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 1996 and 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. They responded 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 colony growth 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 frequently visited by penguins from other colonies, and by rehabilitated birds.     

Surviving off junk: low-energy prey dominates the diet of African penguins Spheniscus demersus at Mercury Island,Namibia, between 1996 and 2009

The diet of African penguins Spheniscus demersus in Namibia consisted mainly of sardine Sardinops sagax in the 1950s. Since the collapse of pelagic fish stocks in the 1970s, birds fed mainly on bearded (pelagic) goby Sufflogobius bibarbatus, a low-energy prey species. We present diet data for African penguins breeding at Mercury Island, the largest colony for this species in Namibia, between 1996 and 2009. Bearded goby was the main prey item throughout the study period, both in terms of frequency of occurrence (67.8%; SD 31.2) and in terms of mass (59.2%; SD 31.5). Diet composition varied throughout the year as well as between years; birds occasionally fed on a variety of fish species other than bearded goby. In Namibia, poor prey abundance is considered as a major factor contributing to the decline of penguin numbers after the collapse of the sardine stocks. However, bearded goby appears to be relatively abundant along Namibia's southern coast and low prey quality rather than low abundance appears to be a key factor influencing population dynamics of African penguins and other marine top predators in southern Namibia.     

Are African penguins worth saving? The ecotourism value of the Boulders Beach colony

Populations of the African penguin Spheniscus demersus have decreased dramatically over the past century, due in part to competition for food with commercial fisheries, and the species is now endangered as a result. Economic arguments are used to favour fisheries over the needs of penguins, but penguins have direct value to the South African economy thanks to penguin-based tourism at several breeding colonies. We estimated the value of African penguins at the most important tourist site for African penguins: Boulders on the Cape Peninsula, South Africa. As a mainland colony close to Cape Town, Boulders is accessible to large numbers of visitors; gate revenues in 2009/2010 alone were R14.5 million (US$2 million). A zonal travel-cost analysis revealed an average consumer surplus among Cape Town residents of some R20 per visit. Penguin-based tourism thus forms an integral part of the R25 billion Western Cape tourism sector. Given that the future survival of this valuable resource is dependent, among others, on the availability of sufficient prey, competition with South Africa's purse-seine fishing fleet should be limited through management strategies that lessen the potential effects of fishing on prey available at the local scale.     

A distance meter for large swimming marine animals

An electronic distance meter for marine animals is described. The device was tested on jackass penguins Spheniscus demersus, but could easily be modified for use on a variety of animals.     

Diving behaviour of African penguins: do they differ from other Spheniscus penguins?

African penguins Spheniscus demersus closely resemble Magellanic S. magellanicus and Humboldt S. humboldti penguins and have similar breeding and feeding ecologies. Adults feed on pelagic schooling fish in continental shelf waters, but African penguins have been reported to have shallower dive angles and remain submerged longer for dives to a given depth than their congeners. The few data for African penguins were gathered using relatively large time-depth recorders. We measured diving behaviour of 36 African penguins provisioning small chicks at three colonies near Cape Town, South Africa. Maximum and mean dive depths were 69m and 14m respectively. Diving took place mainly during the day. Although dive depths differed between colonies, there were no significant differences in dive duration or maximum, median or mean depth. Total dive duration, descent time, bottom time, ascent time and dive angle all were strongly correlated with the maximum depth attained. The diving behaviour of African penguins is similar to that of its congeners. Diving performance probably was compromised by the data-logger used in the previous study. Comparative data from Humboldt penguins also indicate potential biases in an earlier study of this species. Care is needed when comparing the diving performance of penguins measured using different loggers.     

Initial impact of the Treasure oil spill on seabirds off western South Africa

On 23 June 2000, the bulk ore carrier MV Treasure sank off western South Africa between Dassen and Robben islands, which individually currently support the largest and 3^rd largest colonies of African penguins Spheniscus demersus. Subsequently, more than 19 000 penguins were oiled, almost twice the previous highest number of seabirds oiled during a single event in southern Africa (10 000 penguins after the sinking of the Apollo Sea in June 1994). About 19 000 oiled penguins were collected for cleaning and care and about 150 oiled adults died in the wild. Some 19 500 unoiled penguins were caught at Dassen and Robben islands and relocated to Port Elizabeth, 800 km to the east, to remove them from waters affected by the oil. Of all penguins caught, which amounted to 20% of the total species population, less than 2 000 died within the first month, considerably less than in the Apollo Sea spill. This can be attributed to improved transport of penguins and the rapid arrival at rescue centres of experts able to administer emergency care. However, resources were severely extended and mortality would probably have been considerably higher had large numbers of birds not been removed from the area affected by the oil. Many relocated birds returned to their home islands within a month of being released, but considerable disruption of pair bonds is expected to result from mortality, different periods in captivity and disruption of moult cycles. This is likely to result in decreased breeding success. Recruitment to colonies will also be reduced by substantial loss of chicks and eggs. Although more than 3 000 orphaned chicks were collected for captive rearing, an estimated 4 000 died at the islands before they could be rescued. Up to 20% of bank cormorants Phalacrocorax neglectus at Robben Island, the 3^rd largest colony of the species in South Africa, died. There was low success in catching oiled cormorants and in saving those that were caught. Of 53 grown birds of four species of cormorant that were oiled and caught, only 17 survived. Captive rearing of bank cormorant chicks, which it was feared may have been orphaned, proved more successful. Spilt oil had minor impact on gulls, terns and shorebirds in the region.     

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 and 1998. 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 birds had 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.     

Impact of flipper-banding on breeding success of African penguins Spheniscus demersus at Robben Island: comparisons among silicone rubber bands,stainless-steel bands and no bands

From 2001 to 2006, two new designs of flipper bands made from silicone rubbers were tested on African penguins Spheniscus demersus at 365 nests on Robben Island, South Africa. We compared, over six years, the breeding success, from hatching to fledging, of three different groups of penguins: those with rubber bands (117 nests), with conventional stainless-steel bands (103 nests) and without bands (145 nests). There were no significant differences in breeding success between the three groups, suggesting that neither the currently used steel bands, nor either of the new rubber-band designs, were harmful during the seasons investigated. The rubber bands caused less wear of feathers and less drag on a model penguin than the steel bands. In captivity, the behaviours of African penguins fitted with rubber bands were not noticeably different to those of unbanded birds.     

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 20^th 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 20^th 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 21^st century is tenuous.     

Regionally coherent trends in colonies of African penguins Spheniscus demersus in the Western Cape,South Africa, 1987–2005

From 1987 to 2005, numbers of African penguins Spheniscus demersus breeding in South Africa's Western Cape Province increased by about 50%. Numbers decreased at the four northernmost colonies in the region: Lambert's Bay and the three colonies in Saldanha Bay, although at Jutten Island the decrease is inferred from an estimate for 1987, derived from interpolation. Numbers also decreased at Geyser Rock and Dyer Island on the South Coast. At five colonies between Saldanha Bay and Dyer Island there were large increases. At a sixth colony in that region, Seal Island, where Cape fur seals Arctocephalus pusillus pusillus limit breeding space, numbers remained stable. At two colonies that were initiated in the early 1980s, Robben Island and Boulders, increases were initially rapid (>20% per annum) and matched growth of the South African stock of sardine Sardinops sagax. Strong growth at Dassen and Vondeling islands, between Robben Island and Saldanha Bay, was observed from about 1996–2002, when there was a large increase in the biomass of pelagic fish off South Africa. Increases at colonies between Saldanha Bay and Boulders slowed after 2002, whereas the colony at Dyer Island stabilised at that time. In 2003, a new colony was initiated east of Dyer Island at De Hoop Nature Reserve. These latter trends followed an eastward shift in the distribution of sardine. Small penguin colonies may act as foci for growth in a period when the distribution of prey is changing. Hence, it is important that they be maintained, especially those that, if lost, would increase the isolation of regional populations. Some of the small colonies are less susceptible to oil spills than colonies in the proximity of harbours, and for that reason also are important.     

Restoration of oiled African penguins Spheniscus demersus a decade after the Apollo Sea spill

The bulk ore carrier Apollo Sea sank south-west of Dassen Island off western South Africa in June 1994, oiling approximately 10 000 African penguins Spheniscus demersus, most of which were collected from Dassen Island. A total of 4 076 de-oiled penguins was released with flipper bands. From 1994 to 2005, follow-up research using re-sighting and capture-mark-recapture methods indicated that about 73% of the de-oiled penguins observed back at Dassen Island attempted to breed, and were thus successfully restored into the breeding population. For de-oiled breeders, the median interval between their first recorded sighting and first recorded breeding attempt was 11 months, indicating a short-term delay in restoration. At least 45% of the de-oiled breeders were still being re-sighted five years after their release, and a minimum of 4% survived into their ninth year. These results represent the most successful restoration estimates anywhere in the world. The proportion of de-oiled juvenile penguins re-sighted back at Dassen Island and recorded breeding was lower than that of birds in adult plumage. De-oiled non-breeders spent significantly more time along the shore and less time within breeding colonies than de-oiled breeders. The mean proportion of de-oiled breeders that abstained from breeding each year during the study period was greater than expected. There was a negative relationship between breeding and subsequent survival and breeding, suggesting a cost of reproduction for de-oiled birds.     

Breeding success of African penguins Spheniscus demersus at Dassen Island,especially after oiling following the Apollo Sea spill

The reproductive success of African penguins Spheniscus demersus at Dassen Island from 1994 to 2000 was variable, but much higher than previously reported figures for the species. Breeding success was positively related to the abundance of anchovy Engraulis encrasicolus and sardine Sardinops sagax, and the high reproductive output during the study was attributed to the large biomass and high availability of these two species for much of the study period. De-oiled penguins from the Apollo Sea spill had a slightly lower hatching success but a significantly lower overall breeding success than that of un-oiled birds, driven mainly by the reduced number of chicks which fledged. Nests with two de-oiled Apollo Sea parents were less successful than nests with one de-oiled bird. There was increased mortality of chicks 40 days and older in nests with de-oiled birds. Chicks from nests with one de-oiled Apollo Sea parent grew at a similar rate to chicks from nests with no de-oiled parents. However, chicks from nests that comprised two de-oiled Apollo Sea birds had significantly slower growth rates than these other two groups. Breeding success and chick growth at nests with de-oiled birds were more negatively impacted when feeding conditions were less favourable. These results suggest that one of the main reasons for lower breeding success in de-oiled birds was their reduced ability to provision chicks, especially during the period in which the energy demands of the chicks is greatest. The rate of mate fidelity was lower in de-oiled birds than un-oiled birds, and there was a positive association between mate fidelity and breeding success.     

Partitioning of a goby resource amongst four avian predators and evidence for altered trophic flow in the pelagic community of an intense,perennial upwelling system

Pelagic gobies Sufflogobius bibarbatus were numerically the most important prey of jackass penguins Spheniscus demersus. Cape cormorants Phalacrocorax capensis and bank cormorants P. neglectus sampled at islands off the South West African coast during the period 1978–1982. These three seabirds feed on small gobies near the surface and some of them can also dive sufficiently deep to catch larger gobies. Their populations at islands north of Lüderitz, where gobies are abundant, have been increasing. Cape gannets Morus capensis feed only on large gobies that are infrequently at the surface, and the gannet population off South West Africa has shown a large decrease since the collapse of the pilchard Sardinops ocellata resource in the late 1960s and early 1970s. In the intense perennial upwelling system situated between 22 and 27°S gobies are believed to have partially replaced pilchards during the 1970s. Both pilchards and gobies are able to feed on large diatoms of the genera Chaetoceros and Delphineis, which dominated the inshore phytoplankton in the early 1970s when biomass levels of both pilchards and gobies were low.     

Congruent,decreasing trends of gentoo penguins and Crozet shags at sub-Antarctic Marion Island suggest food limitation through common environmental forcing

Numbers of gentoo penguins Pygoscelis papua and Crozet shags Phalacrocorax [atriceps] melanogenis breeding annually at Marion Island, one of South Africa's Prince Edward Islands in the South-West Indian Ocean, were strongly correlated over 19 split-years from 1994/1995 to 2012/2013. Both species decreased between the mid-1990s and the mid-2000s, exhibited a partial recovery in the late-2000s and then decreased to their lowest recorded levels in 2012/2013. In both instances, the partial recoveries in the late-2000s were associated with improved breeding success. At a colony of gentoo penguins, breeding success was negatively correlated with the date of arrival of adults to breed. Gentoo penguins and Crozet shags are demersal feeders in inshore waters around Marion Island and there is considerable overlap in the composition of their diets. Therefore, trends in their populations may be driven by food availability, which is likely to be influenced by benthic production around the island. We propose that, in South Africa, and based on the current assessment, the Crozet shag, which elsewhere breeds only at the Crozet Islands, is now Critically Endangered, and the more widely ranging gentoo penguin is Endangered.     

Populations of Surface-nesting Seabirds at Marion Island, 1994/95–2002/03

During the 1990s and early 2000s, populations of surface-nesting seabirds at Marion Island showed different trends, but for the majority of species numbers decreased. Reduced numbers of gentoo penguins Pygoscelis papua, eastern rockhopper penguins Eudyptes chrysocome filholi, Crozet shags Phalacrocorax [atriceps] melanogenis and probably macaroni penguins E. chrysolophus are most plausibly attributed to an altered availability of food. Decreases in numbers of dark-mantled sooty albatrosses Phoebetria fusca, light-mantled sooty albatrosses P. palpebrata, southern giant petrels Macronectes giganteus and possibly northern giant petrels M. halli may have resulted from mortality of birds in longline fisheries. However, populations of wandering Diomedea exulans and grey-headed Thalassarche chrysostoma albatrosses fluctuated around a stable level. Numbers of Subantarctic skuas Catharacta antarctica and kelp gulls Larus dominicanus breeding at Marion Island also decreased. Kerguelen Sterna virgata and Antarctic S. vittata terns remain scarce at the island. Trends for king penguins Aptenodytes patagonicus were not reliably gauged, but numbers probably remained stable or increased. There were large fluctuations in numbers of king penguin chicks surviving to the end of winter.     

Recent trends in numbers of four species of penguins at the Prince Edward Islands

Four species of penguin breed regularly at South Africa's Prince Edward Islands: king penguin Aptenodytes patagonicus, gentoo penguin Pygoscelis papua, macaroni penguin Eudyptes chrysolophus and southern rockhopper penguin E. chrysocome. In December 2008, it was estimated that some 65 000 pairs of king penguins were incubating eggs at Marion Island, the larger of the two islands in the group, and 2 000 pairs at Prince Edward Island. At Marion Island from 1987 to 2008, there was no long-term trend in numbers of king penguin chicks that survived to the end of the winter period, but there was considerable fluctuation in chick production in the 1990s. It was roughly estimated that on average 88% of king penguin chicks survived the winter period (from April to September/October). Numbers of gentoo penguins at Marion Island decreased from more than 1 300 pairs in the mid-1990s to fewer than 800 pairs in 2003, and then increased to almost 1 100 pairs in 2008 as breeding success improved. Between 1994/1995 and 2008/2009, numbers of macaroni and southern rockhopper penguins at Marion Island decreased by about 30% and 70% respectively. In 2008/2009, some 290 000 pairs of macaroni penguins bred at this island, mostly in two large colonies where there was a progressive decrease in the density of nests. At both these colonies, decreases in numbers breeding followed outbreaks of disease. Inadequate breeding success has influenced the decreases of macaroni and rockhopper penguins. In 2008/2009, some 42 000 pairs of southern rockhopper penguins bred at Marion Island and 12 000 pairs of macaroni penguins and 38 000 pairs of southern rockhopper penguins at Prince Edward Island.     

Fine resolution 3D temperature fields off Kerguelen from instrumented penguins

The use of diving animals as autonomous vectors of oceanographic instruments is rapidly increasing, because this approach yields cost-efficient new information and can be used in previously poorly sampled areas. However, methods for analyzing the collected data are still under development. In particular, difficulties may arise from the heterogeneous data distribution linked to animals’ behavior. Here we show how raw temperature data collected by penguin-borne loggers were transformed to a regular gridded dataset that provided new information on the local circulation off Kerguelen. A total of 16 king penguins (Aptenodytes patagonicus) were equipped with satellite-positioning transmitters and with temperature–time–depth recorders (TTDRs) to record dive depth and sea temperature. The penguins’ foraging trips recorded during five summers ranged from 140 to 600 km from the colony and 11,000 dives >100 m were recorded. Temperature measurements recorded during diving were used to produce detailed 3D temperature fields of the area (0–200 m). The data treatment included dive location, determination of the vertical profile for each dive, averaging and gridding of those profiles onto 0.1°×0.1° cells, and optimal interpolation in both the horizontal and vertical using an objective analysis. Horizontal fields of temperature at the surface and 100 m are presented, as well as a vertical section along the main foraging direction of the penguins. Compared to conventional temperature databases (Levitus World Ocean Atlas and historical stations available in the area), the 3D temperature fields collected from penguins are extremely finely resolved, by one order finer. Although TTDRs were less accurate than conventional instruments, such a high spatial resolution of penguin-derived data provided unprecedented detailed information on the upper level circulation pattern east of Kerguelen, as well as the iron-enrichment mechanism leading to a high primary production over the Kerguelen Plateau.     

Counts of Surface-nesting Seabirds Breeding at Prince Edward Island,Summer 2001/02

The first midsummer survey of surface-nesting seabirds at Prince Edward Island was conducted during December 2001. It was also the first census for most species since the early 1980s. Despite concerns about the impacts of longline fishing mortality on various albatrosses and giant petrels, there was no evidence that populations of these species had decreased. Indeed, the estimate of wandering albatrosses Diomedea exulans (Vulnerable) has increased significantly, making Prince Edward Island equal with Marion Island as supporting the largest single-island populations of this species. Species that underwent significant decreases were macaroni penguins Eudyptes chrysolophus (Vulnerable), Crozet shags Phalacrocorax [atriceps] melanogenis and Kerguelen terns Sterna virgata (Near-Threatened). The reasons for these decreases are unclear, but for macaroni penguins may be partly a consequence of competition for space with the burgeoning population of fur seals Arctocephalus spp. The 2001 survey increased the population estimates for Subantarctic skuas Catharacta antarctica, lightmantled sooty albatrosses Phoebetria palpebrata (Vulnerable) and southern giant petrels Macronectes giganteus (Vulnerable), mostly as a result of greater coverage than for previous counts. The 2001 survey confirms that Prince Edward Island remains a globally important breeding site for seabirds.     

St Helena Bay (southern Benguela) then and now: muted climate signals,large human impact

The development of suitable reference states for ecosystem-based management requires documentation of changes in structure and functioning of marine ecosystems, including assessment of the relative importance of bottom-up and top-down processes as drivers of change. We used monitoring data available from St Helena Bay, the most productive bay and an important nursery area situated on the west coast of South Africa, during 1950–2010 to reveal changes in the abiotic and biotic components. St Helena Bay in the 1950s showed similarities to 2000–2010 in terms of wind patterns, hydrology and phytoplankton. Upwelling, oxygen and nutrient concentrations in subthermocline water displayed pronounced decadal-scale variability. Primary production in St Helena Bay is variable, but consistently higher than that on the adjacent Namaqua shelf. Zooplankton size composition and biomass in August have changed markedly since the 1950s. During 2001–2010, mesozooplankton biomass in autumn was considerably lower than in summer, probably due to predation by small pelagic fish. Pelagic fish catch patterns and distributions have altered dramatically. Conservation measures, implemented to reverse past negative human impact, have benefitted marine mammals, the abundance of which has increased in the area, but additional conservation measures are necessary to reverse the decline in African penguins Spheniscus demersus. St Helena Bay shows a muted response to long-term change in the southern Benguela, with marked decadal variability but no clear long-term trend in oceanography and biogeochemistry. Changes in ecosystem boundary conditions and fishing pressure cannot be ignored as important drivers of change in the southern Benguela since the 1950s.     

Seabirds in the diet of Cape fur seals Arctocephalus pusillus pusillus at three mainland breeding colonies in Namibia

Predation by Cape fur seals Arctocephalus pusillus pusillus on seabirds has previously been attributed to a few individuals, mainly males. Scat samples were collected at three mainland breeding colonies of seals in Namibia to determine the extent of seabird predation. Mainly females were sampled. No scats from the van Reenen Bay and Atlas-Wolf Bay colonies, and only two from Cape Cross, contained feather remains, giving an overall frequency of occurrence of feathers in scats of 0.1%. The feathers found were of African penguin Spheniscus demersus and a species of cormorant (Phalacrocorax sp.).