. The phytoplankton response to this physical forcing is described. Chlorophyll concentration on the inner shelf largely mirrors the pattern of SST variability, similarly dominated by event-scale processes. Over the mid-shelf, higher chlorophyll is observed throughout all seasons, although low biomass occurs during winter. The variability of the offshore extent of SST and chlorophyll is identified at locations of differing shelf width. Cooler upwelled water is confined primarily to the narrow inner-shelf, with event-scale pulses extending considerable distances offshore. Agulhas Current influences are readily observed, even on the Cape Peninsula inner-shelf. Chlorophyll concentrations vary considerably between the locations of differing shelf width. SST, upwelling and phytoplankton indices are derived for selected locations to quantify the intra-seasonal variations. The SST indices show marked temperature changes associated with rapid pulsation on the event scale. No strong seasonal signal is evident. In contrast, the upwelling indices display a strong seasonal signal, with most intense upwelling occurring in spring/summer in the south. The phytoplankton response to the seasonal upwelling index differs between the selected locations. This study concludes that, although low-resolution SST and chlorophyll data may be useful for investigating general patterns over large scales, higher resolution data are necessary to identify finer scale spatial and temporal variability, especially in the inshore coastal zones.     

Climatic regime shifts and their impacts on marine ecosystem and fisheries resources in Korean waters

There were climatic regime shifts over the North Pacific in 1976 and 1988 which affected the dynamics of the marine ecosystem and fisheries resources in Korean waters. Precipitation in Korean waters showed a decadal scale climatic jump, especially of Ullungdo Island, reflecting the regime shift that occurred in the North Pacific. The variation was also detected in East Asian atmospheric systems. The Aleutian Low and North Pacific High Pressure Systems showed substantial changes in 1976 and around 1987–89. 1976 was an unusually warm year for Korea; mean sea surface temperature (SST) was higher than ‘normal’ and was accompanied by a northward shift in the thermal front. Post 1976, the volume transport of the Kuroshio Current increased and higher seawater and air temperatures persisted until 1988. Other shifts occurred after 1976 such as an increase in mixed layer depth (MLD) and biological changes in the ecosystem of Korean waters including decreases in spring primary production and an increase in autumn primary production. Primary production increased again after 1988, and was followed by a significant increase in zooplankton biomass after 1991. The 1976 regime shift was manifested by a decreased biomass and production of saury, but an increase in biomass and production of sardine and filefish in Korean waters. After 1988, recruitment, biomass, and production of sardine collapsed while those of mackerel substantially increased. Based on these observations, hypotheses on the relationship between the climate-driven oceanic changes and changes in fisheries resources were developed and are discussed.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

Relating a recruitment shift of Patagonian grenadier (Macruronus magellanicus Lönnberg) to large-scale environmental changes off Southern Chile

The SE Pacific stock of Patagonian grenadier (Macruronus magellanicus) showed evidence of an abrupt reduction in recruitment after 2000. This drop exceeded expectations from changes in the spawning stock biomass (SSB), indicating a change in the stock-recruitment relationship (S-R). We evaluated whether variability in recruitment could be explained by concurrent changes in three environmental indices: sea-surface temperature anomaly (SSTA); southern oscillation index (SOI); and latitudinal position of the west wind drift bifurcation (WWDL). Continuous and discrete effects of these indices were tested as covariates in linear log-log and non-linear Ricker's S-R models. Discrete effects represented regime shifts detected in SSTA (1998), SOI (1998) and WWDL (1995). While SSTA was the only continuous variable found to be significant, the discrete 1998 regime shift supported the most informative model. The best Ricker's model considered a discrete intercept change in the same year: 1998. Although a spurious correlation between SSTA and S-R changes is possible, SSTA may be reflecting major physical or biological changes relevant to M. magellanicus juveniles in the SE Pacific.