Vertical migration,catchability and acoustic assessment of semi-pelagic Cape horse mackerel Trachurus trachurus capensis in the southern Benguela

The vertical distribution and migration of the Cape horse mackerel Trachurus trachurus capensis were investigated on the basis of midwater and bottom trawl collections taken over diel cycles at two sites on South Africa's south coast. Attempts were also made to investigate the relationship between bottom trawl catch rates and acoustic abundance estimates. During the first cycle, horse mackerel were dispersed in midwater during the night, and towards sunrise they coalesced into schools and migrated to the seabed where they remained throughout the day. During that cycle, horse mackerel did not appear to feed, despite the prevalence of potential food, mainly Calanus copepods and euphausiid larvae, in the water column. In contrast, during the second cycle, horse mackerel remained near the seabed during the 24-h experiment, but they exhibited a marked feeding periodicity, with most feeding taking place in the late afternoon. They appeared to select for larger copepods in the water column. Several reasons are advanced to explain the difference in the vertical migratory behaviour of horse mackerel between the two sites, on the basis of the vertical distribution of their potential prey. Diel differences in acoustic estimates of horse mackerel abundance were significantly reduced after applying an acoustic deadzone correction factor. Trawl catches, however, could not be reconciled with acoustic estimates during the day, when horse mackerel were too close to the bottom for effective echo-integration.     

Reproductive biology of horse mackerel Trachurus capensis and Trachurus trecae in Namibia and Angola

Harvest control measures for Namibia and Angola are defined using reference points for spawning biomass, which requires knowledge of spawning potential and dynamics of fishery stocks. The reproductive biology of two species of horse mackerel, Trachurus capensis and Trachurus trecae, in these waters was investigated for spawning distribution in time and space. Samples were collected during two scientific cruises conducted in February and July/August 2007 off the coast of Namibia and Angola. A total of 396 fish ovaries was examined to determine the spawning dynamics of the two species. The ovaries were histologically analysed and their oocyte distributions mapped. The length (TL)–weight relationships of the two species differed significantly. The observed development of oocyte recruitment and absence of a hiatus in the oocyte distribution indicated that these species are indeterminate spawners. Analyses of the data from the NatMIRC database (2001–2004) indicated annual changes in gonadosomatic index (GSI) for T. capensis at different latitudes, but these changes were small. GSI also fluctuated between years and areas, but there were no clear spatial or temporal trends, with averages ≤4.0% at all stations. The GSI of both species was not indicative of the histologically determined development of gonad maturity. Oocyte maturation for T. capensis indicated that smaller and younger fish spawn first. Batch fecundity (F) for T. trecae was best described by F = 4 986 TL (cm) – 99 645 (r ² = 0.83). This study provided the first analyses of oocyte development in time and space for the two species in the region.     

Growth of Cape horse mackerel Trachurus trachurus capensis off South Africa

Recent studies have provided evidence that the Cape horse mackerel Trachurus trachwus capensis in South African waters is a unit stock. Therefore, an age and growth study of the entire stock throughout its distributional range was undertaken in 1988/89. Growth is described by the equation

L_t = 487,989 (1 – e^{?0.556(t – 0,229)}) mm.     

Variable spatial structure of schooling pelagic fish off Namibia: implications for acoustic surveys

Schools of horse mackerel Trachurus trachurus capensis, sardine Sardinops sagax and round herring Etrumeus whiteheadi were surveyed by conventional echo-integration along systematic transects. The transects were surveyed twice, in opposite directions. Clupeoid biomass was concentrated in a few dense aggregations, whereas horse mackerel were less dense and distributed over larger areas. The influence of aggregation patterns of the different species on the precision of the acoustic estimates was analysed with respect to spatial variability and diurnal effects. Isotropic variograms computed from values of acoustic back-scattering strength showed little or no structure for all three species. Indicator variograms showed relatively large structures (>15 miles) for all three species at the lowest cut-off level, confirming the influence of high values on the variograms. The autocorrelation range decreased with an increase in cut-off value for horse mackerel, whereas variograms of sardine and round herring showed no structure at larger cut-off values. There were large variances in estimated biomass for sardine (77%) and round herring (90%) between surveys, whereas biomass estimates of horse mackerel were relatively similar between surveys.     

Distribution patterns,stock size and life-history strategies of Cape horse mackerel Trachurus trachurus capensis,based on bottom trawl and acoustic surveys

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.     

Comparison of horse mackerel length frequencies obtained from research vessels and commercial midwater trawlers: implications for biomass estimation

The validity of abundance estimates from hydroacoustic surveys relies, inter alia, on the ability of the fishing gear on the research vessel to sample non-selectively. This study compares the length frequencies of Cape horse mackerel Trachurus trachurus capensis taken in Namibian waters by the R.V. Welwitchia and commercial midwater trawlers. Results indicate that the length distributions of catches taken by the Welwitchia were significantly different from those from commercial trawlers, with a greater proportion of fish >25 cm being sampled by the latter. Also, the biomass estimated per length-class from hydroacoustic surveys over the period 1994–2000 was compared with those from annual landings of the commercial fishery. The length distribution of horse mackerel in purse-seine catches compares favourably with those from the research vessel. However, comparisons with the midwater trawl catches indicated that the length frequencies obtained from research vessels during the years 1994–1997 underestimated the number of large fish in the population and biased the biomass in those years. From 1998 to 2000 the bias was negligible. The magnitude of the error varied between years, but it remained fairly low. To compensate for this bias, length distribution data from midwater trawlers should be integrated into the acoustic biomass calculation procedure.     

Stock assessment and management implications of horse mackerel (Trachurus japonicus) in Korean waters, based on the relationships between recruitment and the ocean environment

This study presents an example of horse mackerel (Trachurus japonicus) stock to demonstrate that marine environmental factors are important in stock assessment for the new Korean Total Allowable Catch (TAC)-based fisheries management system. The estimated survival rate (S) of horse mackerel ranged from 0.25 to 0.36. The instantaneous coefficient of natural mortality (M) was 0.48/year, and the age at first capture was 0.83 year. Annual biomass of horse mackerel in Korean waters was estimated by a biomass-based cohort analysis using annual catch in weight at age during 1965–1995. Yield-per-recruit and spawning biomass-per-recruit were estimated under various harvest strategies at F_max, F_0.1, F_30% and F_40%. A method for estimating acceptable biological catch (ABC) is proposed for dealing with the large differences in the quality and quantity of information and data available. Using recruitment of horse mackerel estimated from various spawner–recruitment relationship models combined with salinity, volume transport, and zooplankton biomass as environmental factors, the ABC under the best information available was estimated to range from 3100 to 3800 mt.     

Food of predatory demersal fish in Hauraki Gulf

Feeding was examined in five species of fish associated with snapper Chrysophrys auratus (Forster) trawled from Hauraki Gulf and north‐west Bay of Plenty: trevally Caranx lutescens Richardson, red gurnard Chelidonichthys kumu Lesson and Garnot, horse mackerel Trachurus novaezelandiae Richardson, eagle ray Holorhinus tenuicaudatus (Hector), and john dory Zeus japnnicus Cuvier and Valenciennes. All five species feed on some elements of the bottom fauna; in general, crustaceans were the most important food group.     

Analysis of bycatch in the South African midwater trawl fishery for horse mackerel Trachurus capensis based on observer data

The South African midwater trawl fishery targets adult horse mackerel Trachurus capensis. The bulk of the catch is taken by a single freezer-trawler, the biggest fishing vessel operating in South African waters. As fishing takes place off the south coast in ecologically sensitive areas, there are concerns about the potential impacts of this fishing operation on non-target species. Fishing behaviour and bycatch of this fishery from 2004 to 2014 were investigated by analysing observer records with regard to catch composition, volume and temporal and spatial patterns. The midwater trawl fishery was estimated to have caught 25 415 tonnes annually, with a bycatch of 6.9% of the total catch, by weight. There are species overlaps with various fisheries, namely the demersal trawl, small-pelagic, line, shark longline and squid fisheries, yet the total bycatch estimates from this fishery are generally small relative to catches taken in the target fisheries. Bycatch species with the highest average annual catches were chub mackerel Scomber japonicus, redeye roundherring Etrumeus whiteheadi, ribbonfish Lepidopus caudatus and hake Merluccius spp. Large-fauna bycatch species included sunfish Mola mola as well as a number of CITES II- and IUCN-listed species, such as Cape fur seal Arctocephalus pusillus, dusky shark Carcharhinus obscurus, smooth hammerhead shark Sphyrna zygaena and thresher sharks Alopias spp. The 97.9% observer coverage is high and the 6.9% bycatch rate low compared to other South African fisheries; however, due to the large size of the individual hauls (average of 46.3 t), the average sampling rate of 1.56% is low. Our analyses suggest that bycatch in the South African midwater trawl fishery has been lower than in other South African fisheries and similar fisheries elsewhere, but due to the combination of high catch volumes and low sampling rates, estimation errors for rare species are high and there is a substantial risk of incidental unmonitored bycatch of rare large fauna and aggregations of small fauna. This could be mitigated by spatio-temporal management of this fishery, to avoid fishing in high-risk areas, and the introduction of an electronic monitoring programme.     

Horse mackerel egg identification using DNA methodology

Egg identification in plankton samples is usually needed for purposes of stock assessment. Until recently, only morphological characters were used for identifying the eggs of different fish species. Late developmental stages are easily distinguishable due to pigmentation as well as egg and oil globule size range. However, for early stages, these characteristics are difficult to be discriminated and may overlap with other species. European horse mackerel species (Trachurus trachurus, T. mediterraneus, T. picturatus) overlap significantly in their spawning areas in some European waters. Because of the fact that their eggs are morphologically similar, genetic methodologies are needed to identify eggs and larvae to species correctly. In the present study, formalin‐ and ethanol‐preserved eggs were tested to estimate the efficacy of genetic methodologies for species identification of eggs when different preservatives are used. A 370‐bp fragment of cytochrome b was successfully amplified followed by restriction fragment analysis with two restriction enzymes aiming to identify the eggs to species. Horse mackerel egg identification was accomplished with the maximum success in ethanol‐preserved eggs. However, it seems that various preservatives had different effects on the DNA quality of eggs as genetic identification was less successful in formalin‐preserved eggs. Preserving in ethanol a part of the eggs obtained in plankton surveys is suggested for purposes of accurate genetic identification, even if their morphological features are distorted in time.     

Identification of larvae of snapper Chrysophrys auratus forster by electrophoretic separation of tissue enzymes (note)

A comparison was made between the electrophoretic patterns of isozymes of esterases and glucosephosphate isomerases of adult snapper Chryosophrys auratus, suspected snapper larvae, jack mackerel Trachurus novaezelandiae larvae, and blue mackerel Scomber australasicus larvae. The suspected snapper larvae revealed similar isozyme patterns to those of the adults, but differed from the other two larval species.     

The distribution and abundance of seven commercial trawlfish from the Cape south coast of South Africa, 1986–1990

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.     

Size composition of Jack mackerel catches from the west coast of New Zealand

Catches of iack mackerel Trachurus declivis taken everv 2 months from the west coast of New Zealand show that this species attains lengths of approximately 11 cm and 24 cm at the completion of the first and second years of life respectively. Above these lengths, year classes overlap considerably. Juveniles were found from Westland to North Cape and were most abundant in surface waters.     

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.     

Biochemical‐genetic variation in a wild and a cultured population of the greenshell mussel,Perna canaliculus

A wild and a cultured greenshell mussel (Perna canaliculus) population were compared for biochemical genetic variation at seven polymorphic and four monomorphic allozyme loci. Significant heterozygote deficiencies were observed for all polymorphic loci (La—1, La—2, Lap, Lgg—1, Lgg— 2, Pgi, and Pgm) for both populations (except the Pgi locus of the wild mussel population). Genotypic disequilibrium was calculated for both populations: genotypic frequencies were significantly non‐random at three pairs of loci among the wild mussels, and significantly non‐random at three different pairs of loci among the cultured mussels. All six pairs of loci which exhibited significant genotypic disequilibrium involved amino‐peptidases, suggesting that these loci form a linkage group, and that neither the Pgi nor the Pgm loci are associated with this group. Exact tests for population differentiation based upon population‐specific allele distributions indicated that four of the polymorphic loci were significantly heterogeneous among the two populations, whereas the remaining three polymorphic loci were not. Based upon the private allele system, the number of migrants (N _m) between the populations was estimated to be 2.009, which, according to the private allele system, represents a high level of gene flow. These findings are discussed with regard to the population biology and genetic structure of this species.     

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.     

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.     

Age,growth and natural mortality of blue jack mackerel Trachurus picturatus (Carangidae) from the Canary Islands,Spain (NW Africa)

Status assessment of exploited fish populations is a general requirement to ensure sustainability of fisheries in both global and local contexts. The blue jack mackerel Trachurus picturatus (Bowdich, 1825) remains unassessed in a data-limited fishery in the eastern central Atlantic, within the waters of the Canary Islands (Spain) off the coast of northwest Africa. This is the most southwestern targeted fishery in the species’ distributional range. To obtain basic life-history information, a total of 2 472 specimens were analysed, ranging from 10.2 to 31.9 cm total length, from samples collected monthly between March 2005 and March 2006. Length–weight relationships (a = 0.0023; b = 3.418; R ² = 0.97) showed no significant difference between sexes. Deposition periodicity and coherence of the ageing criteria were verified by analysis of the monthly evolution of the otolith edge. Ages were estimated by counting annual growth rings on whole otoliths, with these fish ranging from age 0 to 6 years. The von Bertalanffy growth model, which was used to estimate growth parameters from both direct readings and back-calculated length-at-age data, gave good statistical fits. The parameters from the direct readings were: L∞ = 34.9 cm; k = 0.214 y^–1; t ₀ = ?2.545 y (R ² = 0.84), and the natural mortality rates estimated from empirical methods varied between 0.20 and 0.55 y^–1.     

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.