Haemoproteus jenniae (Haemoproteidae, Haemosporida) infects gulls (Larus spp.) in South Africa, with redescription of Haemoproteus skuae.

Haemoproteus spp. are dipteran-borne protozoa that infect erythrocytes and reticulo-endothelial cells of birds. These parasites are not usually transmitted between birds belonging to different orders. The suborder Lari (order Charadriiformes) comprises ~170 avian species, the majority of which are aquatic, including gulls, terns, auklets, murres and skuas, among others. In spite of the diversity of this avian group, there is limited known diversity of haemosporidian parasites, with only 4 recorded Haemoproteus morphospecies thus far. We examined the blood smears of 21 kelp gulls (Larus dominicanus) captured at a breeding colony in South Africa, as well as Haemoproteus-positive archival blood smears of 15 kelp gulls and 1 Hartlaub's gull (Larus hartlaubii) sampled while under care at seabird rehabilitation facilities in South Africa. Haemoproteus sp. infection was detected in 19% of wild-caught kelp gulls. All parasites from the gulls were morphologically identified as Haemoproteus jenniae, a species previously recorded in Lari birds at the Galapagos Islands (Ecuador), Rocas Atoll (Brazil) and Poland. Gene sequencing uncovered a new cytochrome b lineage, LARDOM01, which was closely related to the previously reported H. jenniae lineage CREFUR01. Additionally, we evaluated a hapantotype blood smear of Haemoproteus skuae, which had been described infecting a brown skua (Catharacta antarctica) in South Africa. We provide a redescription of H. skuae and discuss the morphological characters distinguishing it from H. jenniae. Further research is necessary to improve our knowledge about the host and geographic distribution, health effects and phylogeny of H. jenniae and H. skuae.

Geographical, temporal, and individual-based differences in the trophic ecology of female Cape fur seals.

Information on resource use and trophic dynamics of marine predators is important for understanding their role in ecosystem functioning and predicting population-level responses to environmental change. Where separate populations experience different local environmental conditions, geographic variability in their foraging ecology is often expected. Within populations, individuals also vary in morphology, physiology, and experience, resulting in specialization in resource use. In this context, isotopic compositions of incrementally grown tissues such as keratinous hairs offer a valuable opportunity to study long-term variation in resource and habitat use. We investigated the trophic ecology of female Cape fur seals (Arctocephalus pusillus pusillus) using carbon and nitrogen isotopic compositions of serially sampled whiskers collected at four breeding sites along the coast of South Africa. Drawing on over 900 isotopic measurements, we assessed geographic variability in isotopic niche width between colonies and the degree of individual specialization. We found slight, but clear geographic differences in isotopic ratios and isotopic niche widths, seemingly related to ecological setting, with niche widths being proportional to the area of available shelf and shelf-slope habitat surrounding the colony. We further identified periodic oscillations in isotopic ratios, which likely reflect temporal patterns in foraging distribution and prey type, linked to shifts in the availability of prey resources and their interaction with constraints on individual females throughout their breeding cycle. Finally, individual specialization indices revealed that each of the study populations contain specialist individuals that utilize only a small subset of the total population niche width. The degree of individual specialization was, however, not consistent across colonies and may reflect an interactive influence between density-dependent effects and habitat heterogeneity. Overall, this study provides important information on the trophic ecology of Cape fur seals breeding in South Africa and highlights the need to consider geographic and individual variability when assessing the foraging ecology of marine predators.

Aquatic birds have middle ears adapted to amphibious lifestyles.

Birds exhibit wide variation in their use of aquatic environments, on a spectrum from entirely terrestrial, through amphibious, to highly aquatic. Although there are limited empirical data on hearing sensitivity of birds underwater, mounting evidence indicates that diving birds detect and respond to sound underwater, suggesting that some modifications of the ear may assist foraging or other behaviors below the surface. In air, the tympanic middle ear acts as an impedance matcher that increases sound pressure and decreases sound vibration velocity between the outside air and the inner ear. Underwater, the impedance-matching task is reversed and the ear is exposed to high hydrostatic pressures. Using micro- and nano-CT (computerized tomography) scans of bird ears in 127 species across 26 taxonomic orders, we measured a suite of morphological traits of importance to aerial and aquatic hearing to test predictions relating to impedance-matching in birds with distinct aquatic lifestyles, while accounting for allometry and phylogeny. Birds that engage in underwater pursuit and deep diving showed the greatest differences in ear structure relative to terrestrial species. In these heavily modified ears, the size of the input areas of both the tympanic membrane and the columella footplate of the middle ear were reduced. Underwater pursuit and diving birds also typically had a shorter extrastapedius, a reduced cranial air volume and connectivity and several modifications in line with reversals of low-to-high impedance-matching. The results confirm adaptations of the middle ear to aquatic lifestyles in multiple independent bird lineages, likely facilitating hearing underwater and baroprotection, while potentially constraining the sensitivity of aerial hearing.

Climate change impacts on seabirds and marine mammals: The importance of study duration, thermal tolerance and generation time.

Understanding climate change impacts on top predators is fundamental to marine biodiversity conservation, due to their increasingly threatened populations and their importance in marine ecosystems. We conducted a systematic review of the effects of climate change (prolonged, directional change) and climate variability on seabirds and marine mammals. We extracted data from 484 studies (4808 published studies were reviewed), comprising 2215 observations on demography, phenology, distribution, diet, behaviour, body condition and physiology. The likelihood of concluding that climate change had an impact increased with study duration. However, the temporal thresholds for the effects of climate change to be discernibly varied from 10 to 29 years depending on the species, the biological response and the oceanic study region. Species with narrow thermal ranges and relatively long generation times were more often reported to be affected by climate change. This provides an important framework for future assessments, with guidance on response- and region-specific temporal dimensions that need to be considered when reporting effects of climate change. Finally, we found that tropical regions and non-breeding life stages were poorly covered in the literature, a concern that should be addressed to enable a better understanding of the vulnerability of marine predators to climate change.

Population structure associated with bioregion and seasonal prey distribution for Indo-Pacific bottlenose dolphins (Tursiops aduncus) in South Africa.

Many marine species exhibit fine-scale population structure despite high mobility and a lack of physical barriers to dispersal, but the evolutionary drivers of differentiation in these systems are generally poorly understood. Here we investigate the potential role of habitat transitions and seasonal prey distributions on the evolution of population structure in the Indo-Pacific bottlenose dolphin, Tursiops aduncus, off South Africa's coast, using double-digest restriction-site associated DNA sequencing. Population structure was identified between the eastern and southern coasts and correlated with the habitat transition between the temperate Agulhas (southern) and subtropical Natal (eastern) Bioregions, suggesting differentiation driven by resource specializations. Differentiation along the Natal coast was comparatively weak, but was evident in some analyses and varied depending on whether the samples were collected during or outside the seasonal sardine (Sardinops sagax) run. This local abundance of prey could influence the ranging patterns and apparent genetic structure of T. aduncus. These findings have significant and transferable management implications, most importantly in terms of differentiating populations inhabiting distinct bioregions and seasonal structural patterns within a region associated with the movement of prey resources.

Spatial segregation in a sexually dimorphic central place forager: Competitive exclusion or niche divergence?

Sexual competition is increasingly recognized as an important selective pressure driving species distributions. However, few studies have investigated the relative importance of interpopulation versus intrapopulation competition in relation to habitat availability and selection. To explain spatial segregation between sexes that often occurs in non-territorial and central place foragers, such as seabirds, two hypotheses are commonly used. The 'competitive exclusion' hypothesis states that dominant individuals should exclude subordinate individuals through direct competition, whereas the 'niche divergence' hypothesis states that segregation occurs due to past competition and habitat specialization. We tested these hypotheses in two populations of an extreme wide-ranging and sexually dimorphic seabird, investigating the relative role of intrapopulation and interpopulation competition in influencing sex-specific distribution and habitat preferences. Using GPS loggers, we tracked 192 wandering albatrosses Diomedea exulans during four consecutive years (2016-2019), from two neighbouring populations in the Southern Ocean (Prince Edward and Crozet archipelagos). We simulated pseudo-tracks to create a null spatial distribution and used Kernel Density Estimates (KDE) and Resource Selection Functions (RSF) to distinguish the relative importance of within- versus between-population competition. Kernel Density Estimates showed that only intrapopulation sexual segregation was significant for each monitoring year, and that tracks between the two colonies resulted in greater overlap than expected from the null distribution, especially for the females. RSF confirmed these results and highlighted key at-sea foraging areas, even if the estimated of at-sea densities were extremely low. These differences in selected areas between sites and sexes were, however, associated with high interannual variability in habitat preferences, with no clear specific preferences per site and sex. Our results suggest that even with low at-sea population densities, historic intrapopulation competition in wide-ranging seabirds may have led to sexual dimorphism and niche specialization, favouring the 'niche divergence' hypothesis. In this study, we provide a protocol to study competition within as well as between populations of central place foragers. This is relevant for understanding their distribution patterns and population regulation, which could potentially improve management of threatened populations.

Foraging in a dynamic environment: Response of four sympatric sub-Antarctic albatross species to interannual environmental variability.

Seasonal and annual climate variations are linked to fluctuations in the abundance and distribution of resources, posing a significant challenge to animals that need to adjust their foraging behavior accordingly. Particularly during adverse conditions, and while energetically constrained when breeding, animals ideally need to be flexible in their foraging behavior. Such behavioral plasticity may separate "winners" from "losers" in light of rapid environmental changes due to climate change. Here, the foraging behavior of four sub-Antarctic albatross species was investigated from 2015/16 to 2017/18, a period characterized by pronounced environmental variability. Over three breeding seasons on Marion Island, Prince Edward Archipelago, incubating wandering (WA, Diomedea exulans; n = 45), grey-headed (GHA, Thalassarche chrysostoma; n = 26), sooty (SA, Phoebetria fusca; n = 23), and light-mantled (LMSA, P. palpebrata; n = 22) albatrosses were tracked with GPS loggers. The response of birds to environmental variability was investigated by quantifying interannual changes in their foraging behavior along two axes: spatial distribution, using kernel density analysis, and foraging habitat preference, using generalized additive mixed models and Bayesian mixed models. All four species were shown to respond behaviorally to environmental variability, but with substantial differences in their foraging strategies. WA was most general in its habitat use defined by sea surface height, eddy kinetic energy, wind speed, ocean floor slope, and sea-level anomaly, with individuals foraging in a range of habitats. In contrast, the three smaller albatrosses exploited two main foraging habitats, with habitat use varying between years. Generalist habitat use by WA and interannually variable use of habitats by GHA, SA, and LMSA would likely offer these species some resilience to predicted changes in climate such as warming seas and strengthening of westerly winds. However, future investigations need to consider other life-history stages coupled with demographic studies, to better understand the link between behavioral plasticity and population responses.

Cross-continental analysis of coastal biodiversity change.

Whereas the anthropogenic impact on marine biodiversity is undebated, the quantification and prediction of this change are not trivial. Simple traditional measures of biodiversity (e.g. richness, diversity indices) do not capture the magnitude and direction of changes in species or functional composition. In this paper, we apply recently developed methods for measuring biodiversity turnover to time-series data of four broad taxonomic groups from two coastal regions: the southern North Sea (Germany) and the South African coast. Both areas share geomorphological features and ecosystem types, allowing for a critical assessment of the most informative metrics of biodiversity change across organism groups. We found little evidence for directional trends in univariate metrics of diversity for either the effective number of taxa or the amount of richness change. However, turnover in composition was high (on average nearly 30% of identities when addressing presence or absence of species) and even higher when taking the relative dominance of species into account. This turnover accumulated over time at similar rates across regions and organism groups. We conclude that biodiversity metrics responsive to turnover provide a more accurate reflection of community change relative to conventional metrics (absolute richness or relative abundance) and are spatially broadly applicable. This article is part of the theme issue 'Integrative research perspectives on marine conservation'.

Foraging behaviour and habitat-use drives niche segregation in sibling seabird species.

To mediate competition, similar sympatric species are assumed to use different resources, or the same but geographically separated resources. The two giant petrels (Macronectes spp.) are intriguing in that they are morphologically similar seabirds with overlapping diets and distributions. To better understand the mechanisms allowing their coexistence, we investigated intra- and interspecific niche segregation at Marion Island (Southern Indian Ocean), one of the few localities where they breed in sympatry. We used GPS tracks from 94 individuals and remote-sensed environmental data to quantify habitat use, combined with blood carbon and nitrogen stable isotope ratios from 90 individuals to characterize their foraging habitat and trophic ecology. Females of both species made distant at-sea foraging trips and fed at a similar trophic level. However, they used distinct pelagic habitats. By contrast, males of both species mainly foraged on or near land, resulting in significant sexual segregation, but high interspecific habitat and diet overlap. However, some males showed flexible behavioural strategies, also making distant, pelagic foraging trips. Using contemporaneous tracking, environmental and stable isotope data we provide a clear example of how sympatric sibling species can be segregated along different foraging behaviour dimensions.

Abundance of Indo-Pacific bottlenose dolphins (Tursiops aduncus) along the south coast of South Africa.

Coastally distributed dolphin species are vulnerable to a variety of anthropogenic pressures, yet a lack of abundance data often prevents data-driven conservation management strategies from being implemented. We investigated the abundance of Indo-Pacific bottlenose dolphins (Tursiops aduncus) along the south coast of South Africa, from the Goukamma Marine Protected Area (MPA) to the Tsitsikamma MPA, between 2014 and 2016. During this period, 662.3h of boat-based photo-identification survey effort was carried out during 189 surveys. The sighting histories of 817 identified individuals were used to estimate abundance using capture-recapture modelling. Using open population (POPAN) models, we estimated that 2,155 individuals (95% CI: 1,873-2,479) occurred in the study area, although many individuals appeared to be transients. We recorded smaller group sizes and an apparent decline in abundance in a subset of the study area (Plettenberg Bay) compared to estimates obtained in 2002-2003 at this location. We recorded declines of more than 70% in both abundance and group size for a subset of the study area (Plettenberg Bay), in relation to estimates obtained in 2002-2003 at this location. We discuss plausible hypotheses for causes of the declines, including anthropogenic pressure, ecosystem change, and methodological inconsistencies. Our study highlights the importance of assessing trends in abundance at other locations to inform data-driven conservation management strategies of T. aduncus in South Africa.

Determinants of external and blood parasite load in African penguins (Spheniscus demersus) admitted for rehabilitation.

We investigate the factors associated with the occurrence and abundance of external and blood parasites in African penguins (Spheniscus demersus), an endangered seabird that breeds exclusively on the coasts of Namibia and South Africa. External parasites were collected using the dust-ruffling method from 171 African Penguins admitted at a rehabilitation facility in the Western Cape, South Africa. Additionally, blood smears were obtained upon admission and weekly during rehabilitation and examined for blood parasites. Fleas Parapsyllus longicornis humboldti, ticks Ornithodoros capensis and lice Austrogoniodes demersus were recovered from 93, 63 and 40%, respectively, of the penguins upon admission to the centre. Rescue location and age group were identified as significant determinants of flea abundance, whereas month of admission was a significant determinant of tick abundance. Blood parasites were also common on admission, with Babesia being the most frequent (46% prevalence) whereas Borrelia was recorded sporadically (1.2%) and Plasmodium was recorded once. The prevalence and abundance of ticks on admission was positively associated with Babesia infection on admission. Our findings demonstrate the variability and contributing factor of parasite infections in an endangered species of penguin, and highlight the need for additional research on the parasite-host dynamics involving these potential disease vectors.

PROGNOSTIC INDICATORS OF IMMATURE REHABILITATED AFRICAN PENGUINS (SPHENISCUS DEMERSUS) IN SOUTH AFRICA.

The Southern African Foundation for the Conservation of Coastal Birds (Cape Town, South Africa) regularly receives African Penguins (Spheniscus demersus) for rehabilitation. The impact of life history and clinical parameters at admission of 3,975 chicks, 1,687 blues (chicks without downy plumage that are about to fledge or have recently fledged), and 850 fledged juveniles admitted over a 12 yr period (2002-13) were assessed in relation to rehabilitation outcomes using multivariate analysis. Younger chicks were more likely to die during rehabilitation compared with older chicks, and chicks admitted because they had been abandoned by their parents were more likely to die than those admitted preemptively (chicks that were removed when rangers considered their nests were in high-risk areas). Blues admitted because of injury and juveniles admitted because of debilitation, injury, and molt were more likely to die or be euthanized relative to the reference category oiling. Plasmodium infection contributed to natural death for all age groups and to the decision to euthanize chicks, whereas Borrelia infection contributed to both natural death and the decision to euthanize blues. Babesia infection was associated with decreased odds of euthanasia among juveniles. Low hematocrit at admission contributed to natural death of chicks and blues and euthanasia of blues. Low total plasma protein, on the other hand, contributed to natural death in chicks, blues, and juveniles, whereas high total plasma protein had a protective effect against natural death in chicks. These results indicate a need to focus on prevention and early intervention among compromised immature penguins in order to increase the rehabilitation success for this endangered species.

Nasal mites (Mesostigmata: Rhinonyssidae) in African penguins (Spheniscus demersus).

Rhinonyssids are obligate haematophagous mites that parasitize the nasal cavity of vertebrates, and occur in a wide range of birds worldwide. Two species of nasal mites are known to occur in penguins: Rhinonyssus sphenisci, which has been recorded from Humboldt and Magellanic penguins (Spheniscus humboldti and S. magellanicus, respectively), and Rhinonyssus schelli, which has been recorded in Adélie and Gentoo penguins (Pygoscelis adeliae and P. papua, respectively). We examined the nasal cavity of African penguins (Spheniscus demersus) that died while under care at a rehabilitation centre (N = 40) or that were found dead at breeding colonies (N = 67). Nasal mites were found in the nasal cavity and/or paranasal of sinuses of 21 penguins, some of which had signs of mild-to-moderate sinusitis. Prevalence was higher in juveniles (29%) and adults (27%) than in chicks (10%). Mean intensity was 5.9 ± 12.9 mites per infected host (range 1-60). The mites presented morphological characteristics that were at times consistent with either R. sphenisci or R. schelli, and therefore we conservatively classified them as 'R. sphenisci sensu lato'. Our morphometric results raise the question of whether the specific status of R. schelli is justified.

First evidence of underwater vocalisations in hunting penguins.

Seabirds are highly vocal on land where acoustic communication plays a crucial role in reproduction. Yet, seabirds spend most of their life at sea. They have developed a number of morphological, physiological and behavioural adaptations to forage in the marine environment. The use of acoustic signals at sea could potentially enhance seabirds' foraging success, but remains largely unexplored. Penguins emit vocalisations from the sea surface when commuting, a behaviour possibly associated with group formation at sea. Still, they are unique in their exceptional diving abilities and feed entirely underwater. Other air-breathing marine predators that feed under water, like cetaceans, pinnipeds and marine turtles, are known to emit sound underwater, but such behaviour has not yet been described in seabirds. We aimed to assess the potential prevalence and diversity of vocalisations emitted underwater by penguins. We chose three study species from three different genera, and equipped foraging adults with video cameras with built-in microphones. We recorded a total of 203 underwater vocalisation from all three species during 4 h 43 min of underwater footage. Vocalisations were very short in duration (0.06 s on average), with a frequency of maximum amplitude averaging 998 Hz, 1097 Hz and 680 Hz for King, Gentoo and Macaroni penguins, respectively. All vocalisations were emitted during feeding dives and more than 50% of them were directly associated with hunting behaviour, preceeded by an acceleration (by 2.2 s on average) and/or followed by a prey capture attempt (after 0.12 s on average). The function of these vocalisations remain speculative. Although it seems to be related to hunting behaviour, these novel observations warrant further investigation.

Empirical Primaquine Treatment of Avian Babesiosis in Seabirds.

Babesia species are tickborne hemoprotozoans. Although experiments have shown that primaquine is highly effective in the treatment of Babesia species infections in mammals, this drug has not been widely used for the treatment of avian babesiosis. Consequently, the treatment of this disease for avian patients has traditionally relied on an empirically established imidocarb treatment. In this study, the authors examined the efficacy of primaquine as a treatment alternative for avian babesiosis (Babesia peircei and Babesia ugwidiensis) in seabirds. Retrospective analysis was performed on the medical records and blood smears of 446 B peircei-positive African penguins (Spheniscus demersus) and 41 B ugwidiensis-positive Cape cormorants (Phalacrocorax capensis) admitted for rehabilitation at the Southern African Foundation for the Conservation of Coastal Birds (SANCCOB, Cape Town, South Africa). Treatment with primaquine (1 mg/kg PO q24h for 10 days) was effective in rapidly and markedly decreasing the proportion of Babesia-positive blood smears in African penguins and Cape cormorants. No regurgitation, loss of appetite, or any other signs after administration of primaquine were observed during the study period. The use of primaquine can be a particularly advantageous treatment alternative for avian babesiosis in circumstances in which it is not possible to determine confidently whether the intraerythrocytic inclusions seen in blood smears correspond to Babesia or Plasmodium or in cases in which there is a coinfection by Babesia and Plasmodium.

A novel candidate species of Anaplasma that infects avian erythrocytes.

BACKGROUND: Anaplasma spp. are Gram-negative obligate intracellular bacteria transmitted by ticks. Even though numerous studies have detected DNA from Anaplasma spp. in the blood of birds, thus far mammals were the only vertebrates demonstrated to serve as competent hosts to these organisms. We report a novel candidate species of Anasplasma that was associated with cytoplasmic inclusions in the erythrocytes of an African penguin (Spheniscus demersus) in South Africa. METHODS: Cytoplasmic inclusions were morphologically characterized from freshly-produced blood smears, and phylogenetic analysis of 16S rRNA and groEL genes were used to evaluate the evolutionary relationships of the organism to other Anaplasmataceae. RESULTS: Dark-purple round or oval inclusions consistent with Anaplasmataceae morulae were observed in the cytoplasm of erythrocytes. Phylogenetic trees produced using different methods agreed that the organism detected in this study belongs to the genus Anaplasma, and suggested that it is most closely related to the cluster comprising A. centrale, A. capra, A. marginale and A. ovis. We propose provisionally naming the strain detected in this study as "Candidatus Anaplasma sphenisci". CONCLUSIONS: This is the first species of Anaplasma shown to produce cytoplasmic inclusions in avian cells, opening the possibility that cytoplasmic inclusions in avian erythrocytes that had previously been attributed to Aegyptianella sp. might in fact correspond to Anaplasma. Further studies on the molecular biology of avian-infecting Anaplasmataceae will be valuable to provide insight into the evolution and epidemiology of these organisms.

Molecular characterization and lesions associated with Diomedenema diomedeae (Aproctoidea: Desmidocercidae) from grey-headed albatrosses (Thalassarche chrysostoma) on Subantarctic Marion Island.

The Prince Edward Islands are Subantarctic islands in the southwest Indian Ocean that are of global importance as seabird nesting sites, and are breeding grounds for five species of albatrosses (Procellariiformes: Diomedeidae). In March-April 2016 numerous chicks of one of these species, the grey-headed albatross (Thalassarche chrysostoma), were found dead at colonies on Marion Island (46°57'S 37°42'E), the larger of the two Prince Edward Islands. Affected chicks were weak, prostrated, apathetic, had drooping wings, and many eventually died while sitting on the nest. Five carcasses were necropsied, and samples were obtained for pathological and parasitological analysis. Four chicks appeared to have died from starvation, and one died due to air-sac helminthiasis, with extensive hemorrhage in the air sacs and multifocal pyogranulomatous air-sacculitis. The air sac parasites were identified as Diomedenema diomedeae (Aproctoidea: Desmidocercidae). Phylogenetic analysis of the nuclear 18S rRNA gene and mitochondrial COI gene confirmed that D. diomedeae belongs to the suborder Spirurina and showed that it is most closely related to the Diplotriaenidae (superfamily Diplotriaenoidea), a family of parasites that infect the air sacs and subcutaneous tissues of a variety of bird species. To our knowledge this is the first record of the occurrence of a nematode in the respiratory tract of an albatross and the first study to provide DNA sequences for a species of the superfamily Aproctoidea.

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.

Reappraisal of the Trophic Ecology of One of the World's Most Threatened Spheniscids, the African Penguin.

Many species of seabirds, including the only penguin species breeding on the African continent, are threatened with extinction. The world population of the endangered African penguin Spheniscus demersus has decreased from more than 1.5 million individuals in the early 1900s to c.a. 23 000 pairs in 2013. Determining the trophic interactions of species, especially those of conservation concern, is important when declining numbers are thought to be driven by food limitation. By and large, African penguin dietary studies have relied on the identification of prey remains from stomach contents. Despite all the advantages of this method, it has well known biases. We therefore assessed the African penguin's diet, using stable isotopes, at two colonies in Algoa Bay (south-east coast of South Africa). These represent over 50% of the world population. Various samples (blood, feathers, egg membranes) were collected for carbon and nitrogen stable isotope analyses. Results indicate that the trophic ecology of African penguins is influenced by colony, season and age class, but not adult sex. Isotopic niches identified by standard Bayesian ellipse areas and convex hulls, highlighted differences among groups and variability among individual penguins. Using Bayesian mixing models it was for the first time shown that adults target chokka squid Loligo reynaudii for self-provisioning during particular stages of their annual cycle, while concurrently feeding their chicks primarily with small pelagic fish. This has important ramifications and means that not only pelagic fish, but also squid stocks, need to be carefully managed in order to allow population recovery of African penguin.