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.     

Genetic diversity in populations of Gentoo penguins (Pygoscelis papua)

RAPD analysis was used to examine the extent of genetic polymorphism in two populations of Gentoo penguin (Pygoscelis papua) from Antarctic Islands (Petermann and Livingston). The chosen two of three 10 mer oligonucleotide primers accordingly to preliminary results showed different levels of polymorphism in Gentoo penguins at Petermann Island (from 23.53 to 42.86%) and Livingston Island (from 52.94 to 57.14%). Nei's similarity coefficients were in range from 0.5606 (when Gentoo genome profiles were compared with RAPD profiles of two related penguin species: Pygoscelis adeliae (Adelie) and Pygoscelis antarctica (Chinstrep)) to 0.9281 among observed Gentoo penguin populations. Nei's distances values ranged from 0.0746 to 0.5787 among the populations and species. The obtained results will be used for further estimation of genetic diversity of Gentoo penguins and determination of their taxonomic status.     

The phylogeny of the living and fossil Sphenisciformes (penguins)

We present the first phylogenetic analysis of the Sphenisciformes that extensively samples fossil taxa. Combined analysis of 181 morphological characters and sequence fragments from mitochondrial and nuclear genes (12S, 16S, COI, cytochrome b, RAG‐1) yields a largely resolved tree. Two species of the New Zealand Waimanu form a trichotomy with all other penguins in our result. The much discussed giant penguins Anthropornis and Pachydyptes are placed in two clades near the base of the tree. Stratigraphic and phylogenetic evidence suggest that some lineages of penguins attained very large body size rapidly and early in the clade's evolutionary history. The only fossil taxa that fall inside the crown clade Spheniscidae are fossil species assigned to the genus Spheniscus. Thus, extant penguin diversity is more accurately viewed as the product of a successful radiation of derived taxa than as an assemblage of survivors belonging to numerous lineages. The success of the Spheniscidae may be due to novel feeding adaptations and a more derived flipper apparatus. We offer a biogeographical scenario for penguins that incorporates fossil distributions and paleogeographic reconstructions of the Southern continent's positions. Our results do not support an expansion of the Spheniscidae from a cooling Continental Antarctica, but instead suggest those species that currently breed in that area are the descendants of colonizers from the Subantarctic. Many important divergence events in the clade Spheniscidae can instead be explained by dispersal along the paths of major ocean currents and the emergence of new islands due to tectonic events. © The Willi Hennig Society 2006.     

Behavioural time budget of breeding king penguins (Aptenodytes patagonica)

As do so many other seabirds, penguins fast when ashore for breeding. For penguins in dense colonies, territory defence seems to imply conflicting energetic requirements because of its assumed high energy cost, when the birds need to limit energy expenditure to cope with their fast. In this context, behavioural time budget over 24 h was investigated during breeding in the king penguin, Aptenodytes putugonicu , by using a remote-controlled videocamera. The comparison of day-night activity was performed in relation to breeding status (incubation vs. brooding) and duration of fasting (beginning vs. end of incubation shift). Five categories of behaviours were quantified: territoly defence, comfort, resting, sleeping and chick-feeding. Breeding king penguins remain active by day as well as by night. Between incubation and brooding we found a three-fold increase in the energy consuming temtory defence, together with a drastic decrease in that posture which corresponds to deep sleep, is. when most energy is saved. These increases in aggressiveness and vigilance may be related to protection of the newly hatched chick. Between the onset and the end of an incubation shift, the time spent in sleep increases three-fold, whereas territory defence remains unchanged. These data for penguins under natural conditions accord with previous studies on captive birds which have shown that an increasing proportion of sleep during the course of fasting may contribute to energy saving. On the other hand, both resting (which is the main component of penguins'time budget; about 65%) and comfort (about 16% of time) show no change either between incubation and brooding or during the course of fasting.     

Health evaluation of penguins (Sphenisciformes) following mortality in the Falklands (South Atlantic)

In the Falklands, heavy mortality of rock-hopper penguins Eudyptes chrysocome occurred during the 1985-86 breeding season. Starvation was diagnosed as the primary cause of death, possibly caused by a shortage of euphausiid crustaceans (krill) due to unusual meterological conditions. 'Puffinosis' may possibly have been a contributory factor; otherwise no conclusive evidence of infectious disease or toxicosis was found and also no evidence of radioactive contamination. In the 1986-87 breeding season no unusual mortality occurred, but 99 apparently healthy penguins were examined, i.e., rockhoppers Eudyptes chrysocome syn E. crestatus, gentoos Pygoscelis papua and Magellanics Spheniscus magellanicus. Full necropsies were carried out on 54. Tissue examinations were made for cadmium, copper, iron, manganese, mercury, lead and zinc. High tissue cadmium concentrations found in healthy birds in 1987 were similar to those in penguins which died in 1986, and therefore not considered to be of pathological significance. Although there has been no repetition of the unusually hot 1985-86 breeding season in the Falklands, penguins and other seabirds have had fluctuating breeding successes since then. The precise cause, including the roles of meteorological conditions and overexploitation of some forms of prey species, is unclear.     

Gastrointestinal helminths of King penguins (Aptenodytes patagonicus) at Crozet Archipelago

Communities of helminths are known to be related to feeding behaviors of hosts. While climate change and overfishing can impact food availability for Antarctic piscivorous predators, knowledge about infectious and parasitic diseases among Antarctic species is scarce or fragmentary. We studied the helminth community of King penguins (Aptenodytes patagonicus) from the Crozet Archipelago, the main breeding area of the species. Based on a sample of 41 individuals found freshly dead from predation or starvation, the gastrointestinal helminth community in King penguins was composed of 1 species of cestode (Tetrabothrius wrighti) and 2 species of nematodes (Tetrameres wetzeli and Contracaecum heardi). Cestodes formed the core of the helminth community (97.5% of worms collected) with a prevalence of infestation of 100% and a mean intensity of 178.6 worms per host. Sources of infestation and pathologies caused by these worms are also discussed.     

Diving behaviour and energetics in breeding little penguins (Eudyptula minor)

We present data on the diving behaviour and the energetics of breeding little penguins in Tasmania, Australia. Using an 18 m long still water canal in conjunction with respirometry, we determined the energy requirements while diving. Using electronic devices measuring dive depth or swimming speed, we investigated the foraging behaviour at sea. Cost of Transport was calculated to be minimal at the speed the birds prefer at sea (1.8 m/s) and averaged 11.1 J/kg/m (power requirements at that speed: 20.0 W/kg). Metabolic rate of little penguins resting in water was found to be 8.5 W/kg. The externally-attached devices had no significant influence on the energy expenditure.
Foraging trips can be divided into four distinct phases with different diving behaviours. A mean of 500 dives was executed per foraging trip lasting about 18 hours with 60% of this time being spent swimming. The total distance travelled averaged 73 km per day, although foraging range was about 12km. Mean swimming speed of little penguins at sea was 1.8 m/s, maximum swimming speed was 3.3 m/s. More than 50% of all dives had maxima not exceeding 2 m. Maximum depth reached was 27 m. Mean dive duration was 21 s. There were inter-sex differences in diving behaviour as well as changes in foraging behaviour over the breeding period. Aerobic dive limits (ADL) in the wild were estimated between 42 and 50 s. From the swim canal experiments we derived an ADL of 44 s. Total oxygen stores were calculated to be 45 ml O₂/kg. Only 2% of all dives exceeded the ADL. FMRs at sea were calculated to be between 1280 and 1500 kJ/kg/d according to chick size. The yearly food requirements of a breeding little penguin amount to 114 kg.     

Hormonal correlates of parental behavior in yellow-eyed penguins (Megadyptes antipodes)

Penguins show varying degrees of brood reduction behavior, from obligate brood reducers to brood maximizers, and we hypothesize that this is associated with differences in prolactin secretion. To address this hypothesis, we determined the breeding season prolactin profile of the yellow-eyed penguin (Megadyptes antipodes) for comparison with those of other penguin species found in the literature. We also measured sex steroid plasma concentrations to better characterize the reproductive cycle of the species. Plasma concentrations of prolactin increased from early in the season, reaching a peak during late incubation, and remained elevated up to the guard period. This general pattern was similar to that of other penguins for which we have corresponding data. However, we found that throughout the laying period, prolactin titers in yellow-eyed penguins remained elevated while they fell to basal levels after the laying of the first egg in macaroni penguins, which corresponds to differences in incubation behavior during this time. We conclude, therefore, that differences in the brood reduction behavior in penguins, may be reflected in the pattern of PRL concentrations around the time of egg laying.     

Heat loss and site-dependent fecundity in chinstrap penguins (Pygoscelis antarctica)

We study the effects of heat loss and nest site quality on fecundity in a chinstrap penguin (Pygoscelis antarctica) colony on Deception Island, Antarctica. During the austral summers of 1990–1991, 1993–1994, and 1995–1996, 441 randomly selected nests were analyzed. Penguins breeding in the center of subcolonies hatched significantly earlier and had larger broods and than those nesting near or at the edge of the subcolonies. These differences, however, were significantly affected by interaction between year and nest location, being highly significant in colder years, when peripheral nests produce fewer young. Analysis of the rate of heat loss showed that penguins breeding at the edge of subcolonies lost heat twice as rapidly as those breeding in the interior of the subcolony. In a re-occupation experiment, evacuated center nests were re-occupied almost ten times as rapidly as edge nests. An analysis of mean fecundity in the period 1991–1996 and mean wind chill suggested that most of the variability in fecundity among years was related to differences in the rate of heat loss. Subcolonies tend to be as circular as possible, thereby decreasing the proportion of edge nests as the size of subcolony increases. Our results support the site-dependent fecundity hypothesis.     

Climate fluctuation effects on breeding of blue penguins (Eudyptula minor)

Abstract

El Niño and La Niña climate perturbations alter sea currents and food availability for seabirds in many areas of the world. This changes their breeding success and mortality. Blue penguin (Eudyptula minor) breeding success is dependent upon whether one or two clutches per season are laid, and the hatching and fledging success of these clutches. This study uses six years of data from five blue penguin breeding colonies, three from Taiaroa Head, Otago Peninsula and two from Oamaru, to examine whether annual variation in breeding success correlates with El Niño/La Niña perturbations. When La Niña conditions prevailed, penguins started breeding later, and there was a lower proportion of double breeders than in El Niño and normal years. The probability of a newly hatched chick surviving to fledging was also dependent on whether large‐scale climatic conditions prevailed, whereas hatching success and overall breeding success (number of fledged chicks per breeding pair) showed no correlation with climate perturbations.     

Multiple cenozoic invasions of Africa by penguins (Aves, Sphenisciformes)

Africa hosts a single breeding species of penguin today, yet the fossil record indicates that a diverse array of now-extinct taxa once inhabited southern African coastlines. Here, we show that the African penguin fauna had a complex history involving multiple dispersals and extinctions. Phylogenetic analyses and biogeographic reconstructions incorporating new fossil material indicate that, contrary to previous hypotheses, the four Early Pliocene African penguin species do not represent an endemic radiation or direct ancestors of the living Spheniscus demersus (blackfooted penguin). A minimum of three dispersals to Africa, probably assisted by the eastward-flowing Antarctic Circumpolar and South Atlantic currents, occurred during the Late Cenozoic. As regional sea-level fall eliminated islands and reduced offshore breeding areas during the Pliocene, all but one penguin lineage ended in extinction, resulting in today's depleted fauna.     

The food and feeding ecology of Adélie penguins (Pygoscelis adeliae) and Chinstrap penguins (P. antarctica) at Signy Island, South Orkney Islands

Adélie Pygoscelis adeliae and Chinstrap P. antarctica penguins are important consumers of Southern Ocean marine resources. The stomach contents of adult penguins at Signy Island, South Orkney Islands, were analysed quantitatively throughout the chick-rearing period. They consisted almost exclusively of Antarctic krill Euphausia superba , Adélies eating 35–63% by number and 23–28% by weight of juvenile krill and Chinstraps 72–87% by number and 90–95% by weight of mature krill, as well as small amounts offish and amphipods. Interspecific dietary differences may partly be attributable to Adélies starting breeding one month before Chinstraps but, as they persist when both are simultaneously rearing chicks, the two species may also forage in somewhat different areas.
Krill data from net hauls indicate a substantial overlap in the size of krill taken by scientific, and probably also commercial, operations and by Adélie and Chinstrap penguins.
Chicks were fed c. 300 g of food 0–5-0-8 times per day, Chinstrap chicks without a sibling being fed most frequently. Chicks of both species were most often fed in the late afternoon, and from estimates of swimming speed and feeding frequency adults may feed quite extensively at night.     

Unexpected microanatomical variation among Eocene Antarctic stem penguins (Aves: Sphenisciformes)

The microanatomical and histological structure of Eocene Antarctic stem penguin tarsometatarsi is examined in order to characterise the bone microstructure. Eight adult tarsometatarsi belonging to eight fossil species (Palaeeudyptes gunnari, Palaeeudyptes klekowskii, Anthropornis grandis, Anthropornis nordenskjoeldi, Archaeospheniscus wimani, Marambiornis exilis, Delphinornis arctowskii and Delphinornis larseni) collected from the Antarctic A. nordenskjoeldi Biozone (La Meseta Formation, ～34.2 Ma) were examined. The thin sections revealed a distinctive microanatomical variation among taxa. Whereas Anthropornis spp., A. wimani and P. gunnari possess massive, clearly osteosclerotic bones (medullary cavities absent or strongly reduced), the bones of Delphinornis spp., P. klekowski and M. exilis exhibit well-developed medullary cavities. The cortical bone in all the specimens consists of primary, well-vascularised fibro-lamellar bone and variable amounts of secondary bone. Medullary cavities are coated by a thick layer of lamellar bone tissue and coarse compacted cancellous bone. Although several causes can explain the striking microanatomical variation (e.g. ontogeny), we interpret that such variation is related to differential adaptations to the aquatic life, for which taxa with more massive bones were possibly adapted to deeper and more prolonged diving excursions.     

Energetics of under-water swimming in Adélie penguins (Pygoscelis adeliae)

Summary The energy consumption of Adélie penguins while at rest in water (8.4 W·kg^-1 at 4°C) or swimming below the surface was determined using a 21 m long canal fitted with respiration chambers at each end. Penguins chose to swim 86% of the time at speeds recorded in nature. Cost of transport was lowest (7.9 J·kg^-1·m^-1) at 1.7–2.3 m·s^-1, corresponding to a power input of 15.8 W·kg^-1, and only 50% as high as previously reported. Assuming a muscle efficiency of 0.25, propulsion efficiency is 0.4 and overall efficiency is 0.1. Calculated food requirements vary between 1060 g krill per adult and foraging trip at the beginning of the breeding season and 2500 g at the period of highest demand, prior to crèching of the chicks.Abbreviations BMR basal metabolic rate - COT cost of transport - DEE daily energy expenditure - DF daily food - M mass - P _i power input - P _o power output - PVC polyvinyl chloride - RMR resting metabolic rate - SE standard error - STPD Standard temperature, pressure and density - VO₂ oxygen consumption - t time     

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.     

Notes on parasites in penguins (Spheniscidae) and petrels (Procellariidae) in the Antarctic and Sub-antarctic

Blood smears were examined from 143 penguins of four species (Aptenodytes patagonicus, Eudyptes chrysolophus, E. schlegeli, and Pygoscelis gentoo) from Sub-antarctic Macquarie Island and Heard Island. No blood parasites were reported. The vectors of Hepatozoon albatrossi (reported from three species of albatross) are probably shared by penguins, and it is suggested that the latter are not susceptible to infection with this protozoan. Cestodes of the genus Tetrabothrius were present in large numbers in the intestines of 17 Antarctic petrels (Thalassoica antarctica), and evidence is presented indicating that euphausiid crustaceans may be intermediate hosts.     

Semen characteristics and artificial insemination in rockhopper penguins (Eudyptes chrysocome chrysocome)

This work was performed as part of a multi-year study to determine the cause of the low fertility in captive rockhopper penguins (Eudyptes chrysocome chrysocome) and attempt to increase the fertility through artificial insemination (AI). Semen collection and characterization was performed on 14 male rockhopper penguins. The samples were evaluated for volume, sperm concentration, and sperm quality (motility, forward motility, viability, and morphology). There was a large variation between individuals and between collections for each individual. Mean volume of ejaculate was 0.24 ml. Mean concentration was 47.09 × 10(6) sperm/ml. Mean number of sperm per collection was 6.57 × 10(6). The mean motility was 49.4%. Mean forward motility was 1.7. Mean percentage of living sperm was 88.9%. Mean percentage of sperm with normal morphology was 69.4%. AI was performed on a total of 10 females using pooled semen samples. The birds were also allowed to naturally mate. Ten eggs were laid and three fertile eggs were produced, one of them hatched but died within 24 hr. Paternity testing was performed using 12 microsatellite loci, but unfortunately due to insufficient variability, the paternity of the chick and two embryos could not be determined.