Age‐specific survival and movement among major African Penguin Spheniscus demersus colonies

Reliable estimates of survival and dispersal are crucial to understanding population dynamics, but for seabirds, in which some individuals spend years away from land, mortality and emigration are often confounded. Multistate mark–recapture methods reduce bias by incorporating movement into the process of estimating survival. We used a multistate model to provide unbiased age‐specific survival and movement probabilities for the Endangered African Penguin Spheniscus demersus based on 5281 nestlings and 31 049 adults flipper‐banded and resighted in the Western Cape, South Africa, between 1994 and 2012. Adult survival was initially high (≥ 0.74) but declined after 2003–2004 coincident with a reduction in the availability of Sardine Sardinops sagax and Anchovy Engraulis encrasicolus on the west coast of South Africa. Juvenile survival was poorly estimated, but was lower and more variable than adult survival. Fidelity to the locality of origin varied over time, but was high in adults at Robben and Dassen islands (≥ 0.88) and above 0.55 for juvenile and immature Penguins at all localities. Movement occurred predominantly during 1994–2003 and was indicative of immigration to Robben and Dassen islands. Our results confirm that a prolonged period of adult mortality contributed to the observed decline in the African Penguin population and suggest a need for approaches operating over large spatial scales to ensure food security for marine top predators.     

Group Size in Foraging African Penguins (Spheniscus demersus)

Diving synchrony was examined for varying group sizes of African penguins (Spheniscus demersus) travelling to their foraging grounds from their breeding islands. Groups of fewer than 12 birds always dived synchronously, whereas groups of more than 17 birds always dived asynchronously. Since travelling penguins do not dive deeply, large groups of birds can remain together irrespective of diving synchronization. Observations from boats showed that foraging penguins rarely occurred in groups of more than 17 birds. We suggest that groups of penguins that do not have synchronized dives cannot forage effectively, because foraging penguins dive deeply.     

Fibre types in primary ‘flight’ muscles of the African Penguin (Spheniscus demersus)

The African penguin (Spheniscus demersus) is an endangered seabird that resides on the temperate southern coast of Africa. Like all penguins it is flightless, instead using its specialized wings for underwater locomotion termed ‘aquatic flight’. While musculature and locomotion of the large Antarctic penguins have been well studied, smaller penguins show different biochemical and behavioural adaptations to their habitats. We used histochemical and immunohistochemical methods to characterize fibre type composition of the African penguin primary flight muscles, the pectoralis and supracoracoideus. We hypothesized the pectoralis would contain predominantly fast oxidative–glycolytic (FOG) fibres, with mainly aerobic subtypes. As the supracoracoideus and pectoralis both power thrust, we further hypothesized these muscles would have a similar fibre type complement. Our results supported these hypotheses, also showing an unexpected slow fibre population in the deep parts of pectoralis and supracoracoideus. The latissimus dorsi was also examined as it may contribute to thrust generation during aquatic flight, and in other avian species typically contains definitive fibre types. Unique among birds studied to date, the African penguin anterior latissimus dorsi was found to consist mainly of fast fibres. This study shows the African penguin has specialized flight musculature distinct from other birds, including large Antarctic penguins.     

Assessment of microsatellite and SNP markers for parentage assignment in ex situ African Penguin (Spheniscus demersus) populations

Captive management of ex situ populations of endangered species is traditionally based on pedigree information derived from studbook data. However, molecular methods could provide a powerful set of complementary tools to verify studbook records and also contribute to improving the understanding of the genetic status of captive populations. Here, we compare the utility of single nucleotide polymorphisms (SNPs) and microsatellites (MS) and two analytical methods for assigning parentage in ten families of captive African penguins held in South African facilities. We found that SNPs performed better than microsatellites under both analytical frameworks, but a combination of all markers was most informative. A subset of combined SNP (n = 14) and MS loci (n = 10) provided robust assessments of parentage. Captive or supportive breeding programs will play an important role in future African penguin conservation efforts as a source of individuals for reintroduction. Cooperation among these captive facilities is essential to facilitate this process and improve management. This study provided us with a useful set of SNP and MS markers for parentage and relatedness testing among these captive populations. Further assessment of the utility of these markers over multiple (>3) generations and the incorporation of a larger variety of relationships among individuals (e.g., half‐siblings or cousins) is strongly suggested.     

Vocal Repertoire of Captive Capybara (Hydrochoerus hydrochaeris): Structure,Context and Function

In this work we describe the structure, behavioral context and functionality of the vocal repertoire of capybaras, a social Caviomorph rodent. Additionally, since territoriality is present in this species, we hypothesize the occurrence of vocal differences among social groups. We analyzed a total of 2069 calls emitted by 28 animals from three different social groups. The capybara’s repertoire is comprised by seven call types (whistle, cry, whine, squeal, bark, click and tooth‐chattering). The vocalizations were functionally categorized as contact, alarm, distress and agonistic calls considering their behavioral contexts. The click calls emitted by the adults of the three captive capybara groups were significantly different, confirming our hypothesis of social groups’ vocal differences. The richness of interactions mediated by vocalization in capybaras suggests that the species’ communication is an important mechanism to regulate social encounters and to alert members of the group about environmental cues.     

Herpesvirus-like infection in black-footed penguins (Spheniscus demersus)

A herpesvirus-like infection is described in the black-footed penguin (Spheniscus demersus). Clinically, the infection was characterized by debilitation and respiratory distress. Histopathological lesions were confined to the respiratory tract and consisted of inflammation and syncytial cell formation with Type A intranuclear inclusions in sinuses, trachea, and mainstem bronchi. Electron microscopy demonstrated polyhedral viral particles 80-140 nm in size consistent with Herpetoviridae. The lesions resembled those seen in infectious laryngotracheitis.     

Identification of land predators of African Penguins Spheniscus demersus through post-mortem examination

The African Penguin Spheniscus demersus is an endangered seabird endemic to southern Africa, and killing sprees by terrestrial predators have been one of the main threats for its mainland colonies. The methods employed to manage predators may differ depending on the species involved, therefore the implementation of strategies to limit the impacts of predation relies on the correct identification of the culprit predator. We report and quantify the lesions seen in African Penguins killed by four species of terrestrial predators: Caracal Caracal caracal (52 kills), Leopard Panthera pardus (27 kills), Domestic Dog Canis lupus familiaris (10 kills), and Cape Grey Mongoose Galerella pulverulenta (4 kills). We discuss patterns of necropsy findings that can be used to identify the predator species involved. Traditional forensic methods are useful tools to direct species-specific management actions for the conservation of the African Penguin and other seabirds so that effective mitigating measures can be deployed quickly to prevent further losses. It should be borne in mind, however, that the age, size and previous hunting experience of the predator are likely to influence the pattern of lesions that will be observed, and not all carcasses will have hallmark lesions or recognisable bite marks.     

Injuries to jackass penguins (Spheniscus demersus): evidence for shark involvement

An analysis is presented of records of injuries and the types of injuries to jackass penguins in Algoa Bay, South Africa, over a 10-year-period. Sharks are implicated in causing most of the injuries, but other possible agents are discussed. The types of injuries were categorized and compared to the dentition of sharks known to occur in the area. Using a life-size model of a jackass penguin and preserved jaws of sharks, bites were simulated to establish the types of injury inflicted by different species. Certain injuries indicate attack by great white sharks, while others cannot be ascribed to any particular species. Most shark injuries show single jaw involvement, and range from small cuts across the feathers to the bird being cut in two. Sharks accounted for the highest number of identifiable natural causes of death, but the significance of shark attack as a mortality factor in jackass penguins is difficult to assess.     

Pond Use by Captive African Penguins (Spheniscus demersus) in an Immersive Exhibit Adjacent to Human Bathers

Nonhuman animals in zoos are exposed to a continuous human presence, which affects their behaviors and welfare. However, little is known about what role the “visitor effect” has on penguins in captivity. The African penguin (Spheniscus demersus) is an endangered species commonly housed in zoos worldwide. The aim of this study was to evaluate whether the abundance of human bathers could reduce the average time spent in the water of a colony of African penguins housed in an exhibit where their pond habitat was adjacent to a swimming pool. Observations were carried out on 7 penguins in the summer of 2009. Data were collected during 3 time periods (Time 1 [T1] = opening of the swimming season, Time 2 [T2] = core of the season, Time 3 [T3] = late season) of 14 days each. The human disturbance caused by bathers strongly reduced the pond use by penguins at T1 and T2, especially when there were large numbers of visitors. However, at T3, we observed that the overall use of the pond by penguins increased, and the average duration of their diving was no longer dependent on the number of visitors.     

Sex determination of African Penguins Spheniscus demersus using bill measurements: method comparisons and implications for use

African Penguins Spheniscus demersus are sexually dimorphic; on average, males are larger than females but measurements overlap making sex determination difficult through observations alone. We developed a discriminant function, using bill length and depth from a sample of birds sexed from gonad visualisation during post-mortem, which correctly classified 93% of the individuals. Cross-validation correctly assigned 90% of DNA-sexed birds and 91% of birds sexed by partner measurement comparisons. The use of discriminant function score cutpoints, while leaving 16% and 29% of birds unclassified, improved accuracy of birds sexed by DNA to 97% and of those sexed by partner comparison to 99%. Bill depth was found to be a discriminating variable. However, two techniques for measuring bill depth are currently in use for African Penguins. While these measurements are correlated (r = 0.85), they differ on average by 1.4?mm hindering accuracy of sex determination when using a discriminant function developed from the other bill depth measurement. Exploration of adult bill morphology of birds sexed from DNA at different colonies suggests the discriminant functions can be applied throughout the African Penguins’ South African range.     

Of squeezers and skippers: factors determining the age at moult of immature African Penguins Spheniscus demersus in Namibia

We used banding and resighting records of 391 African Penguins Spheniscus demersus banded as chicks and later resighted during immature moult to explain the roles of date of fledging and age at moult in determining the season of moult and its timing within the season. Breeding was continuous, but immature moult occurred mainly during spring and summer. Age at immature moult extended over 11 months, from 12 to 23 months after hatching. Birds that fledged during summer and early autumn generally moulted during the next moult season (squeezers), whereas birds that fledged in late autumn, winter and spring skipped the next moult season to moult only the following season (skippers). There was a significant relationship between age at moult and moult date, with young birds moulting later in the season than older birds. The age at moult of immature birds appears to be constrained by minimum age, moult seasonality and plumage wear. Birds that fledged over nearly 2 years moult during one season. Counts of moulting immature African Penguins have not been used to estimate year-class strength and post-fledging survival owing to the wide range of ages at immature moult. Our results provide the means of assigning recruits to specific age groups.     

Evaluation of potential variables contributing to the development and duration of plantar lesions in a population of aquarium-maintained African penguins (Spheniscus demersus)

Bumblefoot (pododermatitis), often described as the most significant environmental disease of captive penguins, is commonly due to excessive pressure or trauma on the plantar surface of the avian foot, resulting in inflammation or necrosis and causing severe swelling, abrasions, or cracks in the skin. Although not formally evaluated in penguins, contributing factors for bumblefoot are thought to be similar to those initiating the condition in raptors and poultry. These factors include substrate, body weight, and lack of exercise. The primary purpose of this retrospective study was to evaluate variables potentially contributing to the development and duration of plantar lesions in aquarium-maintained African penguins (Spheniscus demersus), including sex, weight, age, season, exhibit activity, and territory substrate. Results indicate that males develop significantly more plantar lesions than females. Penguins weighing between 3.51 and 4.0 kg develop plantar lesions significantly more often than penguins weighing between 2.5 and 3.5 kg, and because male African penguins ordinarily weigh significantly more than females, weight is likely a contributing factor in the development of lesions in males compared with females. Significantly more plantar lesions were observed in penguins standing for greater than 50% of their time on exhibit than swimming. Penguins occupying smooth concrete territories developed more plantar lesions compared with penguins occupying grate territories. Recommendations for minimizing bumblefoot in African penguins include training penguins for monthly foot examinations for early detection of plantar lesions predisposing for the disease, encouraging swimming activity, and replacing smooth surfaces on exhibit with surfaces providing variable degrees of pressure and texture on the feet.     

Growth of Jackass penguin chicks (Spheniscus demersus) hand reared on different diets

Jackass penguin chicks from the age of about 10 days were hand reared on different diets: mullet Liza richardsoni , anchovy Engraulis capensis and squid Loligo reynaudi , until they fledged. The weight of food fed and the daily weight increment of the chicks was measured. Excreta were collected every 5–6 days and analysed to establish metabolized energy and metabolic efficiency. Culmen measurements were taken regularly and plotted against age. The age, weight and sum of food fed up to fledging was compared between diets as well as with field data. Compensatory growth was exhibited when stunted chicks on a relatively poor diet had their diet changed to a relatively better diet.     

Foraging ranges and movements of jackass penguins (Spheniscus demersus) established through radio telemetry

In order to study the movements and activities at sea of jackass penguins rearing chicks, a radio telemetry study was undertaken. Twenty five transmitters were attached to breeding adult penguins at their nest sites. These penguins were tracked for a total of 414 hours using a system of antennae on a motor–boat, the island and land–based stations. The foraging paths, foraging duration, foraging distances and swimming speeds were studied. The penguins' foraging patterns were similar; they generally left the island at night and travelled towards their foraging area, actively foraging from first light until they swam back to the island. The foraging durations and distances varied, as did the swimming speeds, according to availability of food.     

Plasma electrolyte concentrations in captive and free‐ranging African penguins (Spheniscus demersus) maintained with and without dietary salt supplements

African penguins (Spheniscus demersus) are common display animals in North American zoos and aquariums. At present, 43 American Zoo and Aquarium Association (AZA)‐accredited institutions maintain over 700 of these birds. A survey of facilities found that only seven institutions maintain their birds in “saltwater habitats” (natural, brine, or artificial seawater systems), while the remaining institutions provide freshwater habitats. Penguins maintained in freshwater habitats commonly receive dietary salt supplements, though evidence is lacking as to the value of this practice. This study was designed to evaluate the necessity of salt supplementation in African penguins. We report the results of a randomized, 10‐month comparison of plasma electrolytes (Na, Cl, and K) between groups of 10 salt‐supplemented and nonsupplemented birds living in a freshwater environment. The results showed no significant differences between the two groups. An interfacility comparison at the same time points revealed temporal pattern differences in analyte values between the facilities; however, the absolute concentrations did not exceed clinical ranges found in healthy birds. Furthermore, single time‐point comparisons between wild and captive African penguins indicated similar sodium concentrations, while potassium and chloride concentrations varied between groups. Finally, the plasma electrolyte levels in 20 birds remained statistically unchanged after a 60‐day withdrawal from salt supplementation. This study provides experimental evidence that African penguins maintained in freshwater exhibits on a herring‐, capelin‐, and squid‐based diet do not require salt supplementation. In addition to the practical implications regarding the need for salt supplementation for captive birds, the results also have theoretical significance. They provide evidence that penguins do not need to ingest high salt concentrations to remain in electrolyte balance. In this regard, the ion regulatory mechanisms in penguins are similar to those in terrestrial mammals and birds. Zoo Biol 23:397–408, 2004. © 2004 Wiley‐Liss, Inc.     

Vocal Repertoire of Cebus capucinus: Acoustic Structure, Context, and Usage

Researchers studying nonhuman primate vocal repertoires suggest that convergent environmental, social, and motivational factors account for intra- and interspecific vocal variation. We provide a detailed overview of the vocal repertoire of white-faced capuchins, including acoustic analyses and contextual information of vocal production and vocal usage by different age-sex classes in social interactions. The repertoire is a mixture of graded and discrete vocalizations. In addition, there is general support for structural variation in vocalizations with changes in arousal level. We also identified several combined vocalizations, which might represent variable underlying motivations. Lastly, by including data on the social contexts and production of vocalizations by different age-sex classes, we provide preliminary information about the function of vocalizations in social interactions for individuals of different rank, age, and sex. Future studies are necessary to explore the function of combined vocalizations and how the social function of vocalizations relate to their acoustic structure, because social use of vocalizations may play an important role in shaping vocal evolution.     

Vocalisation Repertoire of Female Bluefin Gurnard (Chelidonichthys kumu) in Captivity: Sound Structure,Context and Vocal Activity

Fish vocalisation is often a major component of underwater soundscapes. Therefore, interpretation of these soundscapes requires an understanding of the vocalisation characteristics of common soniferous fish species. This study of captive female bluefin gurnard, Chelidonichthys kumu, aims to formally characterise their vocalisation sounds and daily pattern of sound production. Four types of sound were produced and characterised, twice as many as previously reported in this species. These sounds fit two aural categories; grunt and growl, the mean peak frequencies for which ranged between 129 to 215 Hz. This species vocalized throughout the 24 hour period at an average rate of (18.5 ± 2.0 sounds fish^-1 h^-1) with an increase in vocalization rate at dawn and dusk. Competitive feeding did not elevate vocalisation as has been found in other gurnard species. Bluefin gurnard are common in coastal waters of New Zealand, Australia and Japan and, given their vocalization rate, are likely to be significant contributors to ambient underwater soundscape in these areas.