Responses of African penguins to regime changes of sardine and anchovy in the Benguela system

Regional trends in numbers of African penguins Spheniscus demersus conform with an altered distribution of sardine Sardinops sagax and anchovy Engraulis capensis prey. In the 1950s, sardine dominated the pelagic fish component of the Benguela system. Abundance of this fish decreased in the 1960s and early 1970s and it was replaced by anchovy. Beginning in the early 1980s, sardine started to increase and anchovy to decrease in abundance. The decrease of the sardine resulted in collapses of colonies of African penguins between Lüderitz and Table Bay. Although colonies east of Table Bay increased, the overall population of African penguins fell by 25%. As the Benguela system started to revert to one dominated by sardine, penguin colonies between Lüderitz and Table Bay stabilized. Three new colonies were established in the vicinity of Table Bay, but the large colony at Dyer Island underwent a massive decrease. As a result, overall numbers of African penguins decreased by a further 19%. During shifts between regimes dominated by sardine and anchovy, African penguins that are breeding for the first time immigrate to colonies where food is plentiful. Inability of African penguins to cope with recent shifts between regimes may have resulted from increased competition for food with fishermen and seals during the 20th century

Comparison of trends in abundance of guano-producing seabirds in Peru and Southern Africa

The abundant guano-producing seabirds in Peru and southern Africa feed mainly on the large populations of anchovy Engraulis spp. and sardine Sardinops sagax supported by the Humboldt and Benguela upwellingsystems. Numbers of guanay cormorants Phalacrocorax bougainvillii in Peru and the breeding population of Cape cormorants P. capensis in South Africa are significantly related to the biomass of anchovy. For both species,reproductive success decreases in periods of anchovy scarcity, and there may also be substantial adult mortality. There has been long-term stability in numbers of Peruvian boobies Sula variegata, whereas the numbers ofCape gannets Morus capensis decreased as sardine decreased in southern Africa. Numbers of Peruvian pelicans Pelecanus (occidentalis) thagus are significantly related to the combined biomass of anchovy and sardine inPeru. They have been stable in the long-term. There have been ongoing severe decreases in populations of the Humboldt penguin Spheniscus humboldti and the African penguin S. demersus, both of which are nowVulnerable. Common causes for the decreases have been collection of eggs, loss of habitat through exploitation of accumulated deposits of guano and competition with fisheries for food. Short-term decreases in guanay cormorants, Peruvian boobies, Peruvian pelicans and Humboldt penguins have followed El Niño events. Time-series of indices of the abundance of guano-producing seabirds date from 1908 in Peru and 1896 in southern Africa. They are significantly, negatively correlated in the period prior to the development of intensive fisheries on sardineand anchovy, suggesting that the out-of-phase nature of the anchovy and sardine populations in the Peru and Benguela systems pre-dated commercial exploitation of these fish resources

Season of moult of African penguins at Robben Island, South Africa, and its variation, 1988 –1998

Counts of African penguins Spheniscus demersus in immature and adult plumage in the feather-shedding phase of moult were made at Robben Island at two-weekly intervals over a 10-year period between 1988 and1998. For both age-classes, most birds moulted between November and January, although small numbers moulted throughout the year. In most years, the peak moult was in late November or December. Immature birdshad a secondary peak in March. In 1994/95 for adults, and 1995/96 for immature birds, moult was less synchronized than in other years. This probably resulted from oiling of about 2 400 penguins in June 1994, following the sinking of the Apollo Sea. Half of those birds were cleaned and released. Their subsequent moult may have been earlier than normal

Age at first breeding and change in plumage of kelp gulls Larus Dominicanus in South Africa

In South Africa, kelp gulls Larus dominicanus first breed when 4 years old. At least 50% of 4-year-olds breed; virtually all older birds breed annually. Most chicks leave natal colonies in the austral autumn and winter, when about 6 months old, and 60% of them may not return until aged 2 or 3 years. One 3-year-old showed signs of defending territory, but birds only attain full adult plumage when aged 4. The plumage of 3-year-olds is similar to that of adults, but the white spots on the outer primaries are less well developed and the secondaries and inner primaries have no white trailing edge. Chicks are dark, whereas birds aged 1 and 2 years gain increasing amounts of pale or white plumage on the face, neck, underparts and tail

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 1996and 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. Theyresponded 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 colonygrowth 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 frequentlyvisited by penguins from other colonies, and by rehabilitated birds

Bottom-up effects of a no-take zone on endangered penguin demographics.

This is the author accepted manuscript. The final version is available from The Royal Society via the DOI in this record.Marine no-take zones can have positive impacts for target species and are increasingly important management tools. However, whether they indirectly benefit higher order predators remains unclear. The endangered African penguin (Spheniscus demersus) depends on commercially exploited forage fish. We examined how chick survival responded to an experimental 3-year fishery closure around Robben Island, South Africa, controlling for variation in prey biomass and fishery catches. Chick survival increased by 18% when the closure was initiated, which alone led to a predicted 27% higher population compared with continued fishing. However, the modelled population continued to decline, probably because of high adult mortality linked to poor prey availability over larger spatial scales. Our results illustrate that small no-take zones can have bottom-up benefits for highly mobile marine predators, but are only one component of holistic, ecosystem-based management regimes.We thank the Earthwatch Institute, Bristol Zoological Society, Leiden Conservation Foundation and National Research Foundation

Defining ecologically relevant scales for spatial protection with long-term data on an endangered seabird and local prey availability

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Human activities are important drivers of marine ecosystem functioning. However, separating the synergistic effects of fishing and environmental variability on the prey base of nontarget predators is difficult, often because prey availability estimates on appropriate scales are lacking. Understandi ng how prey abundance at different spatial scales links to population change can help integrate the needs of nontarget predators into fisheries management by defining ecologically relevant areas for spatial protection. We investigated the local population response (number of breeders) of the Bank Cormorant (Phalacrocorax neglectus), a range-restricted endangered seabird, to the availability of its prey, the heavily fished west coast rock lobster (Jasus lalandii). Using Bayesian state-space modeled cormorant counts at 3 colonies, 22 years of fisheries-independent data on local lobster abundance, and generalized additive modeling, we determined the spatial scale pertinent to these relationships in areas with different lobster availability. Cormorant numbers responded positively to lobster availability in the regions with intermediate and high abundance but not where regime shifts and fishing pressure had depleted lobster stocks. The relationships were strongest when lobsters 20–30 km offshore of the colony were considered, a distance greater than the Bank Cormorant's foraging range when breeding, and may have been influenced by prey availability for nonbreeding birds, prey switching, or prey ecology. Our results highlight the importance of considering the scale of ecological relationships in marine spatial planning and suggest that designing spatial protection around focal species can benefit marine predators across their full life cycle. We propose the precautionary implementation of small-scale marine protected areas, followed by robust assessment and adaptive-management, to confirm population-level benefits for the cormorants, their prey, and the wider ecosystem, without negative impacts on local fisheries.The National Research Foundation (NRF) SEAChange Grant 79735 and an incentive grant to R.J.M.C., Leiden Conservation Foundation, Claude Leon Foundation, and our institutions provided funding

African penguins as predators and prey — coping (or not) with change

African penguins Spheniscus demersus live in the Benguela and western Agulhas ecosystems off southern Africa. Their numbers decreased throughout the 20th century from at least 1.5 million to about 0.18 million adults, although different regional trends were apparent. They feed to a large extent on shoaling epipelagic fish, notably anchovy Engraulis capensis and sardine Sardinops sagax, and regional trends in the abundance of penguins are associated with trends in the abundance and distribution of these prey fish. Many first-time breeders emigrate from colonies where feeding or other conditions at the time are unfavourable to more favourable breeding localities. This has led to both the extinction and formation of colonies. Food now may limit colonies at relatively small sizes, a fact attributable to industrial fisheries reducing the densities of forage fish. African penguins share their habitat with several other predators, with which they compete for food and breeding space. One of these, the Cape fur seal Arctocephalus p. pusillus, increased through the 20th century to 1.5–2 million animals at its close. Reported observations of predation by fur seals on seabirds have increased in recent decades and threaten the continued existence of small colonies of penguins. Stochastic modelling suggests that colonies of 10 000 pairs have a 9% probability of extinction in 100 years, so smaller populations should be regarded as “Vulnerable”. However, in a period of prolonged food scarcity off southern Namibia, the regional population decreased from more than 40 000 pairs in 1956 to about 1 000 pairs in 2000, and many colonies numbering less than 1 000 pairs became extinct. The minimum viable population for African penguins is currently considered to be >40 000 pairs, likely of the order of 50 000 pairs, a figure equivalent to its level in 2000. The chance of survival of the species through the 21st century is tenuous.African Journal of Marine Science 2001, 23: 435–44

Bayesian inference reveals positive but subtle effects of experimental fishery closures on marine predator demographics

Global forage-fish landings are increasing, with potentially grave consequences for marine ecosystems. Predators of forage fish may be influenced by this harvest, but the nature of these effects is contentious. Experimental fishery manipulations offer the best solution to quantify population-level impacts, but are rare. We used Bayesian inference to examine changes in chick survival, body condition and population growth rate of endangered African penguins Spheniscus demersus in response to 8 years of alternating time–area closures around two pairs of colonies. Our results demonstrate that fishing closures improved chick survival and condition, after controlling for changing prey availability. However, this effect was inconsistent across sites and years, highlighting the difficultly of assessing management interventions in marine ecosystems. Nevertheless, modelled increases in population growth rates exceeded 1% at one colony; i.e. the threshold considered biologically meaningful by fisheries management in South Africa. Fishing closures evidently can improve the population trend of a forage-fish-dependent predator—we therefore recommend they continue in South Africa and support their application elsewhere. However, detecting demographic gains for mobile marine predators from small no-take zones requires experimental time frames and scales that will often exceed those desired by decision makers

Food limitation of seabirds in the Benguela ecosystem and management of their prey base

This is the final version. Available from the Environmental Information Service, Namibia via the URL in this record. Four of seven seabirds that are endemic to the Benguela ecosystem (African Penguin Spheniscus demersus, Cape Gannet Morus capensis, Cape Cormorant Phalacrocorax capensis, Bank Cormorant P. neglectus) compete with fisheries for prey and have an IUCN classification of Endangered. Prey depletion and food resource limitations have been major drivers of recent large population decreases of each of these species. As populations decrease, colony sizes also dwindle rendering them susceptible to Allee effects and higher probabilities of extinction. Therefore, it is necessary to maintain colonies at sizes that minimise their probability of extinction. Means to ensure an adequate availability of food to achieve this goal include closing important seabird foraging areas (often adjacent to key colonies) to relevant fishing, implementing ecosystem thresholds below which such fishing is disallowed (which are also expected to benefit forage resources) and, should there be an altered distribution of prey, attempting to establish seabird colonies close to the new location of forage resources.The Pew Charitable Trust