Laurent Drapeau

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.

Broad-scale distribution patterns of sardine and their predators in relation to remotely sensed environmental conditions during the KwaZulu-Natal sardine run

The annual movement of South African sardine Sardinops sagax up the east coast of South Africa, known as the ‘sardine run’, was investigated using data from aerial surveys for the period 1988–2005 and compared with remotely sensed sea surface temperature (SST) and chlorophyll a data. Sardine sighting rates were highest within the Waterfall Bluff Bight off the Eastern Cape Coast, where conditions appeared to be most favourable. Sardine and predator sightings decreased significantly northwards of Mdoni on the KwaZulu-Natal (KZN) coast, whereas the proportion of nearshore sightings increased. The causal mechanism for this inshore concentration is suggested to be the influx of warm Agulhas Current water from the Durban Eddy that forces sardine shoreward. Cape gannet Morus capensis, common dolphin Delphinus capensis and sardine distributions were associated, and there was an association between SST and sardine and predator distributions. There was a marked increase in bottlenose dolphin Tursiops aduncus sightings upon commencement of the sardine run, with these dolphins being consid-ered to be a ‘migratory’ stock that enters KZN waters every winter.

Quantification and representation of potential spatial interactions in the southern Benguela ecosystem

This work explores the potential spatial interactions between 13 key commercial species of the southern Benguela ecosystem: sardine Sardinops sagax, anchovy Engraulis encrasicolus, round herring Etrumeus whiteheadi, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus, chokka squid Loligo vulgaris reynaudii, kingklip Genypterus capensis, Cape hake Merluccius spp., silver kob Argyrosomus inodorus, snoek Thyrsites atun, albacore Thunnus alalunga, bigeye tuna Thunnus obesus and yellowfin tuna Thunnus albacares. It is based on distribution maps per species after combining available commercial and research databases. The resulting 78 pairs of potential interactions are quantified using three indices: the overlap in area, the overlap in biomass and the weighted kappa index. From additional information on the diet of the different species and trophic models, the main trophic interactions (predation or competition) were identified and mapped. The results are discussed with regard to methodological limitations, habitat selection, fish assemblages, the need for spatial resolution of trophic models and the ecosystem approach to fishery management.

The KwaZulu-Natal sardine run: shoal distribution in relation to nearshore environmental conditions, 1997–2007

The nearshore presence of sardine Sardinops sagax on the KwaZulu-Natal (KZN) coast was investi-gated using sightings data collected by the KZN Sharks Board from 1997 to 2007. The spatio-temporal distribution of sardine was described in relation to that of their predators and to environmental conditions, and subjected to generalised linear model (GLM) and generalised additive model (GAM) analyses. Variables describing spatio-temporal conditions performed best in the models (r2 = 0.52) with seasonal effects, specifically June and July, making the greatest contribution towards sardine presence. The contribution of the years 2003, 2006 and 2007, and the KZN North Coast, was signif-icantly lower. The predator variables were highly significant (r2 = 0.48) with Cape gannets Morus capensis, followed by the sharks/gamefish and common dolphins Delphinus capensis, being most closely associated with sardine presence. Environmental variables were not as influential in the GLM models (r2 = 0.23), but some variables were useful in describing conditions favouring sardine presence, namely calm current conditions, light north-westerly land breezes and stable atmospheric conditions. Increasing sea surface temperature (SST), moderate north to south currents, large swells and turbid water had a negative impact upon sardine presence. North-easterly and north-westerly winds and north to south currents had a cooling effect upon nearshore SSTs, whereas south-easterly winds and increasing air temperatures caused nearshore warming. Results are discussed in the context of developing an understanding of the mechanisms that govern fine-scale movements of sardine shoals during the KZN sardine run, with a view to predicting such movements.

DISTRIBUTION PATTERNS OF KEY FISH SPECIES OF THE SOUTHERN BENGUELA ECOSYSTEM: AN APPROACH COMBINING FISHERY-DEPENDENT AND FISHERY-INDEPENDENT DATA

Within the context of an ecosystem approach for fisheries, there is a need for quantitative information on distributions of key marine species. This information is valuable input for modelling species interactions in the southern Benguela ecosystem. In the present study, a method is described for mapping the density distribution of 15 key species: anchovy Engraulis encrasicolus, sardine Sardinops sagax, round herring Etrumeus whiteheadi, chub mackerel Scomber japonicus, horse mackerel Trachurus trachurus capensis, lanternfish Lampanyctodes hectoris, lightfish Maurolicus muelleri, albacore Thunnus alalunga, bigeye tuna Thunnus obesus, yellowfin tuna Thunnus albacares, silver kob Argyrosomus inodorus, snoek Thyrsites atun, Cape hake Merluccius spp., kingklip Genypterus capensis and chokka squid Loligo vulgaris reynaudi. The purpose was to make use of all available sources of data to extend the spatial and temporal coverage of the southern Benguela. Six sources of data were combined on a 10′ × 10′ cell grid in a Geographical Information System: acoustic and demersal surveys conducted by Marine and Coastal Management (MCM), and pelagic, demersal (including midwater trawl), hake-directed and tuna-directed longline commercial landings data collected by MCM. Comparisons of distributions between two periods (1980s and 1990s) and between two semesters (April – September and October – March) were conducted, but biases as a result of major differences in sampling strategy prevented detailed analysis for certain species. Maps of density distributions are nevertheless presented here and the method to determine them is discussed.

Do inter-colony differences in Cape fur seal foraging behaviour reflect large-scale changes in the northern Benguela ecosystem?

The northern Benguela ecosystem adjoining Namibia has undergone considerable changes in recent decades, with reductions and northwards shifts of key prey species that have had severe implications for marine top predator populations. We investigated how such environmental variability may impact foraging behaviour of the Cape fur seal Arctocephalus pusillus pusillus, using satellite telemetry on animals in northern, central and southern Namibia. We expected that seal foraging behaviour would reflect a gradient of deteriorating feeding conditions from north to south. Results showed that foraging trips were shorter in the central region, where seals fed over the continental shelf, than in the north or south, where seals fed at the shelf edge. However, whereas seals in the north showed strong fidelity to the colony at which they were tagged and to persistent, clustered foraging areas, seals in the south showed weak fidelity both to the colony at which they were tagged and to foraging areas, which were scattered and variable. Seals in the south also tended to migrate northwards to other colonies while concurrently adapting their foraging behaviour to local conditions. Flexible use of foraging space and colonies of Cape fur seals during the three-year period (2002-2004) demonstrates that the species is adapted for variable environments over time and space.

The food of three seabirds at Latham Island, Tanzania, with observations on foraging by masked boobies Sula dactylatra

At Latham Island, Tanzania, in August 2004 and December 2005, flying fish (Exocoetidae) contributed 90% by mass of prey items regurgitated by masked boobies Sula dactylatra. The fish had a mean caudal length of 164mm. Two boobies generally foraged in deep water, away from the Tanzanian coastline, ranging up to 390km from the island. Maximum distances flown on trips exceeded 1 000km during breeding and approached 3 000km when not breeding. Some foraging trips exceeded six days. Swift terns Sterna bergii consumed small fish and mantis shrimps Natosquilla investigatoris. Mantis shrimps recovered from the colony were significantly larger than those stranded on the islands beach, suggesting that swift terns preferentially took larger animals. Sooty terns Sterna fuscata fed on small fish and squid.

Fine-scale spatial variability of different stages of pelagic fish eggs over the western Agulhas Bank, South Africa

Stage-dependent spatial distributions of anchovy Engraulis encrasicolus, sardine Sardinops sagax and round herring Etrumeus whiteheadi eggs over the western Agulhas Bank South Africa were examined from samples collected at a fine-scale (1.8 km) resolution using a continuous underway fish egg sampler (CUFES). Spatial structure in egg distributions was assessed using variograms for three developmental stage groups (early-, middle- and late-stage eggs), and indices of aggregation were also computed for those stage groups. No early- or middle-stage anchovy eggs were collected, but indices of aggregation showed that early-stage sardine eggs were more aggregated than those of round herring, suggesting that adult sardine have a more contiguous distribution during spawning than have round herring. Middle-stage eggs were equally aggregated, whereas late-stage eggs were aggregated differently in all three species. Fitted empirical variograms showed that early-stage sardine eggs have low small-scale variability (nugget effect) (≤10% of total variance) compared with round herring eggs (≥50% of total variance). The autocorrelation range of both those species extends to 3.7 km. Middle-stage sardine eggs have a low nugget effect (≤10%) compared with round herring eggs (≤50%), with autocorrelation ranges of 7.4 km and 5.6 km respectively. Late-stage anchovy eggs have a lower nugget effect (≤20%) than that of sardine (≤50%) and round herring (≥100%) eggs, with autocorrelation ranges of 3.7 km for all three species. The spatial structures of the eggs of the three species were heterogeneous at the sampling scale used, indicating differences in spatial organisation of the adults during spawning. The spatial variability of the eggs of the three species is not well captured using the sampling strategy during current ichthyoplankton surveys in the southern Benguela, in which stations are spaced 18.5 km apart.