Ontogeny of migration destination, route and timing in a partially migratory bird.

In migratory animals, the developmental period from inexperienced juveniles to breeding adults could be a key life stage in shaping population migration patterns. Nevertheless, the development of migration routines in early life remains underexplored. While age-related changes in migration routes and timing have been described in obligate migrants, most investigations into the ontogeny of partial migrants only focused on age-dependency of migration as a binary tactic (migrant or resident), and variations in routes and timing among individuals classified as 'migrants' is rarely considered. To fill this gap, we study the ontogeny of migration destination, route and timing in a partially migratory red kite (Milvus milvus) population. Using an extensive GPS-tracking dataset (292 fledglings and 38 adults, with 1-5 migrations tracked per individual), we studied how nine different migration characteristics changed with age and breeding status in migrant individuals, many of which become resident later in life. Individuals departed later from and arrived earlier at the breeding areas as they aged, resulting in a gradual prolongation of stay in the breeding area by 2 months from the first to the fifth migration. Individuals delayed southward migration in the year prior to territory acquirement, and they further delayed it after occupying a territory. Migration routes became more direct with age. Individuals were highly faithful to their wintering site. Migration distance shortened only slightly with age and was more similar among siblings than among unrelated individuals. The large gradual changes in northward and southward migrations suggest a high degree of plasticity in temporal characteristics during the developmental window. However, the high wintering site fidelity points towards large benefits of site familiarity, prompting spatial migratory plasticity to be expressed through a switch to residency. The contrasting patterns of trajectories of age-related changes between spatial and temporal migration characteristics might reflect different mechanisms underlying the expression of plasticity. Investigating such patterns among species along the entire spectrum of migration tactics would enable further understanding of the plastic responses exhibited by migratory species to rapid environmental changes.

Determinants of departure to natal dispersal across an elevational gradient in a long-lived raptor species.

Attributes of natal habitat often affect early stages of natal dispersal. Thus, environmental gradients at mountain slopes are expected to result in gradients of dispersal behavior and to drive elevational differences in dispersal distances and settlement behavior. However, covariation of environmental factors across elevational gradients complicates the identification of mechanisms underlying the elevational patterns in dispersal behavior. Assuming a decreasing food availability with elevation, we conducted a food supplementation experiment of red kite (Milvus milvus) broods across an elevational gradient toward the upper range margin and we GPS-tagged nestlings to assess their start of dispersal. While considering timing of breeding and breeding density across elevation, this allowed disentangling effects of elevational food gradients from co-varying environmental gradients on the age at departure from the natal home range. We found an effect of food supplementation on age at departure, but no elevational gradient in the effect of food supplementation. Similarly, we found an effect of breeding density on departure age without an underlying elevational gradient. Supplementary-fed juveniles and females in high breeding densities departed at younger age than control juveniles and males in low breeding densities. We only found an elevational gradient in the timing of breeding. Late hatched juveniles, and thus individuals at high elevation, departed at earlier age compared to early hatched juveniles. We conclude that favorable natal food conditions, allow for a young departure age of juvenile red kites. We show that the elevational delay in breeding is compensated by premature departure resulting in an elevational gradient in departure age. Thus, elevational differences in dispersal behaviour likely arise due to climatic factors affecting timing of breeding. However, the results also suggest that spatial differences in food availability and breeding density affect dispersal behavior and that their large-scale gradients within the distributional range might result in differential natal dispersal patterns.

Hungry or angry? Experimental evidence for the effects of food availability on two measures of stress in developing wild raptor nestlings.

Food shortage challenges the development of nestlings; yet, to cope with this stressor, nestlings can induce stress responses to adjust metabolism or behaviour. Food shortage also enhances the antagonism between siblings, but it remains unclear whether the stress response induced by food shortage operates via the individual nutritional state or via the social environment experienced. In addition, the understanding of these processes is hindered by the fact that effects of food availability often co-vary with other environmental factors. We used a food supplementation experiment to test the effect of food availability on two complementary stress measures, feather corticosterone (CORTf) and heterophil/lymphocyte ratio (H/L) in developing red kite (Milvus milvus) nestlings, a species with competitive brood hierarchy. By statistically controlling for the effect of food supplementation on the nestlings' body condition, we disentangled the effects of food and ambient temperature on nestlings during development. Experimental food supplementation increased body condition, and both CORTf and H/L were reduced in nestlings of high body condition. Additionally, CORTf decreased with age in non-supplemented nestlings. H/L decreased with age in all nestlings and was lower in supplemented last-hatched nestlings compared with non-supplemented ones. Ambient temperature showed a negative effect on H/L. Our results indicate that food shortage increases the nestlings' stress levels through a reduced food intake affecting both their nutritional state and their social environment. Thus, food availability in conjunction with ambient temperature shapes between- and within-nest differences in stress load, which may have carry-over effects on behaviour and performance in further life-history stages.

Weather and food availability additively affect reproductive output in an expanding raptor population.

The joint effects of interacting environmental factors on key demographic parameters can exacerbate or mitigate the separate factors' effects on population dynamics. Given ongoing changes in climate and land use, assessing interactions between weather and food availability on reproductive performance is crucial to understand and forecast population dynamics. By conducting a feeding experiment in 4 years with different weather conditions, we were able to disentangle the effects of weather, food availability and their interactions on reproductive parameters in an expanding population of the red kite (Milvus milvus), a conservation-relevant raptor known to be supported by anthropogenic feeding. Brood loss occurred mainly during the incubation phase, and was associated with rainfall and low food availability. In contrast, brood loss during the nestling phase occurred mostly due to low temperatures. Survival of last-hatched nestlings and nestling development was enhanced by food supplementation and reduced by adverse weather conditions. However, we found no support for interactive effects of weather and food availability, suggesting that these factors affect reproduction of red kites additively. The results not only suggest that food-weather interactions are prevented by parental life-history trade-offs, but that food availability and weather conditions are crucial separate determinants of reproductive output, and thus population productivity. Overall, our results suggest that the observed increase in spring temperatures and enhanced anthropogenic food resources have contributed to the elevational expansion and the growth of the study population during the last decades.

Political borders impact associations between habitat suitability predictions and resource availability.

CONTEXT: By linking species of conservation concern to their abiotic and biotic requirements, habitat suitability models (HSM) can assist targeted conservation measures. Yet, conservation measures may fail if HSM are unable to predict crucial resources. HSM are typically developed using remotely sensed land-cover classification data but not information on resources per se. OBJECTIVES: While a certain land-cover class may correlate with crucial resources in the area of calibration, political boundaries can abruptly alter these associations. We investigate this potential discrepancy in a well-known study system highly relevant for farmland bird conservation. METHODS: We compared land cover, land-use intensity and resource availability between plots of highest habitat suitability for little owls (Athene noctua) among two neighbouring, but politically separated areas (i.e. south-western Germany vs. northern Switzerland). RESULTS: Land cover and land-use richness did not differ between German and Swiss plots. Yet there were marked differences in terms of land-use intensity and the availability of resources. Land-use intensity was significantly higher and resource availability lower in Swiss compared to German plots. CONCLUSIONS: While accounting well for remotely sensed data such as land cover, HSM may fail to predict land-use intensity and resources across borders. The relationship between geodata used as proxies and ecologically relevant resources may differ according to history, policies and socio-cultural context, constraining the viability of HSM across political borders. This study emphasises the need for fine-scale resource assessments complementing landscape-scale suitability models. Conservation measures need to consider the availability of crucial resources and their socio-economic moderators to be effective.