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.     

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.     

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.     

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.     

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.     

Prevalence of Kudoa thyrsites (Myxozoa) in South African sardine Sardinops sagax,kingklip Genypterus capensis,and Cape hakes Merluccius capensis and M. paradoxus

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.     

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.     

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.     

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.     

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.     

Bycatch in the gillnet and beach-seine fisheries in the Western Cape,South Africa,with implications for management

Interview questionnaires and access point surveys were conducted in order to describe and quantify the catch composition of the inshore net-fisheries in the Western Cape, South Africa. A total of 138 562 fish, representing 29 species from 20 families, was recorded in 141 monitored commercial gillnet fishing operations between February 1998 and October 1999. Numerically, the legal target species, harders Liza richardsonii, dominated the catches, contributing 94.87% of the total gillnet catch. Elf, Pomatomus saltatrix, horse mackerel Trachurustrachurus capensis, gurnard Chelidonichthys capensis and barbel Galeichthys feliceps were the most common bycatch species, and contributed 4.2% to the total catch numerically and occurred in 12–47% of the marine 44–64-mm gillnet catches that were monitored. Five species most frequently targeted by shore-anglers on the West Coast: galjoen Dichistius capensis, white stumpnose Rhabdosargus globiceps, hottentot Pachymetopon blochii, silver kob Argyrosomus inodorus and white steenbras Lithognathus lithognathus, also occurred in gillnet catches. Most of the bycatch consisting of immature, undersized fish that were often injured during entanglement and were not released alive. L. richardsonii also numerically dominated the beach-seine hauls that were monitored (>99%) with only four bycatch species being recorded in low numbers. Beach-seine questionnaire respondents, however, reported sporadic catches of at least 17 bycatch species, including occasional appreciable catches of the important linefish species L. lithognathus and A. inodorus.     

Evaluation of a class of possible simple interim management procedures for the Namibian hake fishery

During 1997, considerable scientific differences arose about the status of the Namibian hake (Merluccius capensis and M. paradoxus) resource, and as to whether the hake Total Allowable Catch (TAC) should be substantially decreased or increased. These differences revolved primarily around whether or not abundance estimates from the swept-area trawl surveys by the Norwegian research vessel Dr Fridtjof Nansen should be considered as reliable measures of biomass in absolute (as distinct from relative) terms. The paper relates the computations underlying the Interim Management Procedure (IMP) approach that was put forward at that time as a basis to resolve this impasse. The anticipated performance, in terms of catch and risk of resource depletion, of a number of simple candidate IMPs for the Namibian hake resource is evaluated. The IMPs depend on two parameters, whose values are to be chosen by decision-makers, and adjust the TAC up or down from one year to the next according to whether trends in recent commercial catch rate and survey indices of abundance are positive or negative. Performances are evaluated across the then current wide range argued for resource abundance and status. Trade-offs in performance across the candidates considered are discussed. One of the candidates was subsequently selected by a joint meeting of scientists, industry and Ministry officials in February 1998 and served as the basis for scientific recommendations for the TAC for the hake resource for the following three years.     

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

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

Comparison of the abundance of pilchard in Cape gannet diet and commercial catches off the Western Cape,South Africa

A hypothesis is presented that, in the absence of quota restrictions, commercial catches of pilchard Sardinops ocellatus in the western Cape and the abundance of pilchard in the diet of Cape gannet Morus capensis obtained at three South African colonies are directly related to pilchard biomass when the latter is low. There were significant correlations (p < 0,05) between the abundance of pilchard in gannet diet at Lambert's Bay and Malgas Island combined, and the total catch and CPUE at annual, though not monthly, time-scales. Trends in levels of correlation are discussed in relation to short-term variability, sampling intensity, statistical chance and the distribution of pilchard. Gannet diet and commercial catches reflect regional, but not local, changes in biomass. The 1983 pilchard year-class was relatively strong. Gannet diet is suggested to be a reliable monitor of the trend in pilchard biomass at low pilchard biomasses, a time at which direct surveys are least reliable.     

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.     

Seasonally and spatially referenced estimates of recreational shore-angling effort,catch composition,catch rates and total catch in the Goukamma Marine Protected Area,South Africa

A roving creel survey of the recreational shore fishery along the 16.4-km coastline in the Goukamma Marine Protected Area on the south coast of South Africa was conducted from 2009 to 2011. Some 838 patrols were stratified equally among months, areas and years, but intentionally biased towards weekends. Angler densities at Buffalo Bay and Groenvlei were 0.59 and 0.28 anglers km?1, respectively. Weekend densities were double to quadruple weekday densities and fishing during winter was more popular than during summer. Area, habitat and distance to access points explained variation in angler densities. Shannon–Wiener diversity in catches declined from 2.18 in an earlier (1993–2002) survey to 1.79. Although the order of species abundance in the catches remained largely unchanged, blacktail Diplodus capensis dominance increased to 57.3% by number, at the expense of galjoen Dichistius capensis. Habitat explained 27% of the variance in catch composition. The catch per unit effort (CPUE) for the top nine species ranged from 0.19 to 6.35 fish 100-h?1. The CPUE of all species, except spotted grunter Pomadasys commersonnii, declined. Blacktail and galjoen CPUE declined by 17% and 77%, respectively. The total catch estimate was 2 986 fish y?1. Transgressions of size limits were common. The results suggest that the fishery is overexploited and that catch rates are declining.     

Demersal communities off the west coast of South Africa

To describe demersal communities off the west coast of South Africa, collections of all species of fish, cephalopods and crustaceans made during four cruises in the area during January and July have been analysed. Hierarchical classification methods were employed to determine species and sample associations. The results showed that the species could be split into two main groups, the first containing Merluccius paradoxus (the dominant species both in abundance and frequency of occurrence) and other species normally found in deep (> 380 m) water. The second group is represented by species that live on the continental shelf, M. capensis being the most abundant. The boundary separating the deep-sea community from that of the shelf follows approximately the 385-m isobath. Seasonal variations in the number of main associations as well as in their spatial location are documented.     

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.     

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.     

闽东近海游泳动物群落组成、资源量与生物多样性的时空格局研究

采用底拖网方法调查研究了闽东近海春季(5月)和秋季(10月)游泳动物群落组成、生物多样性、资源密度及时空分布格局等问题。调查中出现游泳动物70种(鱼类47种,甲壳类18种,头足类5种),隶属于15目40科64属,以鲈形目种类最多(25种)。鱼类是主要游泳动物群落,春季鱼类个体数和重量分别占总渔获的66.4%和72.3%,甲壳类分别占比33.4%和26.9%,头足类分别占比0.2%和0.8%;秋季,上述三个类群在总渔获中的占比分别为56.6%和60.4%、42.0%和37.8%以及1.4%和1.8%。春季优势种包括红狼牙虎鱼(Odontamblyopus rubicundus)、三疣梭子蟹(Portunus trituberculatus)、孔虎鱼(Trypauchen vagina)等10种,秋季优势种为龙头鱼(Harpadon nehereus)、三疣梭子蟹、六指马鲅(Polynemus sextarius)等7种;两个季节共有优势种包括三疣梭子蟹、日本 (Charybdis japonica)、龙头鱼、棘头梅童鱼(Collichthys lucidus)和六丝钝尾虎鱼(Amblychaeturichthys hexanema)5种。秋季的生物多样性指数和均匀度指数高于春季。春季平均资源密度为1013.7kg/km~2和10.2×10~4ind/km~2,高于秋季的平均值(899.0kg/km~2和6.6×10~4ind/km~2)。总体而言,鱼类是闽东近海主要的游泳动物类群,其资源密度从近岸向外海呈降低趋势,春季的资源密度高值出现在闽东外海中部水域,秋季高值则出现在南部的闽江口等水域;其春季的优势种种类多于秋季,存在一定程度季节性更替现象。