Predatory species left stranded following the collapse of the sardine Sardinops sagax (Pappe, 1854) stock off the northern Benguela upwelling system: A review

Abstract

Abstract Sardine Sardinops sagax (Pappe, 1854) is an important forage species, in the northern Benguela Upwelling System (nBUS). Sardine was a key forage species of the pelagic component in the nBUS, but the population collapsed due to a combination of overfishing and ecosystem change and variability in the nBUS. Multiple predators depended on sardine as high-quality food at the time when it was abundant. This study reviewed the current knowledge of predators diet pre- and post-collapse of the sardine population stock in the nBUS. Data were sourced from previous studies on the trophic interaction in the nBUS. The historical and contemporary diets of commercially exploited species that prey on sardine were found to be relatively well documented while there are data deficiencies for non-commercially exploited species including cetaceans, sharks and some non-commercially exploited fish species. Amongst the species most reliant on sardine as prey are African penguin Spheniscus demersus, Cape gannet Morus capensis, Cape cormorant Phalacrocorax capensis, Cape monkfish Lophius vomerinus, shallow-water hake Merluccius capensis, deep-water hake M. paradoxus, snoek Thyrsites atun, Albacore tuna Thunnus alalunga, Cape fur seal Arctocephalus pusillus, Common dolphins Delphinus delphis, Dusky dolphins Lagenorhynchus obscurus, Heaviside s dolphins Cephalorhynchus heavisidii, six gills shark Hexanchus griseus, copper sharks Carcharhinus brachyurus, school shark Galeorhinus galeus and pelagic sharks Prionace glauca. Some predatory species such as the African penguin, Cape cormorant, Cape gannet, and school shark, are considered endangered as per the International Union for the Conservation of Nature (IUCN) Red List criteria. Populations of Albacore tuna and Cape monkfish are near threatened, while Cape fur seals, common dolphins, and shallow-water hake are considered the least concerned on the IUCN list. Although the collapse of sardine has affected the food webs it has not resulted in collapse, rather there have been changes. Three prey species; bearded goby, Cape horse mackerel, and jellyfish, were identified as alternative prey for some of the predators that previously fed on sardine. This review showed how overexploitation of sardine could have had far-reaching effects on the nBUS through feeding linkages. Goby, although a low-quality prey, is a successful substitute for sardine probably mostly due to its high abundance, tolerance to low oxygen and ability to thrive in warm waters. However, further studies are needed to establish the strength of the ecological linkages of the prey species that replaced sardine in the diets of the predators.