Early Life Conditions and Immune Defense in Nestling Swainson's Hawks.

The quality of perinatal conditions directly influences the physical and immunological development of nestlings, yet it is inherently variable across space and time. Long-term breeding data for a population of Swainson's hawks (Buteo swainsoni) in northern California show a continuum of territory occupancy and productivity values of individual territories and nests. Here we explore effects of variation among territories on immune system development. We hypothesize that nestlings benefitting from favorable conditions will invest in stronger immune systems, a trait with long-term benefits. We used two immunological assays, a bactericidal assay and a hemolytic-complement activity assay, with leukocyte differentials (heterophil∶lymphocyte ratio) to evaluate the constitutive innate immune system. We examined whether early brood-rearing conditions (i.e., number of siblings, hatch date, endoparasite prevalence) were associated with immunological development. Linear mixed-effects models indicated a positive relationship between extended territory occupancy history-an index of habitat quality-and nestling immune function during years with poorer reproduction. There was no association during an exceptionally good reproductive year. Hence, at least under some circumstances, nestling environments or territory characteristics may affect immune function of nestlings. Our study contributes to the growing body of evidence highlighting the importance of facultative allocation to immune traits using long-term demographic data of a top avian predator.

Pheomelanin synthesis varies with protein food abundance in developing goshawks.

The accumulation of the amino acid cysteine in lysosomes produces toxic substances, which are avoided by a gene (CTNS) coding for a transporter that pumps cystine out of lysosomes. Melanosomes are lysosome-related organelles that synthesize melanins, the most widespread pigments in animals. The synthesis of the orange melanin, termed pheomelanin, depends on cysteine levels because the sulfhydryl group is used to form the pigment. Pheomelanin synthesis may, therefore, be affected by cysteine homeostasis, although this has never been explored in a natural system. As diet is an important source of cysteine, here we indirectly tested for such an effect by searching for an association between food abundance and pheomelanin content of feathers in a wild population of Northern goshawk Accipiter gentilis. As predicted on the basis that CTNS expression may inhibit pheomelanin synthesis and increase with food abundance as previously found in other strictly carnivorous birds, we found that the feather pheomelanin content in nestling goshawks, but not in adults, decreased as the abundance of prey available to them increased. In contrast, variation in the feather content of the non-sulphurated melanin form (eumelanin) was only explained by sex in both nestlings and adults. We also found that the feather pheomelanin content of nestlings was negatively related to that of their mothers, suggesting a relevant environmental influence on pheomelanin synthesis. Overall, our findings suggest that variation in pheomelanin synthesis may be a side effect of the maintenance of cysteine homeostasis. This may help explaining variability in the expression of pigmented phenotypes.

Climatic conditions cause complex patterns of covariation between demographic traits in a long-lived raptor.

Environmental variation can induce life-history changes that can last over a large part of the lifetime of an organism. If multiple demographic traits are affected, expected changes in climate may influence environmental covariances among traits in a complex manner. Thus, examining the consequences of environmental fluctuations requires that individual information at multiple life stages is available, which is particularly challenging in long-lived species. Here, we analyse how variation in climatic conditions occurring in the year of hatching of female goshawks Accipiter gentilis (L.) affects age-specific variation in demographic traits and lifetime reproductive success (LRS). LRS decreased with increasing temperature in April in the year of hatching, due to lower breeding frequency and shorter reproductive life span. In contrast, the probability for a female to successfully breed was higher in years with a warm April, but lower LRS of the offspring in these years generated a negative covariance among fecundity rates among generations. The mechanism by which climatic conditions generated cohort effects was likely through influencing the quality of the breeding segment of the population in a given year, as the proportion of pigeons in the diet during the breeding period was positively related to annual and LRS, and the diet of adult females that hatched in warm years contained fewer pigeons. Climatic conditions experienced during different stages of individual life histories caused complex patterns of environmental covariance among demographic traits even across generations. Such environmental covariances may either buffer or amplify impacts of climate change on population growth, emphasizing the importance of considering demographic changes during the complete life history of individuals when predicting the effect of climatic change on population dynamics of long-lived species.

Diet shift induced rapid evolution of size and function in a predatory bird.

A predator's body size correlates with its prey size. Change in the diet may call for changes in the hunting mode and traits determining hunting success. We explored long-term trends in sternum size and shape in the northern goshawk by applying geometric morphometrics. Tetraonids, the primary prey of the goshawk, have decreased and been replaced by smaller birds in the diet. We expected that the size of the goshawk has decreased accordingly more in males than females based on earlier observations of outer morphology. We also expected changes in sternum shape as a function of changes in hunting mode. Size of both sexes has decreased during the preceding decades (1962-2008), seemingly reflecting a shift in prey size and hunting mode. Female goshawks hunting also mammalian prey tend to have a pronouncedly "Buteo-type" sternum compared to males preying upon birds. Interestingly, the shrinkage of body size resulted in an increasingly "Buteo-type" sternum in both sexes. In addition, the sternum shape in birds that died accidentally (i.e., fit individuals) was more Buteo-type than in starved ones, hinting that selection was towards a Buteo-type sternum shape. We conclude that these observed patterns are likely due to directional selection driven by changes in the diet towards smaller and more agile prey. On the other hand, global warming is predicted to also cause a decrease in size, thus these two scenarios are inseparable. Because of difficulties in studying fitness-related phenotypic changes of large raptors in the field, time series of museum exemplars collected over a wide geographical area may give answers to this conundrum.

Density-dependent regulation of population size in colonial breeders: Allee and buffer effects in the migratory Montagu's harrier.

Expanding populations offer an opportunity to uncover the processes driving spatial variation in distribution and abundance. Individual settlement decisions will be influenced by the availability and relative quality of patches, and by how these respond to changes in conspecific density. For example, conspecific presence can alter patch suitability through reductions in resource availability or territorial exclusion, leading to buffer effect patterns of disproportionate population expansion into poorer quality areas. However, conspecific presence can also enhance patch suitability through Allee effect processes, such as transmission of information about resources or improved predator detection and deterrence. Here, we explore the factors underlying the settlement pattern of a growing population of Montagu's harriers (Circus pygargus) in Spain. The population increased exponentially between 1981 and 2001, but stabilised between 2001 and 2004. This population increase occurred alongside a remarkable spatial expansion, with novel site use occurring prior to maximum densities in occupied sites being reached. However, no temporal trends in fecundity were observed and, within sites, average fecundity did not decline with increasing density. Across the population, variance in productivity did increase with population size, suggesting a complex pattern of density-dependent costs and benefits. We suggest that both Allee and buffer effects are operating in this system, with the benefits of conspecific presence counteracting density-dependent declines in resource availability or quality.