Azwianewi B. Makhado

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Collapse of South Africa's penguins in the early 21st century

The number of African penguins Spheniscus demersus breeding in South Africa collapsed from about 56 000 pairs in 2001 to some 21 000 pairs in 2009, a loss of 35 000 pairs (>60%) in eight years. This reduced the global population to 26 000 pairs, when including Namibian breeders, and led to classification of the species as Endangered. In South Africa, penguins breed in two regions, the Western Cape and Algoa Bay (Eastern Cape), their breeding localities in these regions being separated by c. 600 km. Their main food is anchovy Engraulis encrasicolus and sardine Sardinops sagax, which are also the target of purse-seine fisheries. In Algoa Bay, numbers of African penguins halved from 21 000 pairs in 2001 to 10 000 pairs in 2003. In the Western Cape, numbers decreased from a mean of 35 000 pairs in 2001–2005 to 11 000 pairs in 2009. At Dassen Island, the annual survival rate of adult penguins decreased from 0.70 in 2002/2003 to 0.46 in 2006/2007; at Robben Island it decreased from 0.77 to 0.55 in the same period. In both the Western and Eastern Cape provinces, long-term trends in numbers of penguins breeding were significantly related to the combined biomass of anchovy and sardine off South Africa. However, recent decreases in the Western Cape were greater than expected given a continuing high abundance of anchovy. In this province, there was a south-east displacement of prey around 2000, which led to a mismatch in the distributions of prey and the western breeding localities of penguins.

Impact of predation by Cape fur seals Arctocephalus pusillus pusillus on Cape gannets Morus capensis at Malgas Island, Western Cape, South Africa

Cape fur seals Arctocephalus pusillus pusillus were estimated to kill some 6 000 Cape gannet Morus capensis fledglings around Malgas Island in the 2000/01 breeding season, 11 000 in 2003/04 and 10 000 in 2005/06. This amounted to about 29%, 83% and 57% of the overall production of fledglings at the island in these breeding seasons respectively. Preliminary modelling suggests this predation is not sustainable. There was a 25% reduction in the size of the colony, the second largest of only six extant Cape gannet colonies, between 2001/02 and 2005/06. There has been a large increase in predation by Cape fur seals on seabirds around southern African islands since the mid-1980s, coincidental with both an increase in the seal population, altered management of the islands and an altered distribution of prey for gannets and seals. At Malgas Island, most gannet fledglings were killed between 10:00 and 18:00, the period when most are in the water around the island, from mid-January to mid-March, the main fledging period. The Cape gannet is classified as Vulnerable.

Pup growth and maternal attendance patterns in Subantarctic fur seals

Samples of pups from the expanding populations of Subantarctic fur seals (Arctocephalus tropicalis) atMarion and Gough islands were weighed at specific ages between birth and weaning in the period 1993-2000. Growth was estimated and compared between years, sexes, sites of different density, and populations. Pups of comparable age were consistently heavier at Marion Island than at Gough Island, probably reflecting relative prey resource abundance. No changes in pupgrowth rates were apparent over the study period, nor was there any evidence that growth was affected by ENSO events. Measures of pup growth at Marion Island were complemented by measures of maternal attendance behaviour during three summer andwinter seasons. More years of data are required to determine patterns between attendance, growth, resource conditions and population abundance trends.

Mass on arrival of rockhopper penguins at Marion Island correlated with breeding success

For rockhopper penguins Eudyptes chrysocome at Marion Island, there were significant decreases over time in the numbers breeding and breeding success at three monitored colonies from 1985/1986 to 2006/2007, and in mass on arrival for breeding of both males and females from 1994/1995 to 2007/2008. Breeding success decreased by 0.15 chicks pair−1 y−1 over 22 years and was significantly correlated with mass on arrival of males and females. Survival of chicks at the guard stage was low in the late 1990s; hatching success decreased in the early 2000s. It is thought that an increasingly poor parental condition caused birds to abandon breeding at a progressively early stage. Parental condition is influenced by feeding opportunities at overwintering grounds, which have probably been altered by global climate.

Winners and losers – responses to recent environmental change by South African seabirds that compete with purse-seine fisheries for food

There were recent changes in the distributions of the main prey of four seabirds off South Africa, with sardine Sardinops sagax and anchovy Engraulis encrasicolus shifting from South Africas north-west coast towards the south-east. This contributed to numbers of African Penguins Spheniscus demersus and Cape Cormorants Phalacrocorax capensis breeding in South Africa decreasing by more than 50%. By contrast, numbers of Cape Gannets Morus capensis and Swift (Crested) Terns Thalasseus bergii increased. The success of the gannet and the tern in the face of recent change is attributable to a rapid increase in South Africas eastern colony of gannets, aided by high survival of adult birds and perhaps by immigration of first-time breeders from decreasing western colonies, and a probable high production of Swift Terns. The gannet has an extensive foraging range and the tern is nomadic between breeding localities, which gives these species greater flexibility than the other two to adapt to large changes in the distributions of their prey. Additionally, the gannet is able to supplement its diet with fishery discards. A lesser mobility of the African Penguin and the Cape Cormorant rendered them susceptible to a reduced local availability of prey and both species suffered high adult mortality in the last decade.

A changing distribution of seabirds in South Africa—the possible impact of climate and its consequences

In the southern Benguela ecosystem off South Africa, there were recent shifts to the south and east in the distributions of three forage resources (anchovy, sardine, rock lobster), which probably were influenced by environmental change although fishing too may have played a part. In this study, we review information on trends in distributions and numbers of eight seabirds breeding in South Africa. For five species that feed predominantly on anchovy, sardine or rock lobster, their populations off northwest South Africa decreased markedly. For three of these species, which exhibit behavioural inertia and have restricted foraging ranges when breeding (African penguin, Cape cormorant, bank cormorant), there were large decreases in their overall populations in South Africa. Conversely, for two showing more plasticity and able to range over wide areas or move between breeding localities (Cape gannet, swift tern) there were increases. It is thought that movement of forage resources away from the northern islands led to a mismatch in the distributions of breeding localities and prey of dependent seabirds off western South Africa and to attempts by several species to establish colonies on the southern mainland closer to food resources. There also were shifts to the south and east in the distributions of three seabirds that do not compete with fisheries for prey (crowned cormorant, white-breasted cormorant, kelp gull), suggesting some environmental forcing, but decreases of these species off northwest South Africa were less severe and populations in South Africa remained stable or increased in the long term. It is likely, because many fishing plants are located in the northwest, that there was increased competition between seabirds and fisheries for prey as forage resources moved south and east. Potential interventions to mitigate the adverse impacts of distributional changes for seabirds include allocations of allowable catches of shared forage resources at regional levels,

A space oddity: Geographic and specific modulation of migration in Eudyptes penguins

Post-breeding migration in land-based marine animals is thought to offset seasonal deterioration in foraging or other important environmental conditions at the breeding site. However the inter-breeding distribution of such animals may reflect not only their optimal habitat, but more subtle influences on an individual’s migration path, including such factors as the intrinsic influence of each locality’s paleoenvironment, thereby influencing animals’ wintering distribution. In this study we investigated the influence of the regional marine environment on the migration patterns of a poorly known, but important seabird group. We studied the inter-breeding migration patterns in three species of Eudyptes penguins (E. chrysolophus, E. filholi and E. moseleyi), the main marine prey consumers amongst the World’s seabirds. Using ultra-miniaturized logging devices (light-based geolocators) and satellite tags, we tracked 87 migrating individuals originating from 4 sites in the southern Indian Ocean (Marion, Crozet, Kerguelen and Amsterdam Islands) and modelled their wintering habitat using the MADIFA niche modelling technique. For each site, sympatric species followed a similar compass bearing during migration with consistent species-specific latitudinal shifts. Within each species, individuals breeding on different islands showed contrasting migration patterns but similar winter habitat preferences driven by sea-surface temperatures. Our results show that inter-breeding migration patterns in sibling penguin species depend primarily on the site of origin and secondly on the species. Such site-specific migration bearings, together with similar wintering habitat used by parapatrics, support the hypothesis that migration behaviour is affected by the intrinsic characteristics of each site. The paleo-oceanographic conditions (primarily, sea-surface temperatures) when the populations first colonized each of these sites may have been an important determinant of subsequent migration patterns. Based on previous chronological schemes of taxonomic radiation and geographical expansion of the genus Eudyptes, we propose a simple scenario to depict the chronological onset of contrasting migration patterns within this penguin group.

An assessment of the impact of predation by Cape fur seals Arctocephalus pusillus pusillus on seabirds at Dyer Island, South Africa

At Dyer Island, South Africa, observations of predation of seabirds by Cape Fur Seals Arctocephalus pusillus pusillus were made during 2004 and 2006/07. It was estimated that seals killed about 7% of adult African Penguins Spheniscus demersus annually. This may have contributed to the penguin colony not increasing in spite of food becoming more available in its vicinity during an eastward shift of epipelagic fish resources off South Africa at the start of the twenty-first century. Adult penguins were most susceptible to mortality in the breeding season as they returned to feed chicks in the evening. Seals also killed substantial numbers (3–9%) of Cape Cormorant Phalacrocorax capensis fledglings as they left the island, mostly in the morning. The losses of Cape Cormorant fledglings accounted for about 95% of the overall mortality of seabirds attributable to seals. Mortality inflicted by seals on adults of four species of cormorant was negligible and no predation by seals on gulls and terns was observed.

South Africa’s coastal-breeding white-breasted cormorants: population trends, breeding season and movements, and diet

White-breasted cormorants Phalacrocorax [carbo] lucidus breed around South Africas coast and at inland localities. Along the coasts of the Northern, Western and Eastern Cape provinces, numbers breeding were similar during the periods 1977–1981 (1 116 pairs at 41 localities) and 2008–2012 (1 280 pairs at 41 localities). Along the coast of KwaZulu-Natal (not counted in 1977–1981), 197 pairs bred at nine localities in 2008–2012, when the overall number breeding around South Africas coastline was about 1 477 pairs. Between the two study periods, numbers decreased in the Northern and Western Cape provinces following the loss of several breeding localities, but they increased in the Eastern Cape. In the Western Cape, however, numbers were stable east of Cape Agulhas and at nine well-monitored West Coast localities that were surveyed from 1978 to 2012. White-breasted cormorants breed throughout the year, with breeding at some localities more seasonal than at others and the timing of peaks in breeding varying at and between localities. In the vicinity of Saldanha Bay/Langebaan Lagoon (Western Cape), in Algoa Bay (Eastern Cape) and in northern KwaZulu-Natal, it is likely that birds moved between breeding localities in different years, although breeding often occurred at the same locality over several years. Human disturbance, presence of predators, competition for breeding space and occurrence of breeding by other waterbirds may influence movements between colonies. Securing sufficient good habitat at which white-breasted cormorants may breed will be important for conservation of the species. The species may breed at an age of 4 years, possibly younger. The bulk of their diet around South Africas coast consists of inshore marine and estuarine fish species that are not intensively exploited by humans.