Tracing beach litter sources: Drink lids tell a different story from their bottles.

Water and soft drink bottles made from polyethylene terephthalate (PET) sink at sea unless they contain trapped air, whereas their lids are made from polymers that float and can drift long distances. We sampled loose lids and bottles at 21 South African beaches to compare their origins. The proportions of foreign-made bottles and lids were correlated, and increased away from urban centres, indicating that much land-based litter strands close to source areas. Over 80 % of foreign-made drink bottles and 90 % of lids came from Asia, but most bottles were manufactured in China, Malaysia-Singapore and the UAE, and were dumped from ships. By comparison, most loose lids were in poor condition after being carried across the Indian Ocean from Indonesia by the South Equatorial Current. Reducing PET drink bottles at sea requires enforcement of regulations banning dumping at sea, whereas reducing their lids requires better control of littering in source countries.

Effect of Environmental Variables on African Penguin Vocal Activity: Implications for Acoustic Censusing.

Global biodiversity is in rapid decline, and many seabird species have disproportionally poorer conservation statuses than terrestrial birds. A good understanding of population dynamics is necessary for successful conservation efforts, making noninvasive, cost-effective monitoring tools essential. Here, we set out to investigate whether passive acoustic monitoring (PAM) could be used to estimate the number of animals within a set area of an African penguin (Spheniscus demersus) colony in South Africa. We were able to automate the detection of ecstatic display songs (EDSs) in our recordings, thus facilitating the handling of large datasets. This allowed us to show that calling rate increased with wind speed and humidity but decreased with temperature, and to highlight apparent abundance variations between nesting habitat types. We then showed that the number of EDSs in our recordings positively correlated with the number of callers counted during visual observations, indicating that the density could be estimated based on calling rate. Our observations suggest that increasing temperatures may adversely impact penguin calling behaviour, with potential negative consequences for population dynamics, suggesting the importance of effective conservation measures. Crucially, this study shows that PAM could be successfully used to monitor this endangered species' populations with minimal disturbance.

Seabird morphology determines operational wind speeds, tolerable maxima, and responses to extremes.

Storms can cause widespread seabird stranding and wrecking,1,2,3,4,5 yet little is known about the maximum wind speeds that birds are able to tolerate or the conditions they avoid. We analyzed >300,000 h of tracking data from 18 seabird species, including flapping and soaring fliers, to assess how flight morphology affects wind selectivity, both at fine scales (hourly movement steps) and across the breeding season. We found no general preference or avoidance of particular wind speeds within foraging tracks. This suggests seabird flight morphology is adapted to a "wind niche," with higher wing loading being selected in windier environments. In support of this, wing loading was positively related to the median wind speeds on the breeding grounds, as well as the maximum wind speeds in which birds flew. Yet globally, the highest wind speeds occur in the tropics (in association with tropical cyclones) where birds are morphologically adapted to low median wind speeds. Tropical species must therefore show behavioral responses to extreme winds, including long-range avoidance of wind speeds that can be twice their operable maxima. By contrast, Procellariiformes flew in almost all wind speeds they encountered at a seasonal scale. Despite this, we describe a small number of cases where albatrosses avoided strong winds at close range, including by flying into the eye of the storm. Extreme winds appear to pose context-dependent risks to seabirds, and more information is needed on the factors that determine the hierarchy of risk, given the impact of global change on storm intensity.6,7.

Maritime traffic trends around the southern tip of Africa - Did marine noise pollution contribute to the local penguins' collapse?

The rapid increase in seaborn trade since the 1990s has resulted in an increase in vessel-derived noise pollution, yet there is little evidence linking these activities to a decline in many marine taxa, such as seabirds. Algoa Bay, South Africa, is a marine biodiversity hotspot, providing habitats for the largest populations of endangered African Penguins (Spheniscus demersus), as well as other endangered seabirds, cetaceans and seals. The bay is situated on a major shipping route and since 2016 has hosted the first offshore ship-to-ship (STS) bunkering operations in the country, i.e. the supplying of fuel from one ship to another outside of harbours. Using Automatic Identification System (AIS) data, we estimated noise emissions from vessels as a proxy for underwater ambient noise levels within the core penguin utilisation area. Frequency of vessels using the bay doubled during our study, with numbers of bulk carriers increasing ten-fold. Ambient underwater noise levels were generally high in the bay (ca 140 dB re 1 µPa since 2015) but significantly increased by 2 dB SPL after the initiation of STS bunkering in 2016, corresponding to double the underwater noise intensity. This increase coincided with a significant and dramatic decline by 85% in penguin numbers from St Croix Island since 2016. Algoa Bay is now one of the noisiest bays in the world. This is the first study to assess the potential impact of vessel-derived underwater noise levels on a seabird population. Penguins, like marine mammal species, are known to be sensitive to marine noise pollution and urgent management interventions are required to mitigate this recent disturbance, to preserve the remaining stronghold of the African penguin and the marine mammals' populations sharing the penguins' habitat.

Important marine areas for endangered African penguins before and after the crucial stage of moulting.

The population of the Endangered African penguin Spheniscus demersus has decreased by > 65% in the last 20 years. A major driver of this decrease has been the reduced availability of their principal prey, sardine Sardinops sagax and anchovy Engraulis encrasicolus. To date, conservation efforts to improve prey availability have focused on spatial management strategies to reduce resource competition with purse-seine fisheries during the breeding season. However, penguins also undergo an annual catastrophic moult when they are unable to feed for several weeks. Before moulting they must accumulate sufficient energy stores to survive this critical life-history stage. Using GPS tracking data collected between 2012 and 2019, we identify important foraging areas for pre- and post-moult African penguins at three of their major colonies in South Africa: Dassen Island and Stony Point (Western Cape) and Bird Island (Eastern Cape). The foraging ranges of pre- and post-moult adult African penguins (c. 600 km from colony) was far greater than that previously observed for breeding penguins (c. 50 km from colony) and varied considerably between sites, years and pre- and post-moult stages. Despite their more extensive range during the non-breeding season, waters within 20 and 50 km of their breeding colonies were used intensively and represent important foraging areas to pre- and post-moult penguins. Furthermore, penguins in the Western Cape travelled significantly further than those in the Eastern Cape which is likely a reflection of the poor prey availability along the west coast of South Africa. Our findings identify important marine areas for pre- and post-moult African penguins and support for the expansion of fisheries-related spatio-temporal management strategies to help conserve African penguins outside the breeding season.

Seascapes of fear and competition shape regional seabird movement ecology.

Fear effects of predators on prey distributions are seldom considered in marine environments, especially over large spatial scales and in conservation contexts. To fill these major gaps, we tested the Seascape of Fear Hypothesis in the Benguela marine ecosystem off South Africa. Using electronic tracking data, we showed that Cape gannets and their predator, the Cape fur seal, co-occurred in daytime and competed with fisheries within coastal areas. At night, gannets are particularly vulnerable to seals, and 28% of the birds returned to the safety of their breeding colony. The remaining 72% slept at the sea surface, but shifted to offshore areas with lower seal attendance, reducing predation risk by 25%. Overall, our integrative study demonstrates how fear and competition shape the seascape of threatened Cape gannets within a marine environment perturbed by climate change and overfishing. Such knowledge has strong implications for the design of marine protected areas.

Urofaecal glucocorticoid metabolite concentrations in African penguin (Spheniscus demersus) chick populations experiencing different levels of human disturbance.

Despite the importance of ecotourism in species conservation, little is known about the industry's effects on wildlife. In South Africa, some African penguin (Spheniscus demersus) colonies have become tourist attractions. The species is globally endangered, with population sizes decreasing over the past 40 years. As African penguin chicks are altricial and unable to move away from anthropogenic stressors, it is important to evaluate the effect of tourist activities on baseline glucocorticoid levels as a measure of potential disturbance. Chicks at three study sites within two breeding colonies (Robben Island, Stony Point), with varying levels of exposure to tourism (low/moderate/high) were monitored. Urofaecal samples were collected to determine urofaecal glucocorticoid metabolite (ufGCM) concentrations as an indication of baseline stress physiology. Morphometric measurements were taken to compare body condition between sites. Penguin chicks experiencing low, infrequent human presence had significantly higher mean (± standard deviation) ufGCM levels [1.34 ± 1.70 µg/g dry weight (DW)] compared to chicks experiencing both medium (0.50 ± 0.40 µg/g DW, P = 0.001) and high levels of human presence (0.57 ± 0.47 µg/g DW, P = 0.003). There was no difference in chick body condition across sites. These results suggest that exposure to frequent human activity may induce habituation/desensitization in African penguin chicks. Acute, infrequent human presence was likely an important driver for comparatively higher ufGCM levels in chicks, though several other environmental stressors may also play an important role in driving adrenocortical activity. Nevertheless, as unhabituated chicks experiencing infrequent anthropogenic presence showed significantly higher ufGCM levels, managers and legislation should attempt to minimize all forms of activity around important breeding colonies that are not already exposed to regular tourism. Although the results of this study are crucial for developing enhanced conservation and management protocols, additional research on the long-term effect of anthropogenic activities on African penguin physiology is required.

The first baseline estimation of marine litter in Port Elizabeth, South Africa.

Data on marine litter is crucial to guide waste management but is scarce in third-world countries such as South Africa. We established the first baseline measurement of litter accumulation on two beaches differing in public access in Port Elizabeth, Eastern Cape, the poorest province in South Africa. Four 10-day surveys were conducted on each beach between June 2019 and June 2020. Results revealed that most of the litter (95.7%) was of South African origin and likely came from local land-based sources (e.g., beachgoers or a local river). Daily accumulation rates at the study sites ranged between 24.58 and 86.54 items·100 m-1·day-1, an order of magnitude lower than rates from other cities in Africa. However, like elsewhere, plastic (including foam) made up the bulk (74.2%) of litter with food packaging contributing a large portion of this. These findings indicate that local interventions are warranted to reduce litter in Port Elizabeth.

Foraging movements of breeding Kelp Gulls in South Africa.

BACKGROUND: Kelp Gulls Larus dominicanus are one of the most abundant gulls in the Southern Hemisphere and can play an important role in their ecosystem. Understanding their foraging ecology is therefore important, especially in the context of anthropogenic changes of the environment. Over 35,000 Kelp Gulls breed in South Africa but little is known about their habitat use. It has been hypothesised that foraging mainly occurs in natural habitats while provisioning chicks to ensure high quality food, but knowledge on their foraging ecology during the incubation period remains poor. METHODS: We tracked incubating Kelp Gulls from six colonies distributed along the coast of South Africa, varying in their distance to urban areas and landfills, and compared foraging trip patterns and habitat selection between colonies. RESULTS: Gulls from west coast colonies, generally located further from landfills than the other studied colonies, travelled farther from their breeding sites (11.7 ± 9.9-17.8 ± 21.7 km, n = 3 colonies) than birds from Cape Town and south and east coast colonies (1.7 ± 0.8-3.1 ± 3.7 km, n = 3) with birds travelling farthest when foraging at sea. Gulls from all colonies spent more time foraging in marine, coastal, and natural terrestrial environments than scavenging in strongly modified habitats while incubating. CONCLUSIONS: Our results suggest that Kelp Gulls in South Africa are able to exploit various resources from different foraging habitats, regardless of colony location and seem to rely less on anthropogenic habitats than expected.

The conservation status and population decline of the African penguin deconstructed in space and time.

Understanding changes in abundance is crucial for conservation, but population growth rates often vary over space and time. We use 40 years of count data (1979-2019) and Bayesian state-space models to assess the African penguin Spheniscus demersus population under IUCN Red List Criterion A. We deconstruct the overall decline in time and space to identify where urgent conservation action is needed. The global African penguin population met the threshold for Endangered with a high probability (97%), having declined by almost 65% since 1989. An historical low of ~17,700 pairs bred in 2019. Annual changes were faster in the South African population (-4.2%, highest posterior density interval, HPDI: -7.8 to -0.6%) than the Namibian one (-0.3%, HPDI: -3.3 to +2.6%), and since 1999 were almost -10% at South African colonies north of Cape Town. Over the 40-year period, the Eastern Cape colonies went from holding ~25% of the total penguin population to ~40% as numbers decreased more rapidly elsewhere. These changes coincided with an altered abundance and availability of the main prey of African penguins. Our results underline the dynamic nature of population declines in space as well as time and highlight which penguin colonies require urgent conservation attention.

Multi-predator assemblages, dive type, bathymetry and sex influence foraging success and efficiency in African penguins.

Marine predators adapt their hunting techniques to locate and capture prey in response to their surrounding environment. However, little is known about how certain strategies influence foraging success and efficiency. Due to the miniaturisation of animal tracking technologies, a single individual can be equipped with multiple data loggers to obtain multi-scale tracking information. With the addition of animal-borne video data loggers, it is possible to provide context-specific information for movement data obtained over the video recording periods. Through a combination of video data loggers, accelerometers, GPS and depth recorders, this study investigated the influence of habitat, sex and the presence of other predators on the foraging success and efficiency of the endangered African penguin, Spheniscus demersus, from two colonies in Algoa Bay, South Africa. Due to limitations in the battery life of video data loggers, a machine learning model was developed to detect prey captures across full foraging trips. The model was validated using prey capture signals detected in concurrently recording accelerometers and animal-borne cameras and was then applied to detect prey captures throughout the full foraging trip of each individual. Using GPS and bathymetry information to inform the position of dives, individuals were observed to perform both pelagic and benthic diving behaviour. Females were generally more successful on pelagic dives than males, suggesting a trade-off between manoeuvrability and physiological diving capacity. By contrast, males were more successful in benthic dives, at least for Bird Island (BI) birds, possibly due to their larger size compared to females, allowing them to exploit habitat deeper and for longer durations. Both males at BI and both sexes at St Croix (SC) exhibited similar benthic success rates. This may be due to the comparatively shallower seafloor around SC, which could increase the likelihood of females capturing prey on benthic dives. Observation of camera data indicated individuals regularly foraged with a range of other predators including penguins and other seabirds, predatory fish (sharks and tuna) and whales. The presence of other seabirds increased individual foraging success, while predatory fish reduced it, indicating competitive exclusion by larger heterospecifics. This study highlights novel benthic foraging strategies in African penguins and suggests that individuals could buffer the effects of changes to prey availability in response to climate change. Furthermore, although group foraging was prevalent in the present study, its influence on foraging success depends largely on the type of heterospecifics present.

Radar detectors carried by Cape gannets reveal surprisingly few fishing vessel encounters.

Fisheries compete with seabirds for vanishing marine resources, but also produce fishery waste consumed by seabirds. Marine birds may therefore avoid or seek fishing vessels, and have evolved complex, plastic behavioural responses to vessel presence. Understanding these responses is essential to the conservation of a globally declining seabird community. We studied Cape gannets (Morus capensis), which compete with fisheries for reduced sardine (Sardinops sagax) resources in the Benguela upwelling region off South Africa. Using bird-borne GPS trackers coupled with newly-developed ship-radar detectors we show that foraging gannets seldom attended fishing vessels. Rather, they switched from eating scarce sardines or energetically-poor fishery waste to targeting locally abundant saury (Scomberesox saurus). This pelagic fish is brought into the seascape by warm water influx, and is not commercially exploited by fisheries. Cape gannets thereby show dietary plasticity, allowing them to maintain adult body condition and chick growth rates. This diet switch is a strong indicator that Cape gannets forage in an ecologically perturbed marine environment.

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.

Avoidance of seismic survey activities by penguins.

Seismic surveys in search for oil or gas under the seabed, produce the most intense man-made ocean noise with known impacts on invertebrates, fish and marine mammals. No evidence to date exists, however, about potential impacts on seabirds. Penguins may be expected to be particularly affected by loud underwater sounds, due to their largely aquatic existence. This study investigated the behavioural response of breeding endangered African Penguins Spheniscus demersus to seismic surveys within 100 km of their colony in South Africa, using a multi-year GPS tracking dataset. Penguins showed a strong avoidance of their preferred foraging areas during seismic activities, foraging significantly further from the survey vessel when in operation, while increasing their overall foraging effort. The birds reverted to normal behaviour when the operation ceased, although longer-term repercussions on hearing capacities cannot be precluded. The rapid industrialization of the oceans has increased levels of underwater anthropogenic noises globally, a growing concern for a wide range of taxa, now also including seabirds. African penguin numbers have decreased by 70% in the last 10 years, a strong motivation for precautionary management decisions, including the exclusion of seismic exploratory activities within at least 100 km of their breeding colonies.

Group foraging increases foraging efficiency in a piscivorous diver, the African penguin.

Marine piscivores have evolved a variety of morphological and behavioural adaptations, including group foraging, to optimize foraging efficiency when targeting shoaling fish. For penguins that are known to associate at sea and feed on these prey resources, there is nonetheless a lack of empirical evidence to support improved foraging efficiency when foraging with conspecifics. We examined the hunting strategies and foraging performance of breeding African penguins equipped with animal-borne video recorders. Individuals pursued both solitary as well as schooling pelagic fish, and demonstrated independent as well as group foraging behaviour. The most profitable foraging involved herding of fish schools upwards during the ascent phase of a dive where most catches constituted depolarized fish. Catch-per-unit-effort was significantly improved when targeting fish schools as opposed to single fish, especially when foraging in groups. In contrast to more generalist penguin species, African penguins appear to have evolved specialist hunting strategies closely linked to their primary reliance on schooling pelagic fish. The specialist nature of the observed hunting strategies further limits the survival potential of this species if Allee effects reduce group size-related foraging efficiency. This is likely to be exacerbated by diminishing fish stocks due to resource competition and environmental change.

Recreational Fish-Finders--An Inexpensive Alternative to Scientific Echo-Sounders for Unravelling the Links between Marine Top Predators and Their Prey.

Studies investigating how mobile marine predators respond to their prey are limited due to the challenging nature of the environment. While marine top predators are increasingly easy to study thanks to developments in bio-logging technology, typically there is scant information on the distribution and abundance of their prey, largely due to the specialised nature of acquiring this information. We explore the potential of using single-beam recreational fish-finders (RFF) to quantify relative forage fish abundance and draw inferences of the prey distribution at a fine spatial scale. We compared fish school characteristics as inferred from the RFF with that of a calibrated scientific split-beam echo-sounder (SES) by simultaneously operating both systems from the same vessel in Algoa Bay, South Africa. Customized open-source software was developed to extract fish school information from the echo returns of the RFF. For schools insonified by both systems, there was close correspondence between estimates of mean school depth (R2 = 0.98) and school area (R2 = 0.70). Estimates of relative school density (mean volume backscattering strength; Sv) measured by the RFF were negatively biased through saturation of this system given its smaller dynamic range. A correction factor applied to the RFF-derived density estimates improved the comparability between the two systems. Relative abundance estimates using all schools from both systems were congruent at scales from 0.5 km to 18 km with a strong positive linear trend in model fit estimates with increasing scale. Although absolute estimates of fish abundance cannot be derived from these systems, they are effective at describing prey school characteristics and have good potential for mapping forage fish distribution and relative abundance. Using such relatively inexpensive systems could greatly enhance our understanding of predator-prey interactions.

Penguins are attracted to dimethyl sulphide at sea.

Breeding Spheniscus penguins are central place foragers that feed primarily on schooling pelagic fish. They are visual hunters, but it is unclear how they locate prey patches on a coarse scale. Many petrels and storm petrels (Procellariiformes), the penguins' closest relatives, use olfactory cues to locate prey concentrations at sea, but this has not been demonstrated for penguins. Procellariiforms are attracted to a variety of olfactory cues, including dimethyl sulphide (DMS), an organosulphur compound released when phytoplankton is grazed, as well as fish odorants such as cod liver oil. A recent study found that African penguins Spheniscus demersus react to DMS on land. We confirm this result and show that African penguins are also attracted by DMS at sea. DMS-scented oil slicks attracted 2-3 times more penguins than control slicks, whereas penguins showed no response to slicks containing cod liver oil. The number of penguins attracted to DMS increased for at least 30 min, suggesting penguins could travel up to 2 km to reach scent cues. Repeats of land-based trials confirmed previous results showing DMS sensitivity of penguins on land. Our results also support the hypothesis that African penguins use DMS as an olfactory cue to locate prey patches at sea from a distance, which is particularly important given their slow commuting speed relative to that of flying seabirds.

A junk-food hypothesis for gannets feeding on fishery waste.

Worldwide fisheries generate large volumes of fishery waste and it is often assumed that this additional food is beneficial to populations of marine top-predators. We challenge this concept via a detailed study of foraging Cape gannets Morus capensis and of their feeding environment in the Benguela upwelling zone. The natural prey of Cape gannets (pelagic fishes) is depleted and birds now feed extensively on fishery wastes. These are beneficial to non-breeding birds, which show reduced feeding effort and high survival. By contrast, breeding gannets double their diving effort in an attempt to provision their chicks predominantly with high-quality, live pelagic fishes. Owing to a scarcity of this resource, they fail and most chicks die. Our study supports the junk-food hypothesis for Cape gannets since it shows that non-breeding birds can survive when complementing their diet with fishery wastes, but that they struggle to reproduce if live prey is scarce. This is due to the negative impact of low-quality fishery wastes on the growth patterns of gannet chicks. Marine management policies should not assume that fishery waste is generally beneficial to scavenging seabirds and that an abundance of this artificial resource will automatically inflate their populations.