Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

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.     

Depth-stratified density estimates and distribution of the Cape hake Merluccius capensis and M. paradoxus off Namibia deduced from survey data, 1990–1999

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.     

Diel effects on bottom-trawl survey catch rates of shallow- and deep-water Cape hakes Merluccius capensis and M. paradoxus off Namibia,using solar zenith angle

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.     

Suffocation of pilchards (Sardinops sagax) by a green microalgal bloom in Wellington Harbour,New Zealand

A bloom of the green microalga Tetraselmis sp. in a small lagoon in Wellington Harbour, New Zealand, is considered to have caused a fish kill of pilchards (Sardinops sagax). The fish deaths are attributed to anoxia brought about by a combination of reduced dissolved oxygen levels and the microalgae sticking to and clogging the secondary gill lamellae of the fish.     

Population abundance and seasonal migration patterns indicated by commercial catch-per-unit-effort of hakes (Merluccius capensis and M. paradoxus) in the northern Benguela Current Large Marine Ecosystem

We developed generalised additive models (GAMs) to estimate standardised time-series of population abundance indices for assessment purposes and to infer ecological and behavioural information on northern Benguela hakes, Merluccius capensis and M. paradoxus, using haul-by-haul commercial trawl catch-rate data as proxies for hake densities. The modelling indicated that individual ship identifiers should be used rather than general vessel characteristics, such as vessel size. The final models explained 79% and 68% of the variability in the commercial catch rates of M. capensis and M. paradoxus, respectively. The spatial density patterns were consistent and confirmed existing knowledge about these species in the northern Benguela system. Furthermore, seasonal migration patterns were described for the first time and were found to correspond to the known spawning areas and seasons for M. capensis and M. paradoxus. Spatial density patterns were validated using the geostatistical modelling results of fisheries-independent trawl survey data. Improved understanding of the relationships between fleet dynamics and fish movement can be achieved by taking into consideration the present catch-rate model and spatial and seasonal distribution maps. We conclude that the yearly standardised CPUE time-series are problematic as proxies for total stock abundance because of spatial coverage issues. Consequently, such CPUE data should not be used for stock-size assessments and fisheries advice concerning northern Benguela hakes until this is solved. We generally recommend the exclusion of standardised CPUE time-series from stock assessments when important and changing parts of the stock distribution cannot be targeted by the fishery, such as due to closed areas or seasons.     

Occurrence and distribution of Cape hakes Merluccius capensis Cast. and M. paradoxus Franca in the Cape of Good Hope area

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.     

Characterisation of the dietary relationships of two sympatric hake species,Merluccius capensis and M. paradoxus,in the northern Benguela region using fatty acid profiles

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.     

An overview of the living marine resources of Namibia

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.     

Diet of the Cape fur seal Arctocephalus pusillus pusillus at the Robberg Peninsula,Plettenberg Bay,and implications for local fisheries

Cape fur seals Arctocephalus pusillus pusillus were harvested to extirpation on the Robberg Peninsula, Plettenberg Bay, on the south-east coast of South Africa, between the 17th and early 20th centuries. Seals returned to Robberg in small numbers during the early 1990s and their numbers subsequently increased. We studied the diet of this increasing population using faecal (scat) sampling to determine: the species composition and size of prey in the diet of Cape fur seals at Robberg; to explore temporal variation in the diet; and to investigate the potential for competition between seals and the fisheries around Plettenberg Bay. Of the 445 scats collected, 90% contained hard prey remains and 15 teleost prey species were represented in the 3 127 otoliths that could be identified. The seals’ most important prey species in terms of numerical abundance, frequency of occurrence and mass in the diet, were anchovy Engraulis encrasicolus, sardine Sardinops sagax, horse mackerel Trachurus capensis, sand tongue-fish Cynoglossus capensis and shallow-water hake Merluccius capensis (in decreasing order of importance for numerical abundance). The proportion of anchovy in the diet increased during the study period (2003–2008), whereas the proportion of sardine decreased. The estimated average annual consumption of sardine by seals was higher than the average annual catch made by purse-seine fisheries in this area, suggesting resource competition between seals and purse-seiners, especially in the light of continuing growth of seal numbers in the area. However, direct competition between seals and linefisheries appeared to be minimal. Scat sampling of Cape fur seals holds potential to serve a useful and cost effective indicator of temporal changes in sardine abundance.     

Major endoparasites of the Cape hakes Merluccius capensis and M. paradoxus,with brief notes on some conspicuous ectoparasites

The major endoparasites of Cape hake are larvae of the nematode Anisakis sp. and the trypanorhynch cestode Hepatoxvlon trichiuri. Incidence and degree of infestation by Anisakis sp. were high and similar in both species, and the parasite is one of the most important natural enemies of hake. H. trichiuri was not as common or apparently pathologically destructive as Anisakis and infected Merluccius capensis more frequently and intensively than M. paradoxus. Other parasites of Cape hake, both internal and external, are listed.     

Identification of and distinction between the scales of South African pilchard Sardinops ocellatus and Cape anchovy Engraulis capensis

Fossil fish scales hold potential for elucidating past fluctuations in fish populations. Only certain pilchard Sardinops ocellatus and anchovy Engraulis capensis scales can be distinguished easily from each other. Misidentification of the less-typical scales of the two species can introduce a bias in scale-based population studies and, in order to overcome this bias, scales removed from pilchard and anchovy reared in captivity were examined. A complete set of reference photographs of the scales of each species was compiled and study of these showed that both pilchard and anchovy have five distinctive scale types, each of which is found on different parts of the fish. It was also shown that the "typical" scales of each species, i.e. those referred to in the literature, constitute less than 50 per cent of the scales found on the fish.     

Testing for the occurrence of pilchard herpesvirus (PHV) in South African sardine Sardinops sagax

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.     

A note on the diet of Cape cormorant Phalacrocorax capensis at Mercury Island,South West Africa,in November 1978

During November 1978, Cape cormorants Phalacrocorax capensis at Mercury Island, off South West Africa (Namibia), consumed practically exclusively pelagic gobies Sufflogobius bibarbalus. The mean number of fish per regurgitation was 52 with a mean caudal length of 48 mm and an estimated mean mass of 1.7 g. The estimated mean original mass of stomach contents was 75 g. These Cape cormorants took significantly smaller pelagic gobies than did bank cormorants P. neglectus sampled concurrently at the same locality.     

Intraspecific cleaning by juvenile Cape white seabream Diplodus capensis (Sparidae) off eastern South Africa

Juveniles of the Cape white seabream Diplodus capensis were observed cleaning adult conspecifics in a large tidepool off Sodwana Bay, South Africa. Although nine other tropical fish species were present and interacted with a nearby pair of Labroides cleaner wrasses, only adults of D. capensis posed for and were cleaned by the D. capensis juveniles. Such cleaning interactions have not been reported previously for this species or among marine fishes off South Africa, and thus add to the growing list of facultative cleaners globally.     

Spatial variation in meristic and morphometric characteristics of sardine Sardinops sagax around the coast of southern Africa

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.     

Estimation of the annual consumption of food by Cape hake Merluccius capensis and M. paradoxus off the South African west coast

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.     

Merluccius capensis spawn in Namibian waters,but do M. paradoxus?

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.     

Validation of growth zone deposition rate in otoliths and scales of flathead mullet Mugil cephalus and freshwater mullet Myxus capensis from fish of known age

Flathead mullet Mugil cephalus and freshwater mullet Myxus capensis are important components in South African estuarine fish communities and fisheries, but there is little information on their age and growth or age validation. This study validated the periodicity of growth zone formation in sectioned sagittal otoliths and scales of fish of known age that had been at liberty for 10 years. For both M. cephalus and M. capensis, the number of growth zones counted on otolith sections did not differ significantly from the known age of the fish. There were significantly fewer growth zones on scales than the known age of both M. cephalus and M. capensis. Growth zone deposition rate could therefore be validated as annual for otoliths but not for scales of the two species. It is recommended that future ageing studies focus on otoliths rather than scales.     

Composition,abundance, distribution and seasonality of larval fishes in the shallow nearshore of the proposed Greater Addo Marine Reserve,Algoa Bay,South Africa

The larval fish assemblage was investigated in the shallow, nearshore region of a proposed marine protected area in eastern Algoa Bay, temperate South Africa, prior to proclamation. Sampling was conducted at six sites along two different depth contours at ∼5 m and ∼15 m to assess shore association. Larvae were collected by means of stepped oblique bongo net tows deployed off a ski-boat, twice per season for 2 years between 2005 and 2007. In total, 6045 larval fishes were collected representing 32 families and 78 species. The Gobiidae, Cynoglossidae, Clupeidae, Engraulidae and Sparidae were the dominant fish families. Catches varied significantly among seasons peaking in spring with a mean of ∼200 larvae/100 m³. Mean overall larval density was higher along the deeper contour, at ∼15 m (40 larvae/100 m³). The preflexion stage of development dominated catches at the ∼5 m (80%) and ∼15 m (73%) depth contours. Body lengths of Argyrosomus thorpei, Caffrogobius gilchristi, Diplodus capensis, Heteromycteris capensis and Solea turbynei, all estuary associated species, were larger at the shallow sites nearer to shore. Larvae of coastal species that produce benthic eggs dominated catches (75%) in the shallow sites (∼5 m) but were less abundant (32%) farther from shore at the deeper (∼15 m) sites. All developmental stages of D. capensis, Engraulis capensis, H. capensis, Sardinops sagax and two Pomadasys species were found in the study area. It appears that some species use the shallow nearshore as a nursery area.     

Influence of oceanographic fronts and low oxygen on the distribution of ichthyoplankton in the Benguela and southern Angola currents

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.