Experience does not change the importance of wind support for migratory route selection by a soaring bird.

Migration is a complex behaviour that is costly in terms of time, energy and risk of mortality. Thermal soaring birds rely on airflow, specifically wind support and uplift, to offset their energetic costs of flight. Their migratory routes are a record of movement decisions to negotiate the atmospheric environment and achieve efficiency. We expected that, regardless of age, birds use wind support to select their routes. Because thermal soaring is a complex flight behaviour that young birds need to learn, we expected that, as individuals gain more experience, their movement decisions will also increasingly favour the best thermal uplift conditions. We quantified how route choice during autumn migration of young European honey buzzards (Pernis apivorus) was adjusted to wind support and uplift over up to 4 years of migration and compared this with the choices of adult birds. We found that wind support was important in all migrations. However, we did not find an increase in the use of thermal uplifts. This could be due to the species-specific learning period and/or an artefact of the spatio-temporal scale of our uplift proxies.

Paternal transmission of migration knowledge in a long-distance bird migrant.

While advances in biologging have revealed many spectacular animal migrations, it remains poorly understood how young animals learn to migrate. Even in social species, it is unclear how migratory skills are transmitted from one generation to another and what implications this may have. Here we show that in Caspian terns Hydroprogne caspia family groups, genetic and foster male parents carry the main responsibility for migrating with young. During migration, young birds stayed close to an adult at all times, with the bond dissipating on the wintering grounds. Solo-migrating adults migrated faster than did adults accompanying young. Four young that lost contact with their parent at an early stage of migration all died. During their first solo migration, subadult terns remained faithful to routes they took with their parents as young. Our results provide evidence for cultural inheritance of migration knowledge in a long-distance bird migrant and show that sex-biased (allo)parental care en route shapes migration through social learning.

First evidence of neonicotinoid residues in a long-distance migratory raptor, the European honey buzzard (Pernis apivorus).

The evidence of negative impacts of agricultural pesticides on non-target organisms is constantly growing. One of the most widely used group of pesticides are neonicotinoids, used in treatments of various plants, e.g. oilseed crops, corn and apples, to prevent crop damage by agricultural insect pests. Treatment effects have been found to spill over to non-target insects, such as bees, and more recently also to other animal groups, among them passerine birds. Very little is known, however, on the presence of neonicotinoids in other wild species at higher trophic levels. We present results on the presence of neonicotinoid residues in blood samples of a long-distant migratory food-specialist raptor, the European honey buzzard. Further, we investigate the spatial relationship between neonicotinoid residue prevalence in honey buzzards with that of crop fields where neonicotinoids are typically used. A majority of all blood samples contained neonicotinoids, thiacloprid accounting for most of the prevalence. While neonicotinoid residues were detected in both adults and nestlings, the methodological limit of quantification was exceeded only in nestlings. Neonicotinoids were present in all sampled nests. Neonicotinoid presence in honey buzzard nestlings' blood matched spatially with the presence of oilseed plant fields. These are the first observations of neonicotinoids in a diurnal raptor. For better understanding the potential negative sub-lethal of neonicotinoids in wild vertebrates, new (experimental) studies are needed.

Impact of climate change and prey abundance on nesting success of a top predator, the goshawk.

Contemporary research has documented a large number of shifts in spring phenology and changes in distribution range although the average spring temperatures have increased by only 0.3-0.6 °C over the past 100 years. Generally, earlier breeding birds have larger clutch sizes, and the advancing spring could thus potentially increase breeding success. Shifts in spring phenology can, however, be crucial for bird reproduction, and mistiming the breeding event may even have negative consequences for population development. Our aim was to explore how weather and prey abundance relates to the breeding performance of a north European top predator, the northern goshawk Accipiter gentilis. Our nationwide dataset from Finland, spanning the period 1989-2004, shows that ambient weather has a greater impact on the timing and success of breeding than the density of grouse Tetraonidae, the main prey of goshawks. Higher early spring temperatures were associated with advancing hatching date of goshawks. Correspondingly, grouse density and temperature during laying and brooding were positively associated with brood size, while precipitation showed a negative connection. Applying our models to a future scenario of climate warming, combined with a 50 % reduction in grouse density, suggests that average breeding dates will advance only 2.5 days and average breeding success would remain the same. Notably, breeding success was not spatially equal throughout Finland, as northern and eastern populations suffered most from declining grouse densities. The observed pattern is thus the opposite to what is expected from a population situated at the northern edge of its distribution range, and thus may help to understand why populations may not increase at the northern edge of their thermal distribution due to climate change.

Coupling in goshawk and grouse population dynamics in Finland.

Different prey species can vary in their significance to a particular predator. In the simplest case, the total available density or biomass of a guild of several prey species might be most relevant to the predator, but behavioural and ecological traits of different prey species can alter the picture. We studied the population dynamics of a predator-prey setting in Finland by fitting first-order log-linear vector autoregressive models to long-term count data from active breeding sites of the northern goshawk (Accipiter gentilis; 1986-2009), and to three of its main prey species (1983-2010): hazel grouse (Bonasa bonasia), black grouse (Tetrao tetrix) and capercaillie (T. urogallus), which belong to the same forest grouse guild and show synchronous fluctuations. Our focus was on modelling the relative significance of prey species and estimating the tightness of predator-prey coupling in order to explain the observed population dynamics, simultaneously accounting for effects of density dependence, winter severity and spatial correlation. We established nine competing candidate models, where different combinations of grouse species affect goshawk dynamics with lags of 1-3 years. Effects of goshawk on grouse were investigated using one model for each grouse species. The most parsimonious model for goshawk indicated separate density effects of hazel grouse and black grouse, and different effects with lags of 1 and 3 years. Capercaillie showed no effects on goshawk populations, while the effect of goshawk on grouse was clearly negative only in capercaillie. Winter severity had significant adverse effects on goshawk and hazel grouse populations. In combination, large-scale goshawk-grouse population dynamics are coupled, but there are no clear mutual effects for any of the individual guild members. In a broader context, our study suggests that pooling data on closely related, synchronously fluctuating prey species can result in the loss of relevant information, rather than increased model parsimony.

Competitive exclusion within the predator community influences the distribution of a threatened prey species.

While much effort has been made to quantify how landscape composition influences the distribution of species, the possibility that geographical differences in species interactions might affect species distributions has received less attention. Investigating a predator-prey setting in a boreal forest ecosystem, we empirically show that large-scale differences in the predator community structure and small-scale competitive exclusion among predators affect the local distribution of a threatened forest specialist more than does landscape composition. Consequently, even though the landscape parameters affecting Siberian flying squirrel (Pteromys volans) distribution (prey) did not differ between nest sites of the predators Northern Goshawks (Accipiter gentilis) and Ural Owls (Strix uralensis), flying squirrels were heterospecifically attracted by goshawks in a region where both predator species were present. No such effect was found in another region where Ural Owls were absent. These results provide evidence that differences in species interactions over large spatial scales may be a major force influencing the distribution and abundance patterns of species. On the basis of these findings, we suspect that subtle species interactions might be a central reason why landscape models constructed to predict species distributions often fail when applied to wider geographical scales.

The impact of climate and cyclic food abundance on the timing of breeding and brood size in four boreal owl species.

The ongoing climate change has improved our understanding of how climate affects the reproduction of animals. However, the interaction between food availability and climate on breeding has rarely been examined. While it has been shown that breeding of boreal birds of prey is first and foremost determined by prey abundance, little information exists on how climatic conditions influence this relationship. We studied the joint effects of main prey abundance and ambient weather on timing of breeding and reproductive success of two smaller (pygmy owl Glaucidium passerinum and Tengmalm's owl Aegolius funereus) and two larger (tawny owl Strix aluco and Ural owl Strix uralensis) avian predator species using long-term nation-wide datasets during 1973-2004. We found no temporal trend either in vole abundance or in hatching date and brood size of any studied owl species. In the larger species, increasing late winter or early spring temperature advanced breeding at least as much as did high autumn abundance of prey (voles). Furthermore, increasing snow depth delayed breeding of the largest species (Ural owl), presumably by reducing the availability of voles. Brood size was strongly determined by spring vole abundance in all four owl species. These results show that climate directly affects the breeding performance of vole-eating boreal avian predators much more than previously thought. According to earlier studies, small-sized species should advance their breeding more than larger species in response to increasing temperature. However, we found an opposite pattern, with larger species being more sensitive to temperature. We argue that this pattern is caused by a difference in the breeding tactics of larger mostly capital breeding and smaller mostly income breeding owl species.

Food regulates reproduction differently in different habitats: experimental evidence in the Goshawk.

Food supplementation experiments have been widely used to get detailed insight into how food supply contributes to the reproductive performance of wild animals. Surprisingly, even though food seldom is distributed evenly in space, variation in local habitat quality has usually not been controlled for in food supplementation studies. With results from a two-year feeding experiment involving a habitat-sensitive avian top predator, the Northern Goshawk Accipiter gentilis, we show that treatment effects on goshawk reproductive performance are habitat dependent. Extra food reduced nestling mortality in low-quality territories where prime habitat (forest) is scarce, but not in high-quality territories where prime habitat is abundant. Consequently, brood size did not differ between treatment categories in heavily forested territories, but fledgling numbers differed between unfed and fed goshawk pairs breeding in territories where forest is scarce. However, because extra food was not superabundant, this artificial increase in offspring number induced a dramatic decrease in nestling condition in low-quality territories. Treatment effects were detected even after controlling statistically for other potentially confounding effects (year, territory identity) and strongly covaried with territory-specific abundances of the most important summer prey species. These results highlight the importance of acknowledging the effect that small-scale variation in habitat quality and availability of natural food may have on the results of food supplementation experiments. In order to assess the generality of food supplementation effects, the integration of habitat heterogeneity and variation in food abundance is thus needed, especially among species in which small-scale variation in habitat quality influences demographic patterns.

Interactions between habitat heterogeneity and food affect reproductive output in a top predator.

1. Habitat heterogeneity has important repercussions for species abundance, demography and life-history patterns. While habitat effects have been more thoroughly studied in top-down situations (e.g. in association with predation), their role in bottom-up situations (e.g. in association with food abundance) has been less explored and the underlying mechanism(s) behind the ecological patterns have not commonly been identified. 2. With material from 1993 to 2003, we test the hypothesis that the reproduction of Finnish northern goshawks Accipiter gentilis (L.) is bottom-up limited by habitat composition, especially in situations where the density of their main prey (grouse) is low. Special emphasis was placed on identifying the mechanism(s) behind potential habitat effects. 3. While laying date and large-scale variation in the main prey density (but not habitat composition) were related to the number of eggs goshawks laid, small-scale differences in alternative prey density between different territories later influenced how many young were fledged via the mechanism of habitat-dependent partial-brood loss. As a result of this mechanism, a difference in nestling condition also arose between goshawk territories with differing habitat compositions. 4. As the relative proportions of different landscape elements in a given landscape is a function of large-scale differences in geomorphology and land use, this means that the reproductive performance of goshawks as averaged over larger scales can be understood correctly only in respect to the fact that habitat gradients differ across landscapes. 5. In addition to being one of the first papers identifying the mechanism of partial brood loss as being primarily responsible for the habitat-specific differences in the production of young, this study further illustrates the need to identify small-scale mechanisms to correctly understand the large-scale patterns of reproductive performance in territorial species. The repercussions of the observed habitat effect for local population development are discussed.