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.

Sex and size shape the ontogeny of partial migration.

The arrival-time hypothesis of partial seasonal migration proposes that over-winter residence is driven by reproductive benefits of early presence on the breeding grounds. Thus, it predicts increased occurrence of residence at reproductive age. In contrast, the body size hypothesis proposes age-independent benefits of residence for large individuals, who should exhibit greater winter tolerance. Despite different expectations in age patterns for the two hypotheses in long-lived partially migrant species, there is little empirical work investigating the ontogeny of migratory phenotypic expression, that is the expression of residence or migration. We investigated the influence of age, sex and body size on migratory phenotype throughout ontogeny (from first year to early adulthood) in a long-lived partially migrant species, the red kite Milvus milvus. We GPS-tracked 311 individuals tagged as juveniles and 70 individuals tagged as adults over multiple years, yielding 881 observed annual cycles. From this data, we estimated age-dependent probabilities of the transition to residence and of survival in migrants and residents using a Bayesian multistate capture-recapture model, as well as the probability of resuming migration once resident. We then calculated the resulting proportion of residents per age class. In both sexes, almost all juveniles migrated in their first winter, after which the probability of becoming resident gradually increased with age class to approximately 0.3 in adults (>3 calendar years). A size effect in third calendar year females suggests that large females adopt residence earlier in life than small females. The transition from residence back to migration only occurred with a probability of 0.15 across all resident individuals. In addition, survival was notably reduced in adult male migrants compared to adult male residents. These results are largely consistent with the arrival-time and body size hypotheses, simultaneously. Our results reveal a plastic, yet primarily directional within-individual change in migratory phenotype towards more residence with increasing age, varying between sexes and between individuals of different size. This study highlights that different individual characteristics can jointly shape the ontogeny of migratory behaviour and result in complex within-population patterns and persistence of migratory phenotypes.

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.

Temporal and spatial variation in reproductive benefits in a partial migrant.

In partial migrant systems, where residents and migrants coexist within a population, residents are commonly predicted to gain a reproductive advantage over migrants through priority access to high-quality territories and an earlier breeding start. Annual variation in reproductive benefits has been suggested to be important for the coexistence of both strategies in a population, as differences in wintering conditions experienced by the two strategies may result in a periodic reproductive advantage for migrants. However, the importance of spatial environmental variation for reproductive output in partially migrant populations remains largely unexplored. We investigated variation in the reproductive output of migrants and residents in a population of Swiss red kites (Milvus milvus) both temporally, across and within years, and spatially, along an elevational gradient. We gathered 4 years of reproductive data combined with 183 GPS-derived full annual cycles from individuals breeding in the Swiss Alpine foothills. At low, but not high, elevations, residents produced more fledglings than migrants. We also found evidence for annual variation in the reproductive advantage of the two strategies. Furthermore, while reproductive output did decline with a later breeding start, there was no difference in the start of breeding between the two migration strategies. The results of this study suggest that differences in reproductive output between migrants and residents in partial migrant populations can vary both due to the use of spatially distinct overwintering grounds and because the strategies are differently affected by spatial variables in the breeding area, such as elevation. The study emphasizes that spatial and temporal variation in reproductive benefits must be considered when predicting how migratory species will respond to future environmental change.

Imprinted habitat selection varies across dispersal phases in a raptor species.

Natal Habitat Preference Induction (NHPI) plays a significant role in shaping settlement decisions in dispersive animals. Despite its importance, limited research has explored how NHPI varies during natal dispersal phases and across different types of natal habitats. In this study, we examined NHPI in 77 GPS-tagged juvenile red kites (Milvus milvus) originating from different natal habitats along an elevational gradient in Switzerland. We applied individual-based step selection analysis to investigate habitat selection from independence to settlement. We found that during the prospecting phase, individuals predominantly selected habitats similar to their natal environments. However, this pattern changed in the settlement phase: individuals fledged from habitats at higher elevations or closer to urban areas mostly avoided similar habitats (negative NHPI), while those from areas with more farmlands or pastures (combined with forests) showed a preference for similar habitats (positive NHPI). Moreover, the magnitude and individual variation in NHPI differed depending on the natal habitat types from which individuals originated. These findings highlight that strength, direction, and individual variation in NHPI differ between natal habitat types and dispersal phases. Natal habitats therefore can have pervasive legacy effects on subsequent habitat selection, likely affecting population and range dynamics.

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.