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.     

Factors influencing growth of the African penguin colony at Boulders,South Africa, 1985–1999

This paper reports on growth of the Boulders colony of African penguins Spheniscus demersus from inception in 1985 to the present. More than 900 pairs now breed there. Growth of the colony slowed in 1995 and 1996 and reversed in 1998, coinciding with periods of low abundance of Cape anchovy Engraulis capensis off South Africa. In December 1996, penguins were excluded from a portion of land where they had formerly bred. They responded by increasing the density of their nests in other areas and expanding their area of breeding longshore. These patterns indicate that food and not space are currently controlling colony growth rate. Much of the colony growth probably results from immigration of first-time breeders from other colonies. Of immigrants, 70–80% may be from Dyer Island to the south-east, where numbers of penguins have decreased. Boulders also is frequently visited by penguins from other colonies, and by rehabilitated birds.     

Remotely sensed variability of temperature and chlorophyll in the southern Benguela: upwelling frequency and phytoplankton response

High-resolution (1km) satellite data from the NOAA AVHRR (Advanced Very High Resolution Radiometer) and OrbView-2 SeaWiFS (Sea-viewing Wide Field-of-view Sensor) are used to investigate the upper layer dynamics of the southern Benguela ecosystem in more detailed space and time scales than previously undertaken. A consistent time-series of daily sea surface temperature (SST) and chlorophyll a concentration images is generated for the period July 1998–June 2003, and a quantitative analysis undertaken. The variability in SST, upwelling and phytoplankton biomass is explored for selected biogeographic regions, with particular focus on intra-seasonal time scales. The location and emergence of upwelling cells are clearly identified along the length of the southern Benguela, being distinct on the narrow inner and the mid-continental shelves. Most notable is the rapidly pulsating nature of the upwelling, with intense warm/cold events clearly distinguished. The phytoplankton response to this physical forcing is described. Chlorophyll concentration on the inner shelf largely mirrors the pattern of SST variability, similarly dominated by event-scale processes. Over the mid-shelf, higher chlorophyll is observed throughout all seasons, although low biomass occurs during winter. The variability of the offshore extent of SST and chlorophyll is identified at locations of differing shelf width. Cooler upwelled water is confined primarily to the narrow inner-shelf, with event-scale pulses extending considerable distances offshore. Agulhas Current influences are readily observed, even on the Cape Peninsula inner-shelf. Chlorophyll concentrations vary considerably between the locations of differing shelf width. SST, upwelling and phytoplankton indices are derived for selected locations to quantify the intra-seasonal variations. The SST indices show marked temperature changes associated with rapid pulsation on the event scale. No strong seasonal signal is evident. In contrast, the upwelling indices display a strong seasonal signal, with most intense upwelling occurring in spring/summer in the south. The phytoplankton response to the seasonal upwelling index differs between the selected locations. This study concludes that, although low-resolution SST and chlorophyll data may be useful for investigating general patterns over large scales, higher resolution data are necessary to identify finer scale spatial and temporal variability, especially in the inshore coastal zones.     

Comparative ecology of the copepods Calanoides carinatus and Calanus agulhensis — the influence of temperature and food

Hypotheses regarding temperature, food abundance and food size were tested to explore niche separation between Calanoides carinatus, an abundant copepod in the cool and food-rich southern Benguela upwelling system, and Calanus agulhensis, the dominant copepod on the warmer, relatively food-poor Agulhas Bank off the south coast of South Africa. Under non-limiting food conditions, egg production by both species increased linearly with temperatures between 9°C and 18°C. Egg production by C. carinatus was relatively faster at 21°C, but was offset by greater mortality. Both species showed similar functional responses to food concentration in the field, reaching satiation at ～15mg Chl a m^?3, or ～3–4ppm. Food abundance was the most important predictor of egg production, whether measured as Chl a or as particle volume. Both species preferred larger particles that dominated the biomass peak, but particle size appeared more important for C. carinatus, with increasingly faster rates of egg production as the proportion of large cells (>10μm) exceeded 50%, and slower ingestion of small (<10μm) cells. Omnivory may be more important to C. agulhensis. Niche separation between the two species appears unrelated to temperature, food abundance or diel vertical migratory behaviour, and is more likely a function of variability in food availability, although food size may also play a role.     

Application of the sequential t-test algorithm for analysing regime shifts to the southern Benguela ecosystem

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

Abrupt environmental shift associated with changes in the distribution of Cape anchovy Engraulis encrasicolus spawners in the southern Benguela

Cape anchovy Engraulis encrasicolus spawners in the southern Benguela showed an eastward shift in their distribution on the Agulhas Bank that occurred abruptly in 1996 and has since persisted. We assessed whether this shift was environmentally mediated by examining sea surface temperature data from different regions of the Agulhas Bank, which showed that in 1996 the inner shelf of the Agulhas Bank to the east of Cape Agulhas abruptly became 0.5°C colder than in previous years and has since remained that way. In addition, signals, coherent with the 1996 shift recorded in sea surface temperatures, were also found in atmospheric surface pressure and zonal wind data for that region; interannual coastal SST variability is also shown to be correlated with zonal wind-stress forcing. As a result, increased wind-induced coastal upwelling east of Cape Agulhas is proposed as the main driver of the observed cooling in the coastal region. The synchrony between the environmental and biological signals suggests that the eastward shift in anchovy spawner distribution was environmentally mediated and arose from a change in environmental forcing that altered the relative favourability for spawning between regions to the west and east of Cape Agulhas. The results highlight how a relatively minor change in environmental conditions can lead to a drastic spatial reorganisation of the life history of one species in an ecosystem.     

Influence of buoyancy and vertical distribution of sardine Sardinops sagax eggs and larvae on their transport in the northern Benguela ecosystem

In recent years, sardine Sardinops sagax spawning has been recorded inshore off central Namibia. Field observations on eggs and laboratory measurements show that spawning, demonstrated by the distribution of newly spawned eggs, takes place just below the upper mixed layer. The high positive buoyancy of the eggs causes them to ascend rapidly to the surface layer, where they are moved offshore by upwelling-induced offshore transport. However, increased wind-induced mixing also influences the vertical distribution of eggs, causing them to be partly mixed down below the layer moving offshore and into the layer moving inshore. This mechanism acts to retard the transport and offshore loss of eggs from the spawning areas. The vertical distribution of sardine larvae, with highest concentrations deeper than 20 m, indicates active movement out of the layer moving offshore, and this tendency seems to be more pronounced for older larvae. Hence, vertical migration of larvae is an additional factor mitigating their loss from nearshore. Taken together, these features seem to minimize the offshore loss of offspring, particularly in periods of low stock biomass when spawning close to the shore seems to be common.     

A comparison of condition factor and gonadosomatic index of sardine Sardinops sagax stocks in the northern and southern Benguela upwelling ecosystems, 1984–1999

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.     

A method of age determination for the South West African pilchard Sardinops ocellata

A method of age determination, based on recent research on growth and ring formation, is described for the pilchard of South West Africa. Dates of ring formation from 1971 to 1981 are documented. Illustrated examples of otolith interpretations are given.     

Genetic and morphological variation of the lanternfish Lampanyctodes hectoris (Myctophiformes: Myctophidae) off southern Africa

Genetic and morphological variation within the southern African population of Lampanyctodes hectoris was analysed. A total of 15 enzymes, encoding 22 isozyme loci, was examined (n = 327); seven were polymorphic. The percentage of polymorphic loci ranged from 13.6 to 27.3%. The mean heterozygosity was generally low and ranged from 0.003 to 0.005. Genetic divergences between specimens from four areas along the west coast of southern Africa were negligible, genetic distance values (D) ranging from 0 to 0.00011. The results showed the population of L. hectoris to be genetically invariant. Principal Component Analysis was performed separately on ratios of 13 morphometric and 6 meristic variables (n = 446), and there was evidence of extensive overlap between fish from all areas. However, discriminant analysis suggested some morphological variability within this population. Although genetically the population of L. hectoris seems to be homogenous, morphologically it appears to be variable. A combination of these results suggests that there is no clear genetic basis for the slight morphological differentiation within the population.