Responses of <Emphasis Type="Italic">Pygoscelis adeliae</Emphasis> and <Emphasis Type="Italic">P. papua</Emphasis> populations to environmental changes at Isla 25 de Mayo (King George Island)

As part of a monitoring study of Adélie and Gentoo penguin colonies, birds occupying nests with eggs and chicks in crèches were counted annually from the 1995/1996 to the 2006/2007 seasons at Stranger Point, Isla 25 de Mayo (King George Island), Antarctica. During the study period the Adélie penguin population showed a decrease of 62%. The number of chicks in crèches followed a similar trend, the smallest number occurring in 2002, when it was 63% lower than in 1995/1996. In contrast, the Gentoo breeding population size increased by 68%, while chicks produced increased by 63%. Despite the opposing trends in population size between species, there was a positive relation in their interannual variation, although the extent, and for some years the direction, of the change observed always favoured Gentoo penguins. Breeding success (chicks in crèches/nests with eggs) fluctuated between 0.65 and 1.26 for Adélies and between 0.76 and 1.27 for Gentoo penguins, and did not differ significantly between species. The similar breeding success of these species suggests that the contrasting population trends observed were driven by factors operating over winter. We suggest that current changes in environmental conditions may affect adult birds of both species during the previous winter with different intensity but in a roughly similar way, but that juvenile survival of both species and thus the recruitment of new breeders might be affected differentially, with a much lower survival rate of juvenile Adélie penguins.     

Leucocyte profiles and corticosterone in chicks of southern rockhopper penguins

The immune system is essential for health and survival of vertebrates, yet still little is known about the ontogeny of the immune system in wild birds. The southern rockhopper penguin (Eudyptes chrysocome chrysocome) is a semi-altricial seabird with a long developmental period and reversed hatching asynchrony, favouring the survival of B-chicks. We compared leucocyte counts and baseline corticosterone levels of southern rockhopper penguin chicks under different preconditions such as sex and origin from an A- or B-egg from 4 to 51 days of age. We conducted an experiment to compare leucocyte profiles and baseline corticosterone levels in A- and B-chicks in single-egg clutches as well as in B-chicks from normal two egg-clutches (one A- and one B-egg). None of these treatments influenced leucocyte counts or corticosterone levels, indicating a similar investment in the immune system. Our main finding was an increase of leucocytes/10,000 erythrocytes with age, which was mainly caused by an increase in lymphocyte numbers. This suggests differential investment into acquired immunity at this stage of development. As the granulocyte/lymphocyte (G/L) ratio did not change with age or body condition, G/L ratios seem not to reflect stress caused by poor provisioning of penguin chicks. This was also reinforced by the decrease of plasma corticosterone levels with age. Body condition was negatively correlated with monocyte numbers, suggesting a poorer health status of penguin chicks in poorer body condition. Yet, there was no link between body condition and other leucocyte parameters, indicating that chicks in a good body condition did not additionally invest into their immune system.     

Is cell-mediated immunity related to the evolution of life-history strategies in birds?

According to life-history theory, the development of immune function should be balanced through evolutionary optimization of the allocation of resources to reproduction and through mechanisms that promote survival. We investigated interspecific variability in cell-mediated immune response (CMI), as measured by the phytohaemagglutinin (PHA) assay, in relation to clutch size, longevity and other life-history traits in 50 species of birds. CMI exhibited significant repeatability within species, and PHA responses in chicks were consistently stronger than in adults. Univariate tests showed a variety of significant relationships between the CMI of both chicks and adults with respect to size, development period and lifespan, but not clutch size or prevalence of blood parasites in adults. Multivariate analyses confirmed these patterns but independent variables were too highly correlated to isolate unique influences on CMI. The positive relationship of chick CMI to nestling period is further complicated by a parallel relationship of chick CMI to the age at testing. However, multivariate analysis showed that chick CMI varies uniquely with length of the nestling period. Adult CMI was associated with a strong life-history axis of body size, development rate and longevity. Therefore, adult CMI may be associated with prevention and repair mechanisms related to long lifespan, but it also may be allometrically related to body size through other pathways. Neither chick CMI nor adult CMI was related to clutch size, contradicting previous results linking parasite-related mortality to CMI and the evolution of clutch size (reproductive investment) in birds.     

Growth, developmental stability and immune response in juvenile Japanese quails (Coturnix coturnix japonica).

Stresses are environmental factors which restrict growth or cause a potentially adverse change in an organism. The exposure of developing organisms to environmental stresses may have several physiological consequences including a decrease in immunocompetence. However, mounting an immune response against a foreign antigen may in itself constitute a cost for developing organisms. This cost has potentially long-term consequences for adult function and fitness. This study examines the growth and developmental stability of Japanese quail++ chicks challenged by three non-pathogenic antigens: sheep red blood cells, which assess T-cell-dependent immune responses, and Mycoplasma synoviae and Newcastle disease virus, which assess T-cell-independent responses. Increases in both body mass and wing length were significantly reduced in antigen-challenged birds compared to control birds. Fluctuating asymmetry (FA) in the masses of primary feathers increased from the innermost (1) to the outermost (10) position on the wing. In addition, antigen challenge by M. synoviae and sheep red blood cells was associated with an increase in FA. The cell-mediated response measured by reaction to phytohaemagglutinin was significantly depressed in M. synoviae-challenged birds. White blood cell counts, except for monocytes, were elevated in response to all three antigen treatments. Total plasma protein and haematocrit also differed between treatments but exhibited no clear relationship to antigen challenge. Immune responses clearly impose a stress on developing chicks. Additional research will be required to determine the long-term consequences of developmental stress and assess the selective forces that influence the strength of the immune responses of chicks.     

Chick parasitism by blowflies affects feeding rates in a Mediterranean population of blue tits

Offspring fitness depends on interactions between parental care and environmental constraints. It has been suggested that in altricial birds parents are able to compensate for the detrimental effects of ectoparasites by improving food provisioning. We tested this prediction in a population of blue tits highly parasitized by blowfly larvae. The frequency of parental feeding visits was significantly higher in parasitized broods than in broods experimentally deparasitized. Despite a strong increase in parental care, chicks of parasitized broods were lighter, smaller, and more anaemic than chicks in deparasitized broods. Parents invest more in feeding parasitized young but cannot fully compensate for the negative effects of parasites, hence young are in poor condition at fledging.     

Experimental assessment of the effects of gastrointestinal parasites on offspring quality in chinstrap penguins (Pygoscelis antarctica)

Parasites reduce host fitness and consequently impose strong selection pressures on their hosts. It has been hypothesized that parasites are scarcer and their overall effect on hosts is weaker at higher latitudes. Although Antarctic birds have relatively low numbers of parasites, their effect on host fitness has rarely been investigated. The effect of helminth parasitism on growth rate was experimentally studied in chinstrap penguin (Pygoscelis antarctica) nestlings. In a total of 22 two-nestling broods, 1 nestling was treated with anthelminthics (for cestodes and nematodes) while its sibling was left as a control. Increased growth rate was predicted in de-wormed nestlings compared to their siblings. As expected, 15 days after treatment, the experimental nestlings had increased body mass more than their siblings. These results show a non-negligible negative effect of helminth parasites on nestling body condition that would presumably affect future survival and thus fitness, and it has been suggested there is a strong relationship between body mass and mortality in chinstrap penguins.     

A comprehensive demographic assessment of the endangered Fiordland crested penguin Eudyptes pachyrhynchus

ABSTRACT

Many species of Eudyptes penguin have shown substantial population declines and in response, there have been efforts to identify the key demographic parameters. Here, we present the demographic parameters of one of the least well known and the least abundant species of crested penguin, the endangered Fiordland crested penguin Eudyptes pachyrhynchus. A population study incorporating mark–recapture, nest occupancy and breeding success was conducted over 16 years at several sites in the northern half of the range of the species. Survival probabilities were calculated using standard Cormack–Jolly–Seber models and the Burnham Live and Dead model. The annual probability of true survival for banded birds and apparent survival for birds with transponders were both estimated at 89% during their adult years, which is similar to that reported for Eudyptes penguin species inhabiting more southerly latitudes. Annual juvenile survival was assessed for Fiordland crested penguins until their first return at 77%. The mean breeding success (0.61?±?0.02 chicks/pair) was higher than is observed for other crested penguin species, except the southern rockhopper penguin, which may be due to having lower A-egg ejection rates and higher rates of fledging two chicks per pair. Breeding success was related to the niche of predators present.     

Secondary phytohaemagglutinin (PHA) swelling response is a good indicator of T‐cell‐mediated immunity in free‐living birds

The validity of using the phytohaemagglutinin (PHA) test to measure acquired immunity, one of the most widely used methods, is currently being debated due to new knowledge on the complex physiology of the process. As a greater secondary response to repeated challenges linked to increases of circulating lymphocyte levels would be indicative of a T‐cell‐mediated immune response, we performed for the first time an experiment under natural conditions with repeated PHA challenges in free‐living adult birds and chicks to shed light on this topic. We found significantly stronger secondary response to PHA injection independent of sex or age, while controlling for body condition, the second response being on average 90% larger than the first. Likewise, lymphocyte counts were significantly higher in the second PHA challenge, whereas no significant differences were found among untreated birds. Significant positive correlations between the PHA response and both lymphocyte counts and plasma protein levels (mainly albumin, globulin precursor) were recovered, whereas no significant differences were recovered in plasma protein levels between challenges. Our results are consistent with those from captive birds, supporting the validity of the PHA skin‐swelling test as an accurate gauge of acquired T‐cell‐mediated immunity in birds.     

Do individual branches of immune defence correlate? A comparative case study of scavenging and non‐scavenging birds

Costs of immunity are widely believed to play an important role in life history evolution, but most studies of ecological immunology have considered only single aspects of immune function. It is unclear whether we should expect correlated responses in other aspects of immune function not measured, because individual branches of immune defence may differ in their running costs and thus may compete unequally for limiting resources, resulting in negatively correlated evolution. In theory such selection pressure may be most intense where species are hosts to more virulent parasites, thus facing a higher potential cost of parasitism. These issues are relatively unstudied, but could influence the efficacy of attempting to estimate the scale and cost of host investment in immune defence. Here, in a comparative study of birds we found that species that scavenge at carcasses, that were hypothesised to be hosts to virulent parasites, had larger spleens for their body size and higher blood total leukocyte concentrations (general measures of immune function) than non-scavengers. These results support the hypothesis that scavengers are subject to strong parasite-mediated selection on immune defences. However, measures of specific branches of immune function revealed that scavengers had a relatively lower proportion of lymphocytes than phagocytic types of leukocytes, suggesting robust front line immune defences that could potentially reduce the need for mounting relatively energetically costly lymphocyte-dependent immune responses. Following experimental inoculation, scavengers produced significantly larger humoral immune responses, but not cell-mediated immune responses, than non-scavengers. However, the sizes of cell-mediated and humoral immune responses were not correlated across species. These results suggest that single measures of immune defence may not characterise the overall immune strategy, or reveal the likely costs involved.     

Effects of ectoparasites of young on parents' behaviour in a Mediterranean population of Blue Tits

In altricial birds, adults are expected to alleviate the impact of parasitism on their young by increasing parental care. We experimentally investigated the effects of blowfly parasitism on adult behaviour in the nest cavity in a population of Blue Tits in which nestlings are heavily parasitized by blowfly larvae. The behaviour of males and females was different, since only the females carried out nest sanitation activities related to the presence of parasites. The time females devoted to nest sanitation was higher in parasitized nests than in experimentally deparasitized ones, suggesting that chicks and adults might share the costs of blowfly parasitism.     

Health impact of blood parasites in breeding great tits

Hypotheses of hemoparasite-mediated sexual selection and reproductive costs rely on the assumption that avian blood parasite infections are harmful to their hosts. To test the validity of this assumption, we examined the health impact of Haemoproteus blood parasites on their great tit (Parus major) host. We hypothesised that if blood parasites impose any serious health impact on their avian hosts, then infected individuals must differ from uninfected ones in respect to hemato-serological general health and immune parameters. A 3-year study of two great tit populations, breeding in contrasting (urban and rural) habitats in south-east Estonia, revealed that Haemoproteus blood parasites affected the health state of their avian hosts. Infected individuals had elevated lymphocyte hemoconcentration and plasma gamma-globulin levels, indicating that both cell-mediated and humoral immune response mechanisms are involved in host defence. The effect of parasites on cell-mediated immunity was both age- and sex-specific, as infection status affected peripheral blood lymphocyte counts only in males, and among these, the magnitude of response was greater in old individuals than yearlings. Heterophile hemoconcentration and plasma albumin levels were not affected by infection status, suggesting that blood stages of Haemoproteus infection do not cause a severe inflammatory response. Parasitism was not related to hematocrit values, indicating that Haemoproteus infection does not cause anemia. In two years, infected individuals were heavier than uninfected ones in the urban but not in the rural study area. This suggests, that under certain circumstances (possibly related to reproductive tactics), breeding great tits may avoid losing body mass in order to save resources for an anti-parasite immune response. Received: 16 February 1998 / Accepted: 22 May 1998     

Coevolution of host immune defence and parasite-induced mortality: relative spleen size and mortality in altricial birds

Animal hosts defend themselves against parasites by the antibodies produced by an immune system. It is an inherent assumption that parasites constitute the selection pressure that has given rise to and maintains the immune system. Thus, greater impact by parasites on host fitness should result in greater investment in immune function across species. We tested this prediction by using field estimates of parasite-induced nestling mortality in altricial birds as an estimate of the fitness cost of parasitism and the relative size of the spleen as an estimate of investment in immune function. The spleen is a peripheral lymphoid tissue that acts as the main site of lymphocyte differentiation and proliferation, and these B- and T-cells are involved in the production of humoral and cell-mediated immune responses. In a comparative study of 21 species of altricial birds we found a significant positive relationship between relative spleen size and parasite-induced mortality, accounting for a third of the variance, even when controlling for potentially confounding variables. This finding provides evidence for the level of investment in immune function being related to the natural selection pressure imposed by parasites.     

Cell-mediated immunosenescence in birds

The phytohaemagglutinin (PHA) skin test response, used to assess cell-mediated immunity, is known to vary with many social and energetic factors, but the effects of age have received little attention. We found that the PHA response of immature birds was lower than those of the youngest breeding adults and were decreased in adults. Whenever possible, age should be included as a covariate when the PHA skin test is used to assess immunocompetence in ecological immunology. The rate of decline in PHA response differed between species and was inversely correlated with survival. The decrease in the PHA response averaged 57% over an average 80% of the maximum life span, but the absolute rate varied with species lifespan such that the short-lived species showed a greater loss per year than the long-lived species. This link between declining immune function and survival may reflect differences in resource partitioning between species, and suggests that selection may act on investment in immune function to influence maximum life span.     

Hand-Rearing,Release and Survival of African Penguin Chicks Abandoned Before Independence by Moulting Parents

The African penguin Spheniscus demersus has an ‘Endangered’ conservation status and a decreasing population. Following abandonment, 841 African penguin chicks in 2006 and 481 in 2007 were admitted to SANCCOB (Southern African Foundation for the Conservation of Coastal Birds) for hand-rearing from colonies in the Western Cape, South Africa, after large numbers of breeding adults commenced moult with chicks still in the nest. Of those admitted, 91% and 73% respectively were released into the wild. There were veterinary concerns about avian malaria, airsacculitis and pneumonia, feather-loss and pododermatitis (bumblefoot). Post-release juvenile (0.32, s.e.  = 0.08) and adult (0.76, s.e.  = 0.10) survival rates were similar to African penguin chicks reared after oil spills and to recent survival rates recorded for naturally-reared birds. By December 2012, 12 birds had bred, six at their colony of origin, and the apparent recruitment rate was 0.11 (s.e.  = 0.03). Hand-rearing of abandoned penguin chicks is recommended as a conservation tool to limit mortality and to bolster the population at specific colonies. The feasibility of conservation translocations for the creation of new colonies for this species using hand-reared chicks warrants investigation. Any such programme would be predicated on adequate disease surveillance programmes established to minimise the risk of disease introduction to wild birds.     

Bacteria divert resources from growth for magellanic penguin chicks

The influence of bacteria on the growth of their wild avian hosts is unknown. We tested experimentally whether administration of a wide‐spectrum antibiotic (cephalosporine) during early development of magellanic penguin (Spheniscus magellanicus) chicks had any effect on their growth rates in the wild. Chicks that were injected in two occasions with cephalosporine grew faster than control untreated chicks. The positive effect of medication on nestling growth disappeared after the treatment ceased, did not alter haematological indices indicative of health status, had no influence on chick survival until near independence and was related to a changed bacterial composition of the faecal microbiota of treated chicks when compared with that from control chicks. These results were similar to those obtained for poultry with antimicrobials promoting growth and chick nutrient assimilation rates. Gram‐positive bacilli in the diphtheroid genus Corynebacterium are likely candidates to cause decreased growth rates in magellanic penguin chicks.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

High blowfly parasitic loads affect breeding success in a Mediterranean population of blue tits

The detrimental effects of ectoparasites on the breeding success of birds have been especially well demonstrated in the case of ectoparasites that affect both chicks and their parents. Since blowfly larvae of the genus Protocalliphora attack only nestling birds, they represent a good model for testing the consequences of parasitism on nestlings. A Corsican population of blue tit suffers extremely high rates of infestation by blowflies, which are suspected to negatively affect young birds. Comparing experimentally deparasitized (treated) and naturally infested (control) broods, we showed that the attack by Protocalliphora causes anaemia and an important disturbance to the chicks. Therefore, we expected that these effects would have a negative impact on body condition and survival in the infested broods. Although we did not find any effect of treatment on fledging success, our predictions were confirmed by lower growth rate, body mass at fledging and tarsus length at fledging in the control compared with the treated group. This suggests that in this population, blowflies decrease the probability of recruitment of young blue tits. Received: 6 May 1997 / Accepted: 14 July 1997     

Energetic costs of raising Pygoscelid penguin chicks

Energy requirements of resting Adélie (Pygoscelis adeliae), Gentoo (P. papua) and Chinstrap (P. antarcticd) penguin chicks were determined with respect to body mass via respirometry in Antarctica. Resting metabolic rates of all Pygoscelid penguin chicks were similar (ANOVA p=0.91) and best described by E=0.0096 M^0.98 (n=24, r²=0.97), where E is power (W) and M is mass (g). Using the results obtained here and data published in the literature, I determined a) the amount of food needed from hatching to fledging as 29.8, 31.7 and 56.4 kg per chick for Adélie, Chinstrap and Gentoo penguins, respectively and b) the average amount of food left daily to the parent after feeding the brood throughout the breeding period. Parents keep only a minimum of food for themselves just prior to the time when chicks begin forming crèches. Thereafter, nest relief intervals are increased, and the amount of food parents can keep for themselves rises. The results of both models are discussed with respect to available data on Pygoscelid penguin food requirements.     

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.