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1.
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.  相似文献   

2.
Diel patterns in survey trawl catches for the Cape hakes Merluccius capensis and M. paradoxus off Namibia were studied in order to examine the effect of diel bias on catchability, and its implication for survey abundance estimation and the consistency of the survey time-series. Catch rates (numbers per haul) by species and length from summer demersal biomass surveys conducted during the period 2002–2013 were used, together with a computation of the corresponding light-level data from which the solar zenith angles were obtained. Generalised additive models were fitted to assess the relationship between the catches and a number of explanatory variables. Significant covariates were zenith angle, depth and geographical position. The final models explained 78% and 59% of the variability in catch rates of M. capensis and M. paradoxus, respectively. For M. capensis, the response to zenith angle increased sharply for values above 100°, which represents the time between sunset and sunrise. For M. paradoxus there was a moderate increase in the response to zenith angle during the night. In cases where some fishing took place at night in shallow water, the survey results for M. capensis were more greatly affected than was the case for M. paradoxus, which is related to the different depth preference of the two species. Fishing in depths shallower than 400?m outside daylight hours should therefore be avoided in order to reduce bias and ensure consistency in abundance estimates from surveys.  相似文献   

3.
The two sympatric species of Cape hake, Merluccius capensis and M. paradoxus, have been the main targets of bottom-trawl fisheries off Namibia for several decades. The feeding ecology of these hakes has been studied mainly using stomach content analyses and thus there remain some gaps in our knowledge about food assimilated over the longer term. In this study, we used fatty acid (FA) profiles to characterise the dietary relationships of M. capensis and M. paradoxus. Muscle samples from hake (n=110) and their known prey (n=68) were collected during trawl surveys off Namibia during 2011. Significant differences between the neutral FA profiles of the hake populations were detected in December 2011 but not in January 2011, an indication of temporal variations in diet and resource partitioning. Comparisons of the neutral FAs in hake and the total FAs of potential prey showed no clear trophic connections, with the exception of flying squid Todarodes sagittatus, which had FA profiles very similar to those of M. paradoxus in December 2011. Our results highlight the complex and temporally shifting relationships that exist between hake and the large pool of prey available to them, and between the two hake species that overlap in their feeding habits and distribution within the highly productive Benguela Current region.  相似文献   

4.
Optimal and sustainable management of fish resources cannot be ensured without a thorough understanding of the migration patterns and population (demographic stock) structure. Recent studies suggest that these aspects of the economically and ecologically important deepwater hake Merluccius paradoxus are not reflected in the current assessment and management practices for the Benguela Current Large Marine Ecosystem. In this study, we compiled data from multiple demersal trawl surveys from the entire distribution area and applied state-of-the-art geostatistical population modelling (GeoPop) to estimate growth rate, mortality, and spatial and temporal distribution patterns of M. paradoxus. The data and the model enabled us to follow temporal and spatial changes in the distribution and infer movements from the recruitment/nursery areas, through the juvenile phase and the adults’ migration to the spawning areas outside/upstream of the nursery areas. The results indicated one primary recruitment/nursery area on the west coast of South Africa and a secondary less-productive recruitment/nursery area on the south coast near Port Elizabeth. Juveniles initially migrated away from the main recruitment area, followed by natal homing by larger individuals. This pattern was highly consistent through the time-series of the study. This perception of a, primarily, panmictic population that performs transboundary migrations between Namibia and South Africa corresponds largely to the hypothesis and data plots given in recent studies. We recommend that fisheries assessment, advice and management take into consideration these aspects of the distribution and population (stock) structure of M. paradoxus.  相似文献   

5.
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   

6.
A brief history of research into, and the fishery for, Cape hake is given. From twelve monthly surveys in 1972/73, abundance of the two species (Merluccius capensis and M. paradoxus) at depth was found to vary, larger individuals of both species being found deeper than the smaller ones. Off the Cape of Good Hope, M. capensis was never found deeper than 440 m, and there was significant species overlap between 150 and 440 m. M. paradoxus was numerically dominant except at the shallowest station sampled (150 m). Little overlap of mature specimens of each species was recorded. The research data were applied to commercial trawl statistics, revealing that M. capensis accounted for only 12,9 per cent by mass and 4,8 per cent by number of the commercially caught hake in the years 1955–1974. Such low percentages justify considering the Cape hake fishery as a single-species entity for the purposes of stock assessment and management.  相似文献   

7.
Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.  相似文献   

8.
Prevalence of infection with the myxozoan parasite Kudoa thyrsites is not well documented in sardine Sardinops sagax, the Cape hakes Merluccius capensis and M. paradoxus, and kingklip Genypterus capensis. Quantitative polymerase chain reaction assays were used to determine the prevalence of K. thyrsites in wild-caught S. sagax (n?=?296), Merluccius spp. (n?=?162) and G. capensis (n?=?70), harvested in 2015. Sardinops sagax were harvested from the east, south and west coasts of South Africa, whereas M. capensis, M. paradoxus and G. capensis were taken from the west coast only. Relationships between K. thyrsites infection prevalence and fish capture locality, season, sex and size were determined with Chi-square tests. Prevalence was highest in S. sagax (91%), followed by M. capensis and M. paradoxus (87%) and G. capensis (40%). Prevalence was independent of sex of the host for all fish species. In the case of S. sagax only, the prevalence of infection was independent of locality of capture but depended on season, with prevalence lowest during spring and highest during autumn. No significant relationship (p?>?0.05) was found between infection prevalence and host size for any of the fish species.  相似文献   

9.
Spawning time and areas, and the length of the spawning season of shallow-water (Merluccius capensis) and deep-water (M. paradoxus) hake, were investigated from bottom trawl collections taken in Namibian waters between September 1998 and October 2000 and from August to November 2001. The gonadosomatic index (GSI) and the incidence of advanced maturity stages of M. paradoxus were low along the entire coast throughout the year, but with a few exceptions in the south (between Lüderitz and Orange River). By contrast, M. capensis spawn in the area throughout the year, but mainly between July and October. Evaluating the accuracy of visual maturity staging by comparing results with those of image and histological analyses revealed few errors in classifications of maturity stage in the field, with the exception of Stage 5 (spent and resting) in M. paradoxus. Specific GSI values were intercalibrated with the appearance of developing oocytes. As the GSI is quickly estimated, this new concept, designated here as a 'maturity reference line', could enhance understanding of the spawning biology of other species with a similarly complex, indeterminate spawning strategy. It is concluded that M. paradoxus do not appear to spawn in Namibian waters.  相似文献   

10.
In this study demersal survey data for the period 1990–1999 are used to investigate the average distribution of the Cape hake Merluccius capensis and M. paradoxus off Namibia in terms of density and mean length. Further, biomass estimates are compared on the basis of depth and density stratification. The main distribution of M. capensis was north of 27°S and that of M. paradoxus south of 24°S. M. paradoxus was deeper than M. capensis. For both species, average length increased with depth. M. paradoxus expanded its range to the north through the 1990s as its population size (off South Africa and Namibia) increased. In Namibian waters, small M. paradoxus were found only south of 25°S. Mean length of M. capensis increased north of 21°S, largely as a result of decreased numbers of small fish in shallower water. Abundance estimates stratified by depth were no different from those post-stratified on similar densities.  相似文献   

11.
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   

12.
Spatial and size/age distributions of seven teleost species are described from a series of five annual trawling surveys made over the continental shelf and upper slope (20–500 m) of the South African south coast between Cape Agulhas and Port Alfred. The catch was quantified and scaled up to an estimate of an annual biomass index for each of the principal species. The fish studied were the two species of Cape hake Merluccius capensis and M. paradoxus, kingklip Genypterus capensis, Agulhas sole Austroglossus pectoralis, Cape horse mackerel Trachurus trachurus capensis, monk Lophius vomerinus and panga Pterogymnus laniarius. Each showed varying patterns of biomass fluctuation annually, although the surveys constituted too short a data series to reveal significant trends. Nevertheless, declines in abundance of panga and kingklip supported data collected from other sources over a longer period. Interpretation of the data is tempered by a realization that species which are patchily distributed or less vulnerable to the sampling gear may have been inadequately sampled. The information can be used for more-effective management of the multi-species demersal trawl fishery of the region.  相似文献   

13.
Information on hake stomach contents collected during research cruises off the west coast of South Africa between 1988 and 1990 is analysed. Estimates of the annual consumption and daily ration of the Cape hakes Merluccius capensis and M. paradoxus are obtained. Coefficients of variation (CVs) for these estimates are estimated by a bootstrap variance-estimation procedure. Cape hakes are opportunistic feeders. Juveniles feed mainly on crustaceans and the diet becomes increasingly piscivorous with age. Anchovy were the dominant dietary item of juvenile M. capensis, although this may be a reflection of the fact that anchovy were readily available at the time of the surveys. The large number of anchovy found in the diet during this study highlights the dangers of extrapolating consumption estimates for opportunistic feeders to other time periods. Pooling of data across geographic and seasonal strata introduces substantial bias in estimates of consumption and daily ration for only the pelagic prey species. Estimates of annual consumption are highly sensitive to the cruise selected to provide the estimates of numbers-at-length, because these latter estimates vary substantially between cruises. Furthermore, estimates of annual consumption and daily ration by prey species are very imprecise due to the effects of small sample sizes, the opportunistic nature of and natural variability associated with feeding. This implies that it is unlikely that model-estimation procedures which utilize these data will be able to provide particularly precise predictions. Therefore, before any larger-scale stomach collection exercise is undertaken, it is advisable to perform simulation studies to assess the sampling intensity required to achieve the desired levels of accuracy and precision for predictions from multispecies model-estimation procedures which make use of such data.  相似文献   

14.
Research surveys of Cape horse mackerel Trachurus trachurus capensis abundance on the south coast of South Africa are complicated because changes in the species' vertical and horizontal distribution limit the value of stock assessments based a single survey method. Annual bottom trawl surveys conducted in spring provide estimates of the abundance of fish close to the bottom over trawlable grounds. Between 1991 and 1994, hydroacoustic surveys conducted in spring have been used to estimate the pelagic portion of the stock, as well as the portion over untrawlable grounds. These two research datasets, as well as data from purse-seine, mid water and bottom trawl commercial landings, are reviewed to elucidate distribution patterns of horse mackerel and their migratory and spawning strategies. The problems and advantages of bottom trawl and acoustic surveys are discussed in the context of fluctuations in estimates of the size of the stock between 1991 and 1994 and the prevailing environmental conditions. It is concluded that combined acoustic and bottom trawl surveys are the only effective means of surveying horse mackerel, and that effort should be concentrated east of 22°E to assess the spawner stock. It is suggested that research effort directed at improving understanding of exchanges between West Coast (including Namibia) and South Coast population of horse mackerel, as well as of the role of vertical migrations in modulating these exchanges, would be beneficial.  相似文献   

15.
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   

16.
During intensive biannual demersal fish-sampling cruises since January 1983 over the South African west coast shelf region (Orange River to Cape Agulhas, coast to 500-m isobath), feeding studies on both species of Cape hake (Merluccius capensis and M. paradoxus) were undertaken. These studies were of two forms, random sampling of the dietary preference and constituents of the hake caught during daylight only and intensive sampling of stomachs with food throughout a 28-hour cycle. From the index of relative importance, crustaceans were shown to be most important in the majority of length classes of hake studied, with mesopelagics (fish and cephalopods) and other hake also important. Differences by hake species, by predator size, by geographical location and by season were evident. The Cape hakes, which constitute some three-quarters of the demersal fish biomass in the area, are deduced to play a major role in the marine faunal environment, both as predators and prey, and some base-line figures of the quantities of different food items consumed by the resource are presented. Hake are largely opportunistic feeders and it is concluded that they can adapt to perturbations in the availability of prey, whether they be triggered by environmental cause or by, for example, purse-seining of particularly mesopelagic fish.  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.
Indicators of abundance for American lobster (Homarus americanus) based on 8 years of trap catch rates (catch‐per‐unit‐effort, CPUE) were evaluated. Volunteer harvesters recorded count, sex and size of lobsters captured in standard traps on a daily basis during the fishing season in coastal Nova Scotia, Canada. We examined the extent to which standardised CPUEs of prerecruits predict the future catches of legal sizes and explored spatial patterns in the abundance of lobsters of different size and reproductive status. The standardised CPUE of prerecruits was correlated with legal size catches in only one of five areas examined. This area had a strong signal of incoming recruitment. Improving the capacity of prerecruit CPUE for predicting legal size catches several years later most likely lies with model incorporation of variables associated with catchability. The spatial distribution of catch rates showed that the area with the highest historical landings per unit area also had the highest relative abundance of prerecruits. The spatial distribution data point to further areas of research related to recruitment processes in lobster in coastal Nova Scotia.  相似文献   

19.
An eight-year research angling dataset collected between February 1998 and December 2005 in the Tsitsikamma National Park marine protected area (MPA), along the south-eastern Cape coast of South Africa, was examined to provide estimates of important fishery and population parameters for eight important shore-angling species. Five different estimates of natural mortality (M), and the coefficients of variation (CVs) of catch per unit effort (CPUE) and size measurements were calculated. There was no consistency in estimates of M between methods, highlighting the difficulty in obtaining reliable estimates of this parameter for per-recruit assessments. The CVs of CPUE measurements were generally higher than those for size measurements. Estimates obtained for the most abundant species, blacktail Diplodus capensis, with a mean CPUE of 0.252 fish h–1, yielded the lowest CVs for CPUE (0.806) and size (0.130) measurements. Therefore, blacktail is considered the preferred indicator species for making comparisons with estimates obtained for shore-angling fish in exploited areas and other MPAs along the South African coastline. The large inter annual variations in mean CPUE and size for all species indicated that a minimum assessment period of four years is required to obtain reliable estimates.  相似文献   

20.
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   

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