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.     

A knowledge base for management of the capital-intensive fishery for small pelagic fish off South Africa

As a contribution to South Africa's move towards an ecosystem approach to fisheries management, this study explores the existence of common perceptions about South Africa's pelagic fishery between resource users and scientists. It represents a collaborative research effort of social and natural scientists. A brief overview is given of the southern Benguela upwelling ecosystem and small pelagic fish resources, the fishery, and management of the fishery. Stakeholder knowledge and views were determined by conducting open-ended qualitative local knowledge interviews. Candidate indicators to address five major issues raised in the interviews were selected: length-at-50% maturity, total mortality, exploitation rate, proportion of bycatch, mean length of catch, and centre of gravity of catches. The indicator approach is shown to be a useful tool to manage the South African small pelagic fishery, and can be made compatible with existing management approaches. The foundation of a good adaptive fisheries management system is a data collection system that enables multi-disciplinary analysis and provides a basis on which decisions can be made.     

Revised estimates of abundance of South African sardine and anchovy from acoustic surveys adjusting for echosounder saturation in earlier surveys and attenuation effects for sardine

Hydro-acoustic surveys have been used to provide annual estimates of May recruitment and November spawner biomass of the South African sardine Sardinops sagax and anchovy Engraulis encrasicolus resources since 1984. These time-series of abundance estimates form the backbone of the assessment of these resources, and consequently the management of the South African sardine and anchovy is critically dependent on them. Upgrades to survey equipment over time have resulted in recent surveys providing more accurate estimates of abundance, yet in order to maintain comparability across the full time-series, estimates of biomass mimicking the old equipment were used for a number of years. In this paper we develop a method to revise the earlier part of the time-series to correct for receiver saturation in the older generation SIMRAD EK400 and EKS-38 echo sounders and to account for attenuation in dense sardine schools. This is applied to provide a revised time-series of biomass estimates for the South African sardine and anchovy resources with associated variance–covariance matrices. Furthermore, the time-series presented here are based on updated acoustic target strength estimates, making this the most reliable time-series currently available for both resources.     

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.     

Feeding ecology of South African Argyrosomus japonicus (Pisces: Sciaenidae), with emphasis on the Eastern Cape surf zone

A Monte-Carlo simulation approach is used to investigate the relative robustness of the results of the Butterworth–Andrew (B ₁ = K and B ₁ estimated) dynamic production-model observation-error estimators to different "true" values of the ratio of initial biomass B ₁ to unexploited equilibrium biomass K. The simulations use an underlying operating model typical of that appropriate for stocks of Cape hake off southern Africa. The B ₁ = K variant of the estimator results in lower estimated expected discrepancies for a number of management quantities over a wide range of the value of the "true" ratio. In addition, it is less likely to provide severe overestimates of the f _0.1 total allowable catch (TAC). Not all management quantities are equally sensitive to the value of the "true" ratio. In particular, f _0.1) TACs can be severely overestimated, although the maximum sustainable yield (MSY) is relatively well determined over a wide range of values of the ratio. It is therefore recommended that harvesting strategies which do not permit TACs to be set in excess of the estimated MSY be preferred.     

Changes in abundance of the northern Benguela sardine stock during the decade 1990–2000, with comments on the relative importance of fishing and the environment

The northern Benguela stock of sardine Sardinops sagax used to be considered one of the major clupeoid stocks of the world; it supported an average annual catch of >700 000 tons throughout the 1960s. The stock has been in a depressed state for more than two decades, as demonstrated by annual catches that averaged around 50 000 tons between 1978 and 1989 and only slightly more in the 1990s. It has experienced fluctuations in abundance of several orders of magnitude during the most recent decade. Population size increased until 1992, when the acoustic estimate of biomass was about 750 000 tons. Catches increased accordingly, averaging 100 000 tons between 1992 and 1995, but from 1992 to 1996 the stock was in decline and the lowest annual catch in the history of the fishery was taken in 1996. Although there was a small increase during the last three years of the decade, the stock remains seriously depleted. Survey-based recruitment indices suggest that the changes in the 1990s were initiated by fluctuations in recruitment, but the decline was almost certainly exacerbated by continued fishing. Poor recruitment and decreasing catch rates between 1993 and 1996 in a number of other key resources suggest that system-wide environmental changes were an important factor in the decline of the sardine stock at that time. Anomalous oceanographic conditions, such as extensive hypoxic shelf waters in 1993/94 and a Benguela Niño in 1995, support this conclusion.     

Characterising and comparing the spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem were characterised by comparing their egg abundances with environmental variables measured concomitantly during two different survey programmes: the South African Sardine and Anchovy Recruitment Programme (SARP), which comprised monthly surveys conducted during the austral summers of 1993/94 and 1994/95; and annual pelagic spawner biomass surveys conducted in early summer (November/December) from 1984 to 1999. Eggs were collected using a CalVET net. Physical variables measured included sea surface temperature (SST), surface salinity, water depth, mixed-layer depth, and current and wind speeds; biological variables measured included phytoplankton biomass, and zooplankton biomass and production. Spawning habitat was identified by construction of quotient curves derived from egg abundance data and individual environmental variables, and relationships between these variables were determined using multivariate co-inertia analysis. SARP data showed that anchovy spawning was associated with cool water and moderate wind and current speeds, whereas sardine spawning was related to warmer water and more turbulent and unstable conditions (i.e. high wind speeds and strong currents) than for anchovy. SARP data also showed significant differences in selection of spawning habitat of the two species for all environmental variables. The relationship between anchovy egg abundance and salinity was strongly positive, but strongly negative with water depth, phytoplankton biomass and zooplankton production. Sardine egg abundance was strongly positively related to current speed. The spawner biomass survey data demonstrated that the spawning habitat of anchovy was characterised by warm water and high salinity, whereas sardine spawning was associated with cool water and low salinity. The survey data showed significant differences in spawning habitat selection by anchovy and sardine for SST, salinity and zooplankton biomass, but not for the other environmental variables. There was a positive relationship between anchovy egg abundance and SST, salinity and mixed-layer depth, and a negative relationship with water depth, phytoplankton biomass and zooplankton production. For sardine there was a strong positive relationship between egg abundance and current speed and wind speed. Differences in the results between the two survey programmes could be attributable to differences in their spatio-temporal coverage. Spawning habitats of anchovy and sardine appear to be substantially different, with anchovy being more specific than sardine in their preference of various environmental conditions.     

Comparison of assemblages and some life-history traits of seabirds in the Humboldt and Benguela systems

There are 21 and 15 species of seabirds that breed in the Humboldt and Benguela upwelling systems respectively. Only two species of gull are common to both systems, one as an endemic subspecies to the Benguela system. Eleven species and two subspecies are endemic (or nearly so) to the Humboldt system; seven species and one subspecies to the Benguela system. Each system has an endemic penguin, sulid, cormorant and tern that feed mainly on anchovy Engraulis spp., sardine Sardinops sagax or both these fish. The Peruvian pelican Pelecanus thagus also feeds primarily on these prey items. A plentiful availability of food has resulted in many of these seabirds attaining high levels of abundance. For the four pairs of species that feed on anchovy and sardine, those in the Humboldt system all have a biology that enables them to increase more rapidly than their Benguela counter-parts. This reflects the higher frequency of environmental perturbations that depress seabird populations in the Humboldt system. In addition, both systems have a small endemic cormorant that feeds near the coast and a small endemic tern that breeds in the adjacent mainland desert and feeds at the sea surface. Several seabirds endemic to a system have no obvious ecological equivalent in the other system: the pelican, a diving-petrel, four storm-petrels and a gull in the Humboldt system; a cormorant and a gull in the Benguela system. Some species with tropical or subantarctic affinities breed at the boundaries of the systems. Others breed also in freshwater systems. The grey gull Larus modestus, which feeds in the Humboldt system, breeds in montane deserts.