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1.
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.  相似文献   

2.
We examined spatial variability in meristic and morphological characteristics of the branchial basket of sardine Sardinops sagax collected from four geographical regions around the southern African coast, namely Namibia and the South African west, south and east coasts. Our analysis tested the hypothesis of three putative sardine stocks off South Africa, one in each of the three geographical regions. We therefore collected fish data from Namibia to compare with South Africa, because sardine from the two countries are considered to be separate stocks. Morphometric measurements (gill arch length and gill raker spacing) and meristic data (number of gill rakers) were collected from the left side of the first gill arch from a total of 377 sardine, approximately equally divided between the regions. A multivariate general linear model with caudal length as covariate was used to assess differences among fish from the four regions and significant differences were observed, although not always consistently across all fish size classes. Small South Coast sardine had shorter gill arches than small West Coast sardine, but adults had gill arches of similar length, longer than those from Namibia and the East Coast. Small sardine from the South Coast had fewer gill rakers than small sardine from the West Coast, but larger fish had similar numbers of gill rakers, significantly more than sardine from Namibia and the East Coast. Sardine from the West and South coasts had similar gill raker spacings, which were smaller than those of fish from the East Coast and Namibia. Despite spatially and particularly temporally unbalanced sampling, we consider that these differences provide evidence of spatial variation in Benguela sardine phenotype and that it would support the hypothesis of discrete sardine stocks off Namibia and South Africa. The results are consistent with the hypothesis of three sardine stocks within the southern Benguela.  相似文献   

3.
The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.  相似文献   

4.
The development of suitable reference states for ecosystem-based management requires documentation of changes in structure and functioning of marine ecosystems, including assessment of the relative importance of bottom-up and top-down processes as drivers of change. We used monitoring data available from St Helena Bay, the most productive bay and an important nursery area situated on the west coast of South Africa, during 1950–2010 to reveal changes in the abiotic and biotic components. St Helena Bay in the 1950s showed similarities to 2000–2010 in terms of wind patterns, hydrology and phytoplankton. Upwelling, oxygen and nutrient concentrations in subthermocline water displayed pronounced decadal-scale variability. Primary production in St Helena Bay is variable, but consistently higher than that on the adjacent Namaqua shelf. Zooplankton size composition and biomass in August have changed markedly since the 1950s. During 2001–2010, mesozooplankton biomass in autumn was considerably lower than in summer, probably due to predation by small pelagic fish. Pelagic fish catch patterns and distributions have altered dramatically. Conservation measures, implemented to reverse past negative human impact, have benefitted marine mammals, the abundance of which has increased in the area, but additional conservation measures are necessary to reverse the decline in African penguins Spheniscus demersus. St Helena Bay shows a muted response to long-term change in the southern Benguela, with marked decadal variability but no clear long-term trend in oceanography and biogeochemistry. Changes in ecosystem boundary conditions and fishing pressure cannot be ignored as important drivers of change in the southern Benguela since the 1950s.  相似文献   

5.
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.  相似文献   

6.
The results of detailed morphological, experimental, field and modelling studies on various aspects of the trophic ecology of sardine Sardinops sagax and anchovy Engraulis encrasicolus in the Benguela ecosystem are synthesised, and differences in the trophodynamics of these two species are highlighted. Anchovy possess a relatively coarse branchial apparatus; feed predominantly by particulate-feeding and maximise their net energetic gain by using this feeding mode; show higher weight-standardised clearance rates than do sardine for prey >580μm; are most efficient at assimilating nitrogen from zooplankton and excrete <50% of ingested nitrogen; feed inefficiently on phytoplankton and derive the bulk of their dietary input from larger zooplankton; and maximise their scope for growth on mesozooplankton. In contrast, sardine possess a relatively fine branchial apparatus; feed predominantly by filter-feeding and maximise their net energetic gain by using this feeding mode; show higher weightstandardised clearance rates than do anchovy for prey <580μm; are most efficient at assimilating nitrogen from zooplankton but excrete >50% of ingested nitrogen; are able to feed on phytoplankton but derive the bulk of their dietary input from smaller zooplankton; and maximise their scope for growth on microzooplankton. These differences provide compelling evidence that anchovy and sardine are trophically distinct, and indicate that the two species show resource partitioning based on zooplankton size. The implications of these trophic differences for ecosystem functioning are discussed, and support the hypothesis that species alternations between anchovy and sardine, both in the southern Benguela and in other upwelling ecosystems, are likely to be trophodynamically mediated.  相似文献   

7.
Long-term ecosystem changes, such as regime shifts, have occurred in several marine ecosystems world-wide. Multivariate statistical methods have been used to detect such changes. A new method known as the sequential t-test algorithm for analysing regime shifts (STARS) is applied to a set of biological state variables as well as environmental and anthropogenic forcing variables in the southern Benguela. The method is able to correct for auto-correlation within time-series by a process known as prewhitening. All variables were tested with and without prewhitening. Shifts that were detected with both methods were termed robust. The STARS method detected shifts in relatively short time-series and identified when these shifts occurred without a priori hypotheses. Shifts were generally well detected at the end of time-series, but further development of the method is needed to enhance its performance for auto-correlated time-series. Since 1950, two major long-term ecosystem changes were identified for the southern Benguela. The first change occurred during the 1960s, caused predominantly by heavy fishing pressure but with some environmental forcing. The second change occurred in the early 2000s, caused mainly by environmental forcing. To strengthen these findings, further analyses should be carried out using different methods.  相似文献   

8.
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.  相似文献   

9.
A recently developed technique to estimate the lipid content of pelagic fish, using macroscopic visual assessment of their mesenteric fat, was applied to samples of southern Benguela sardine Sardinops sagax collected from commercial catches over the period 1996–2004. The data were analysed using general linear modelling, and temporal and spatial variably in lipid content was examined and compared with that of condition factor derived from length/mass relationships. Lipid content showed a clear seasonal cycle, being highest in May and lowest in August. This pattern was comparable with seasonal cycles in condition factor, although lipid content peaked 1–5 months later than did condition factor. A strong spatial effect was apparent in model results, with both lipid content and condition factor being highest in sardine from the South-East Coast. The correspondence in results from the fat staging technique, and those derived using condition-factor data for longer time-series, validate the new technique, and the higher level of resolution of the former compared with the latter argue for broader application of this technique.  相似文献   

10.
The study focuses on ichthyoplankton populations in the southern Angola Current, the Angola-Benguela Front and the coastal upwelling area of the northern and central Benguela Current. The horizontal and vertical distributions of eggs and larvae of sardine Sardinops sagax, anchovy Engraulis encrasicolus and horse mackerel Trachurus trachurus capensis were examined in relation to distribution patterns of temperature, salinity and dissolved oxygen. Samples were collected during February–March 2002 using a Bongo net and a multiple opening/closing net in five depth layers (0–30m, 30–60m, 60–100m, 100–140m and 140–220m). Eggs and larvae were restricted mainly to the shelf areas in the Angola-Benguela Font and southern Angola Current waters. Their vertical distributions in the Angola-Benguela Front and northern Benguela upwelling region showed a sharp decline in abundance at about 60m depth. This decline appeared to be correlated to the distribution of dissolved oxygen in the water column It is hypothesised that low oxygen concentrations have a strong impact on the development and survival of the early life-cycle stages of fish, and that recruitment of sardine and other pelagic species relies more on the upward extension of the oxygen minimum layer than was previously thought.  相似文献   

11.
The diet of African penguins Spheniscus demersus in Namibia consisted mainly of sardine Sardinops sagax in the 1950s. Since the collapse of pelagic fish stocks in the 1970s, birds fed mainly on bearded (pelagic) goby Sufflogobius bibarbatus, a low-energy prey species. We present diet data for African penguins breeding at Mercury Island, the largest colony for this species in Namibia, between 1996 and 2009. Bearded goby was the main prey item throughout the study period, both in terms of frequency of occurrence (67.8%; SD 31.2) and in terms of mass (59.2%; SD 31.5). Diet composition varied throughout the year as well as between years; birds occasionally fed on a variety of fish species other than bearded goby. In Namibia, poor prey abundance is considered as a major factor contributing to the decline of penguin numbers after the collapse of the sardine stocks. However, bearded goby appears to be relatively abundant along Namibia's southern coast and low prey quality rather than low abundance appears to be a key factor influencing population dynamics of African penguins and other marine top predators in southern Namibia.  相似文献   

12.
Spatial variability in phenotypic characteristics within a fish population may be used to infer the existence of multiple stocks, and knowledge of the population structure of exploited species is important for their sustainable management. In this study we investigated geographic variability in meristic (vertebral count) and morphometric (body shape) characteristics of sardine Sardinops sagax from three southern African regions: Namibia, the South African west coast, and the South African south coast. There were significant regional differences in vertebral counts, with Namibian fish tending to have more vertebrae than South African fish, but with no difference between sardine off the South African west and south coasts. Body shape was characterised using morphometric landmarks, and shape changes were investigated using geometric morphometrics, with influences on fish shape assessed using multivariate regressions. To remove the effects of allometry, the 22 shape variables (represented by Procrustes coordinates) were regressed against fish size (represented by centroid size). There was a significant allometric effect, which was removed by using the residuals of these regressions in further analyses. A multivariate multiple regression was applied to the 22 size-corrected shape variables and to three covariates: condition factor, stomach fullness and fat stage. The significant influence of these covariates was removed by using the residuals from this regression in a canonical variate analysis, where corrected shape variables were classified in relation to region, year and sex. The Mahalanobis distances from the canonical variate analysis differed significantly for most combinations of classifier variables, with clear separations among the three regions. Sardinops sagax off Namibia had thicker bodies and more contracted tail regions than South African fish, whereas S. sagax off the South African south coast had thinner bodies and smaller heads than those off the west coast. These results support the hypothesis of multiple sardine stocks around southern Africa.  相似文献   

13.
The northern Benguela stock of sardine Sardinops sagax used to be considered one of the major clupeoid stocks of the world; it supported an average annual catch of >700 000 tons throughout the 1960s. The stock has been in a depressed state for more than two decades, as demonstrated by annual catches that averaged around 50 000 tons between 1978 and 1989 and only slightly more in the 1990s. It has experienced fluctuations in abundance of several orders of magnitude during the most recent decade. Population size increased until 1992, when the acoustic estimate of biomass was about 750 000 tons. Catches increased accordingly, averaging 100 000 tons between 1992 and 1995, but from 1992 to 1996 the stock was in decline and the lowest annual catch in the history of the fishery was taken in 1996. Although there was a small increase during the last three years of the decade, the stock remains seriously depleted. Survey-based recruitment indices suggest that the changes in the 1990s were initiated by fluctuations in recruitment, but the decline was almost certainly exacerbated by continued fishing. Poor recruitment and decreasing catch rates between 1993 and 1996 in a number of other key resources suggest that system-wide environmental changes were an important factor in the decline of the sardine stock at that time. Anomalous oceanographic conditions, such as extensive hypoxic shelf waters in 1993/94 and a Benguela Niño in 1995, support this conclusion.  相似文献   

14.
Despite much public awareness surrounding the annual migration of sardine Sardinops sagax northward along the east coast of South Africa in winter each year, relatively little research effort has been expended to improve understanding of the ‘sardine run’. For this reason, a dedicated multidisciplinary survey, timed to coincide with the annual sardine run, was conducted off the East Coast in June and July of 2005. The major objective of the survey was to estimate the biomass of sardine off the East Coast during the run, and to compare this with biomass estimates collected during previous surveys conducted in this area during the late 1980s when the South African sardine population was at a considerably smaller size. We also collected data on the distribution of sardine and other small pelagic fish species and their eggs, the biological characteristics of sardine during the run, and data on the hydrography (temperature and currents) and lower trophic levels (phytoplankton and zooplankton) of the region. Results suggest that the biomass of sardine off the East Coast in winter remains relatively small and consistent, regardless of overall sardine population size. The narrow continental shelf to the east of Port Alfred, which is dominated offshore by the fast-flowing warm Agulhas Current, constrains the amount of suitable habitat for sardine and other clupeoids such as anchovy Engraulis encrasicolus, West Coast round herring Etrumeus whiteheadi and East Coast round herring Etrumeus teres, and hence precludes these species from attaining a high biomass in this region. Additionally, primary and secondary productivity levels are much lower than elsewhere on the western and eastern Agulhas Bank off the south coast of South Africa, suggesting that the sardine run is not a feeding migration. A previous hypothesis that the run is mainly a result of an expansion of the distributional range of these fish as conditions become favourable in winter due to sporadic cooling off the East Coast is also not entirely supported by results from the survey. It is suggested that a migration for the purposes of spawning off this coast when conditions become favourable is a more likely incentive for sardine to undertake this arduous journey, despite increased predation and poor feeding conditions.  相似文献   

15.
This paper gives an overview of the main living marine resources of Namibia. It focuses on the scientific research conducted during the past decade as input to the management of these resources. The distribution and habitats of the most important harvested species and the main seabird populations are briefly described and discussed. The life histories of the major exploited species are summarized, with emphasis on spatial and temporal spawning patterns, dispersal of early life stages, migration patterns of recruits and adults, and diet, the latter particularly as it relates to potential competition between species. A number of commercially important species, such as the hake Merluccius capensis and M. paradoxus, deep-sea red crab Chaceon maritae, West Coast rock lobster Jasus lalandii, skipjack tuna Katsuwonus pelamis, southern albacore Thunnus alalunga and to a lesser extent Cape horse mackerel Trachurus capensis, southern African sardine Sardinops sagax and Cape anchovy Engraulis capensis, are distributed across national boundaries, requiring regional cooperation in research and management. The history and current status of the major fisheries is discussed. Over the past 30–40 years total annual catches have declined from a peak of around 2 million tons in the late 1960s to less than a million tons in the 1990s. This decline has been due, mainly, to a collapse in the sardine stock in the late 1960s and 1970s, and a reduction in the catches of hake and horse mackerel under a conservative management strategy in the past decade. Changes in the abundance and distribution of commercially important species, as determined by acoustic and trawl surveys and catch-based analytical methods, are presented. The effect of major environmental anomalies on the distribution and abundance of the resources in recent years is discussed. The most dramatic anomaly in recent years was the wide-scale advection of low-oxygen water into the northern Benguela from the Angola Dome in 1994, and the subsequent Benguela Niño of 1995, which appear to have severely impacted the Namibian sardine population and many other resources. The present socio-economic value of the Namibian fishing industry is given together with the broad policy, legislation and formal structures for managing the living marine resources.  相似文献   

16.
Once one of the most numerous seabirds of the Benguela upwelling system, the population of Cape cormorants Phalacrocorax capensis has decreased by 60% in the past three decades and the species is listed as Near Threatened. Declines in prey availability and/or abundance brought about by recent changes in the distribution of pelagic fish stocks and industrial purse-seine fishing are hypothesised to be a key driver of seabird population decreases in the southern Benguela. We investigated the foraging behaviour of breeding Cape cormorants by deploying GPS and temperature–depth recorders on 24 breeding adults from three islands off the coast of South Africa, two of them to the north of Cape Point and a third farther south on the western Agulhas Bank. This provided the first measures of foraging dispersal by a cormorant in the Benguela system, and enabled a comparison of foraging behaviour between birds from these islands. Foraging trips of Cape cormorants lasted between 17 min and >7 h, at a maximum distance of between 2 and 58 km away from their colony. Foraging effort was significantly greater for birds from farther north off the West Coast in terms of trip duration, distance travelled, number of dives and time spent flying compared to those from the southernmost island (Dyer), which is probably a response to low prey availability in the north. Coastal reserves that exclude pelagic fishing from inshore feeding grounds around Cape cormorant breeding colonies may result in increased local prey availability, which would benefit Cape cormorant populations.  相似文献   

17.
We synthesise and update results from the suite of biophysical, larval-dispersal models developed in the Benguela Current ecosystem. Biophysical models of larval dispersal use outputs of physical hydrodynamic models as inputs to individual-based models in which biological processes acting during the larval life are included. In the Benguela, such models were first applied to simulate the dispersal of anchovy Engraulis encrasicolus and sardine Sardinops sagax ichthyoplankton, and more recently of the early life stages of chokka-squid Loligo reynaudii and Cape hakes Merluccius spp. We identify how the models have helped advance understanding of key processes for these species. We then discuss which aspects of the early life of marine species in the Benguela Current ecosystem are still not well understood and could benefit from new modelling studies.  相似文献   

18.
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 Corg 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 Corg 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.  相似文献   

19.
Catches of South African sardine Sardinops sagax have declined in recent years from about 200 000 t harvested annually during the period 2002–2006 to less than 100 000 t. Consequently, some companies are now importing sardine from sources elsewhere in the world to meet local demand for canned sardine and bait. This importation has the potential for the introduction of sardine pathogens, in particular the pilchard herpesvirus (PHV), which could have a negative impact on the currently small South African sardine population. The aims of the current study were to determine whether PHV is present in the local sardine population and to assess the extent to which sardine is being imported into the country and whether imported fish are from countries where the virus is known to be endemic. Fish sampled from South Africa’s western (n = 150), southern (n = 182) and eastern (n = 96) putative stocks of S. sagax were analysed for the presence of PHV using real-time quantitative polymerase chain reaction (qPCR). The origin and amount of potentially infected material imported into South Africa during the period 2010–2014 was also assessed. None of the South African sardine collected during this study tested positive for PHV, suggesting that active PHV was not prevalent in the local population of S. sagax at the time of this study. Between 56 000 and 71 000 t of frozen sardine was imported annually into South Africa from countries where S. sagax occurs, including some from areas (Australia and New Zealand) where sardine infection by PHV is known to be endemic. Hence, it is plausible that the PHV pathogen, capable of perpetuating infections in local sardine populations, could be imported into South Africa along with the importation of frozen sardine. Should local sardine be naïve to the virus, as suggested by this study, then the population is at risk of infection and precautions against such must be taken.  相似文献   

20.
Discussion of mesopelagic fish within the southern Benguela system necessitates recognition of both oceanic and neritic realms, demarcated by the continental shelf-break. In the eastern South Atlantic (FAO Area 47), the stock size of two oceanic, mesopelagic families (Myctophidae, Sternoptychidae) have been estimated at 8–12 million and 750 000 metric tons respectively. Each possesses pseudoceanic species, populations of which constitute a possible alternative fishery resource to conventional pelagic stocks. Of these upper-slope/shelf derivatives, Lampanytodes hectoris (Myctophidae) and Maurolicus muelleri (Sternoptychidae) are most abundant. In the southern Benguela system (Orange River mouth to Cape Point), both species appear to be restricted mainly to the region east of the offshore divergence, corresponding roughly to the continental shelf-break. Maurolicus muelleri has not been taken by the fishery, but purse-seine landings of Lampanyctodes hectoris have accounted for up to some 10 per cent of the total South African pelagic catch. Anomalies in the commercial catch (sexual disparity, sexual immaturity) have been linked to expatriatism. This phenomenon is re-examined in the light of more recent data from research sampling in the southern Benguela region, and alternative biological hypotheses are presented to explain the anomalies. It is suggested that the energy "bonus" derived from vertical migration in summer is enhanced both by a physical component, termed "cycle riding", and by a biochemical component, linked to long-chain wax esters. Seasonal variation in percentage lipid concentration also appears to be associated with reproductive activity in the late winter-spring spawning period. Vertical migration may be reduced at this time, in tandem with the cessation of upwelling within the system and paucity of food in the upper water column. Comparative data for Maurolicus muelleri are presented.  相似文献   

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