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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.

The rare Egyptian vulture (Neophron percnopterus) stands out among the Old World vultures (Family Accipitridae) because of its brightly ornamented head, which is coloured yellow by carotenoid pigments, and its practice of feeding on faeces. Here we show that Egyptian vultures obtain these pigments from the excrement of ungulates. To our knowledge, this is the first demonstration that faeces can be used as a source of carotenoids by a vertebrate.

Feather mites (Astigmata) are distributed around the world, living on the feathers of birds, but their mechanisms for transmission among hosts are not fully understood. There is anecdotal evidence of feather mites attached to louseflies (Diptera: Hippoboscidae), suggesting that feather mites may use these flies as a mode of phoretic transmission among birds. Two bird-lousefly associations (alpine swift Apus melba-Crataerina melbae and feral pigeon Columba livia-Pseudolynchia canariensis) were inspected to test the hypothesis that feather mites use hippoboscid flies as major mode of transmission. Both bird species showed a high prevalence and abundance of feather mites and louseflies. However, no feather mites were found attached to the 405 louseflies inspected, although skin mites (Epidermoptidae and Cheyletiellidae) were found on louseflies collected from feral pigeons. This study suggests that feather mites do not use hippoboscid flies as a major mode of transmission among birds.

Why avian colonies vary in size and how food competition among nearby colonies affects offspring quality are still not completely understood. We simultaneously examined the effects of four scales of breeding density on two measures of offspring viability (body condition and T-cell-mediated immunity) in the colonial Magellanic penguin. Body condition of fledglings was inversely correlated with breeding density within 100 m(2) of nests, and decreased with increasing numbers of breeding pairs competing within the parental foraging ranges (100 km), probably as a result of density-dependent food depletion. The T-cell-mediated immune response was positively correlated with body condition, reflecting, to some extent, the previous breeding-density effects, and was negatively correlated with colony size, which may be related to social stress. However, given the effect of protein intake on cell immunity, this result could also indicate a thus far neglected cost of coloniality, namely the consumption of low-protein food to compensate for the depletion of optimal prey. These results were not influenced by other traits, nor by the current exposure of birds to parasites and diseases, as measured by serological variables. Since body condition and the T-cell-mediated immune response of fledgling birds are indicators of their survival and recruitment prospects, the costs we have identified can explain variability in colony size in relation to food competition with surrounding colonies, as well as the skewed distribution toward small colonies in this species.

Single cell gel electrophoresis, the so-called "Comet" assay, was performed as a genotoxicity test in white storks sampled in an area heavily contaminated after the ecological disaster in south western Spain. This disaster occurred as a consequence of a massive toxic spillage of acid waste rich in heavy metals that impacted on the Doñana National Park. The importance of this protected area as a breeding and wintering site for many endangered bird species makes this analysis of DNA damage of special interest. Our results clearly show that white storks born in the contaminated area 1 year after the toxic spill bear a high burden of genetic damage as compared with control individuals. The possible implications for future survival as well as reproductive rate are discussed.

In April 1998, an ecological disaster resulting from a massive toxic spill of mining acid waste rich in heavy metals posed a serious threat to the Doñana National Park in southwestern Spain. This especially important protected area is the nesting and breeding site for many endangered bird species; white storks (Ciconia ciconia) and black kites (Milvus migrans) are considered the more representative. The suitability of the Comet assay as a biomarker for genotoxic analysis in environmental biomonitoring has been recently validated in studies using different sentinel organisms such as fish, amphibians, rodents, or mollusks. Birds preying on a variety of invertebrate and vertebrate species in the marshlands are appropriate for evaluating the potential deleterious effects of the toxic spill on wildlife of the Dofiana area. Our study on wetland birds high on the aquatic trophic chain sampled within a few months after the toxic spill in the area around Doñana National Park has shown the accumulation of heavy metals. Fourteen months after the mine waste spill, blood samples from white storks and kites collected in the neighborhood of the park and from control birds at reference areas for comparison were examined by fluorescence image analysis after lymphocyte isolation, and by subsequent alkaline single-cell gel (SCG) electrophoresis, known as the Comet assay. Our results indicate that the exposed birds had a significantly increased level of genotoxic damage compared with control animals from noncontaminated locations.

Aggregation of Crataerina melbae flies on breeding adult alpine swifts (Apus melba) was low when compared with other host-parasite systems and varied with sampling date, year, and sex of the flies. Generalized linear models were performed to ascertain which factors, extrinsic and/or intrinsic to the host, explained variability in the number of louse flies present on a single host, i.e., abundance. Overall abundance was unrelated to any host characteristic but varied slightly among years. Abundance of female flies varied among years, but also with date of sampling, the number of females increasing as the breeding season advanced. In contrast, abundance of males decreased as the season progressed, independently of host characteristics. Despite these different patterns, the number of flies of each sex on a given host was strongly intercorrelated. These results suggest that mate attraction may explain aggregation patterns in this louse fly species. Overall sex ratio of louse flies did not differ from unity. However, the proportion of males decreased during the breeding season, as a consequence of the opposite sex-related seasonal patterns in parasite abundance. Sex-ratio variability was not related to host characteristics or to infrapopulation sizes.

Carotenoids are important as pigments for bright coloration of animals, and as physiologically active compounds with a wide array of health-related functions. Carotenoid-dependent coloration may have evolved as a signal to conspecifics; however, factors that may limit availability of carotenoids are poorly known. We investigated how the acquisition of carotenoids may be constrained by availability in the environment, diet, genetic make-up and health status of wild American kestrels (Falco sparverius). Plasma concentrations of siblings at the time of fledging showed a high degree of resemblance; however, a cross-fostering experiment revealed that variance was largely explained by nest of rearing, rather than nest of origin, thus indicating a low genetic component. A multivariate analysis of attributes of nestlings (sex, size, plasma proteins, immune function), parental reproduction (laying date, clutch size) and rearing conditions (brood size, size hierarchy, nestling mortality) showed only a small significant effect of leucocyte differentials on carotenoid concentrations of nestlings. A strong environmental effect on plasma carotenoids was demonstrated by levels of adult kestrels being correlated within mated pairs, and having a significant association with the abundance of voles, the primary prey species, per territory.

We investigated genetic and environmental components of variance in avian T-cell-mediated immune response (CMI) through a cross-fostering experiment conducted on wild American kestrels (Falco sparverius). CMI was evaluated in vivo by an experimental challenge with phytohaemagglutinin, a T-cell mitogen, injected intradermally in fledglings. Additionally, we assessed two measures of nutritional condition (body mass and circulating plasma proteins) which could influence the variance components of CMI. A two-way nested ANOVA indicated that CMI of fledgling kestrels was explained more by the nest where the bird was reared (33% of the explained variance) than by the nest of origin (12%). Body mass was explained equally by familial and environmental components, while plasma proteins were only related to the rearing environment. CMI of fledglings was not related to their circulating plasma proteins, but was positively correlated with their body mass. Fledgling body mass seemed to be influenced by pre-hatching or post-hatching maternal effects prior to manipulation since resemblance in body mass of sibships at the age of manipulation was high (h 2 ≤0.58), and body mass at this age predicted body mass at fledging. Therefore, pre-manipulation parental effects on body mass, such as investment in egg size, could have inflated the familial effects on body mass of fledglings and then on its correlated CMI. When controlling for body mass, most of the variation in CMI of fledglings was explained by the nest where the bird was reared (36.6%), while the variance explained by the nest of origin (4%) was not significant. This means that environmental influences are major determinants of offspring CMI. The low proportion of variance explained by the familial component may have been due to the high correlation of CMI to fitness.

Life-history theory predicts that parents face a trade-off between the number and viability of the progeny they produce. We found evidence for an apparent trade-off in a free-living population of American kestrels (Falco sparverius), as larger clutches produced more but lighter fledglings. However, while the body mass of fledglings has traditionally been used as a measure of survival prospect, offspring immunocompetence should also play an important role. We thus measured the T-cell-mediated immune response of fledgling kestrels in relation to brood traits and nest-rearing conditions through a cross-fostering experiment. The immune response was positively correlated with the body condition of fledglings, but was also higher in those hatched from five-egg than four-egg clutches. These results were not influenced by other brood traits, nor by current exposure to stressors and infectious agents, as measured by serological variables. Such ability to resist pathogens may account for why the probability of offspring returning to the study area in subsequent years, when controlling for brood size, was higher for five-egg than four-egg clutches. These results suggest an optimal clutch size through maternal effects on offspring immunocompetence rather than a trade-off between the number and quality of the offspring.

Louse flies were collected from 401 birds of 32 species captured in autumn of 1996 in Baja California Sur (Mexico). Only one louse fly species (Microlynchia pusilla) was found. It occurred in four of the 164 common ground doves (Columbina passerina) collected. This is a new a host species for this louse fly.

Carotenoids have a dietary origin in birds, but mechanisms by which they are absorbed in the gut, transported in the blood, metabolized at various sites, and deposited in the integument remain poorly understood. Variation in both plasma carotenoid levels and external color may reflect different access to dietary carotenoids or individual physiological differences in the uptake and deposition of carotenoids. We compared total plasma carotenoid concentration in nestling white storks (Ciconia ciconia) from 11 Spanish colonies in two consecutive years. The main food item in one of the colonies was the red swamp crayfish (Procambarus clarkii), a recently introduced species. Storks in the remaining colonies ate a variety of foods but no crayfish. Total plasma carotenoid levels in the colony where crayfish were consumed were about five times higher than in any other colony. These differences were maintained after controlling for the significant interyear variability, as well as for sex, age, and body mass of birds. Skin pigmentation also differed, being intensely orange in storks that consumed crayfish but white (unpigmented) in the remaining individuals. With thin-layer chromatography (TLC) and electronic absorption spectroscopy, astaxanthin was confirmed as the major carotenoid in crayfish as well as in the plasma, skin, and body fat of crayfish-eating storks, whereas lutein was the main carotenoid in plasma samples from the other colonies. These results indicate that a newly available carotenoid in the environment, astaxanthin, can be absorbed in large quantities from the gut and be transported in the blood before deposition in different tissues.

We assessed the value of communal roosting for mating and territory acquisition in nonbreeding red-billed choughs, Pyrrhocorax pyrrhocorax. Between 1991 and 1996 we surveyed all communal roosts, breeding territories and vacant nest sites in an area of 250 km2in Los Monegros (northeast Spain), as well as other communal roosts in the surrounding 500 km2from 1987 to 1996. Main roosts were used traditionally and throughout the year and gathered more choughs than subroosts, which were used only during the nonbreeding season. Nonbreeding choughs were socially segregated in these roosts according to their age and breeding prospects. Compared with random potential roost sites, main roosts included more first-year birds and were in areas of higher breeding density. Subroosts were mainly joined by choughs of breeding age (at least 2 years old) whose morphology (males) and body condition (females) were similar to those of established breeders, and were in areas with breeding densities as high as around main roosts but with more available nest sites. Nonbreeding choughs that used main roosts later used subroosts, while the converse was rare. More choughs joining subroosts acquired mates than those joining main roosts. Choughs mostly paired with roostmates but also with widowed territorial breeders. Finally, choughs from subroosts acquired territories closer to the roost than those who mated in main roosts, supporting the hypothesis that subroosts reduce the costs of mating and territory acquisition. Copyright 1999 The Association for the Study of Animal Behaviour.

The factors explaining interspecific differences in prevalences of blood parasites in birds are poorly known. We simultaneously assessed 20 social, ecological, life history, and sampling-related variables that could influence hemoparasite prevalences among diurnal birds of prey in Spain. Our results show that multiple factors are responsible for the studied host-parasite association. We confirmed for the first time that prevalence is inversely correlated to the embryonic development period, and thus probably to immune performance, even among closely related birds. Macrohabitat features related to vector availability are also important, prevalences being higher in species breeding in forested habitats. Finally, prevalence is positively correlated with the host's world geographic range. We hypothesize that larger geographic ranges offered more opportunities for host-vector-hemoparasite associations to become established. The results from our multivariate analyses differ from those obtained through univariate ones, showing that all potential factors should be assessed jointly when testing any ecological or evolutionary hypothesis dealing with parasites.

The study of the role of carotenoids on the physiology and evolutionary ecology of birds demands methods for their quantification in the bloodstream. We compared color-chart scores of plasma hue with the actual concentration of plasma carotenoids obtained by spectrophotometry in 356 wild birds from 26 species. Repeatability of chart scores between three independent observers was high. However, color scores did not correlate with the spectrophotometric results in interspecific analyses. Within species (n = 3), one showed no relationship and two showed weak but significant positive correlations. Hemoglobin, and probably other substances, may mask the color of carotenoids, making the accurate use of color charts difficult. Spectrophotometry should be the method of choice as it permits precise quantifications of total plasma carotenoids and objective comparisons among studies.

We looked for louse flies (Diptera, Hippoboscidae) in a mixed colony of 9 species of birds from 1991 to 1997. Alpine swifts (Apus melba) exhibited an unusually high prevalence (85.9%) by Crataerina melbae (Rondani, 1879). No birds from the other 8 breeding bird species were parasitized by this louse fly. We suggest that the number of potential swiftlike hosts as well as the size of hosts may account for the differential prevalence of the louse flies within the colony.

Some bird species may breed close to aggressive predators to reduce predation risk by more dangerous, generalist predators. We tested this protective nesting association hypothesis by studying solitary choughs, Pyrrhocorax pyrrhocoraxbreeding within or outside lesser kestrel, Falco naumannicolonies in northern Spain (1993-1994). We found 27 potential predators of choughs and kestrels and confirmed predation on adults and nests of both species by at least eight common predators. We experimentally assessed the defence investment of choughs and kestrels towards a stuffed eagle owl, Bubo buboin 19 buildings shared by both species. Lesser kestrels were more efficient at detecting the predator, and defended more vigorously than choughs. Choughs clearly selected for breeding buildings where lesser kestrel colonies were installed. Breeding success of these choughs was much higher than that of choughs breeding alone, because of a lower percentage of nest failure. Benefits to choughs probably accrued from both the 'parasitism' of the kestrels' nest defence and the dilution of predation risk in the colonies. Since lesser kestrels do not prey on choughs, this could be a good example of protective nesting association in birds.

The prevalence of hematozoa in two populations of red-billed choughs (Pyrrhocorax pyrrhocorax) was sampled in 1992 and 1994 in Spain. Two blood parasites infected red-billed choughs. A species of Plasmodium, possibly Plasmodium relictum, and the piroplasm Babesia frugilegica, are described for the first time from this host. Low prevalence (1/178, < 1%) of hematozoa in these populations, was evidence for a lack of effects of blood parasites on the life history and conservation of this threatened species in at least the two populations studied.