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.     

Long-term trends in cetacean occurrence during the annual sardine run off the Wild Coast,South Africa

During the austral winter, cetaceans and other apex predators follow the annual northeastward movement of shoaling sardines, known as the sardine run, along the southeast coast of South Africa, including a 400-km stretch called the Wild Coast. In total, 131 opportunistic aerial surveys were conducted between May and July, from 1996 to 2014, to monitor sardine movement. Cetacean-sighting data from these surveys were analysed, focusing on long-term trends in frequencies of the cetaceans. In total, 630 sightings involving five cetacean species were recorded: 268 (approximately 32 400 individuals) of Indo-Pacific bottlenose dolphin Tursiops aduncus, 108 (approximately 79 400 individuals) of long-beaked common dolphin Delphinus capensis, 242 (approximately 670 individuals) of humpback whale Megaptera novaeangliae, 1 (two individuals) of southern right whale Eubalaena australis, and 11 (16 individuals) of Bryde’s whale Balaenoptera edeni. The occurrence of common dolphins, typically associated with sardines, decreased significantly in average group size over the study period (p = 0.0343); bottlenose dolphins, considered generalist feeders, demonstrated no such trend (p = 0.916). Humpback whales were most frequently sighted between 2010 and 2014, and with significantly larger groups observed towards the end of the study period (p = 0.0121). For all these species, more sightings were made inside than outside of marine protected areas (>70% of the dolphin species, and >65% of the humpback whales), both pre- and post-2005 (from 2005 the size of the survey area increased). The results indicate that movements of the common dolphin may be employed as a proxy for sardine occurrence. Long-term trends evident in the data also demonstrate the importance of this coastal region for bottlenose dolphins as well as use as a migratory corridor for humpback whales.     

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.     

Assessment and management of Cape horse mackerel Trachurus capensis off Namibia based on a fleet-disaggregated age-structured production model

The Cape horse mackerel stock off Namibia is assessed using a fleet-disaggregated age-structured production model including historical International Commission for the Southeast Atlantic Fisheries (ICSEAF) and current data. Data from acoustic biomass surveys, on commercial catch per unit effort and on commercial catch-at-age, were used to estimate current stock status over a range of model specifications. A management tool, the so-called management monitor graph, was introduced in order to track past management relative to stock status. Target, limit and precautionary reference points, decision rules and harvesting strategies, according to which the stock should be managed, are described and tested. The target reference point was set at maximum sustainable yield (MSY), the precautionary reference point at 80% of MSY and the limit reference point at 30% of MSY. Owing to some inconsistencies in the dataseries, evaluation of the current state of the resource is challenging, but estimates are that the resource is around the MSY level, somewhere between 250 000 and 300 000 t.     

A hard-knock life: the foraging ecology of Cape cormorants amidst shifting prey resources and industrial fishing pressure

Once one of the most numerous seabirds of the Benguela upwelling system, the population of Cape cormorants Phalacrocorax capensis has decreased by 60% in the past three decades and the species is listed as Near Threatened. Declines in prey availability and/or abundance brought about by recent changes in the distribution of pelagic fish stocks and industrial purse-seine fishing are hypothesised to be a key driver of seabird population decreases in the southern Benguela. We investigated the foraging behaviour of breeding Cape cormorants by deploying GPS and temperature–depth recorders on 24 breeding adults from three islands off the coast of South Africa, two of them to the north of Cape Point and a third farther south on the western Agulhas Bank. This provided the first measures of foraging dispersal by a cormorant in the Benguela system, and enabled a comparison of foraging behaviour between birds from these islands. Foraging trips of Cape cormorants lasted between 17 min and >7 h, at a maximum distance of between 2 and 58 km away from their colony. Foraging effort was significantly greater for birds from farther north off the West Coast in terms of trip duration, distance travelled, number of dives and time spent flying compared to those from the southernmost island (Dyer), which is probably a response to low prey availability in the north. Coastal reserves that exclude pelagic fishing from inshore feeding grounds around Cape cormorant breeding colonies may result in increased local prey availability, which would benefit Cape cormorant populations.     

Cape cormorants decrease,move east and adapt foraging strategies following eastward displacement of their main prey

Numbers of Cape cormorants Phalacrocorax capensis breeding in South Africa decreased by nearly 50% from approximately 107 000 pairs in 1977–1981 to 57 000 pairs in 2010–2014. Although four colonies had >10 000 pairs in 1977–1981, there was just one such colony in 2010–2014. Almost all the decrease occurred after the early 1990s off north-west South Africa, between the Orange River estuary and Dassen Island. South of this, the number breeding in the two periods was stable, with some colonies being formed or growing rapidly in the 2000s. The proportion of South Africa’s Cape cormorants that bred south of Dassen Island increased from 35% in 1977–1981 to 66% in 2010–2014, with the opposite situation observed in the north-west. This matched a shift to the south and east in the distributions of two of the Cape cormorant’s main prey species, anchovy Engraulis encrasicolus and sardine Sardinops sagax. In 2014, an apparent scarcity of prey in the north-west resulted in Cape cormorants attempting to take bait from hooks of fishing lines over an extended period, a behaviour not previously recorded. The number of Cape cormorants breeding in the south may be constrained by the absence of large islands between Dyer Island in the west and Algoa Bay in the east. If so, it may be possible to bolster the southern population through the provision of appropriate breeding habitat, such as platforms, or restricting human disturbance at suitable mainland cliff breeding sites.     

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.     

Trends in numbers of crowned cormorants in South Africa,with information on diet

During 2008–2012, the number of crowned cormorants Phalacrocorax coronatus breeding in South Africa was c. 1 900 pairs, compared to 1 700 pairs for 1977–1981. Numbers at 10 islands in the Western Cape province fluctuated around a level of 1 100 pairs from 1991/1992 to 2011/2012, 300 more than from 1978/1979 to 1990/1991. These increases are attributable to the discovery of more colonies and an increased frequency of counting at the 10 islands after 1990/1991. The overall number of crowned cormorants breeding in South Africa is thought stable in the long term. Crowned cormorants feed mainly on small, inshore fish species that are not harvested by humans. Clinidae dominated the diet at 10 colonies adjoining the open sea, whereas Gobiidae contributed most food of birds at three colonies in a lagoon. The stability of the crowned cormorant population contrasts with decreases of some other seabirds endemic to southern Africa that feed primarily on prey that is exploited by fisheries. The crowned cormorant population decreased in the Northern Cape and small numbers initiated breeding at colonies to the east of Cape Agulhas at the turn of the century, but most of the population continues to breed to the west of Cape Agulhas. In some instances the availability of suitable breeding habitat may limit numbers breeding.