Annual variation in breeding success in boreal forest grouse: Four decades of monitoring reveals bottom-up drivers to be more important than predation.

Knowledge of the temporal variation in reproductive success and its key driving factors is crucial in predicting animal population persistence. Few studies have examined the effects of a range of explanatory factors operating simultaneously on the same population over a long period. Based on 41 years of monitoring (1979-2019), we tested prevailing hypotheses about drivers of annual variation in breeding success in two sympatric species of boreal forest grouse-the capercaillie (Tetrao urogallus) and the black grouse (T. tetrix)-in a 45 km2 boreal forest landscape. From counts in early August, we measured breeding success (chicks/hen) along with potential determining factors. We formulated five main hypotheses on causes of variation (hen condition, chick weather, chick food, predation, demographic characteristics) and derived 13 associated explanatory variables for analysis. We first tested the five hypotheses separately and then used model selection (AICc) to rank the best predictive models irrespective of hypotheses. Lastly, we used path analysis to illuminate potential causal relationships. Barring demographic characteristics, all hypotheses were supported, most strongly for chick food and predation. Among predictor variables, chick food (insect larvae and bilberry fruit crops), vole and fox abundances, the winter-NAO index, and temperature after hatching, had the strongest effect sizes in both species. Precipitation after hatching had no detectable effect. Model selection indicated bottom-up factors to be more important than predation, but confounding complicated interpretation. Path analysis suggested that the high explanatory power of bilberry fruiting was due not only to its direct positive effect on chick food quality but also to an indirect positive effect on vole abundance, which buffers predation. The two components of breeding success-proportion of hens with broods and number of chicks per brood-were uncorrelated, the former having the strongest effect. The two components had different ecological correlates that often varied asynchronously, resulting in overall breeding success fluctuating around low to moderate levels. Our study highlights the complexity of key explanatory drivers and the importance of considering multiple hypotheses of breeding success. Although chick food appeared to equal or surpass predation in explaining the annual variation in breeding success, predation may still be the overall limiting factor. Comparative and experimental studies of confounded variables (bilberry fruiting, voles, and larvae) are needed to disentangle causes of variation in breeding success of boreal forest grouse.

Climate change and bird reproduction: warmer springs benefit breeding success in boreal forest grouse.

Global warming is predicted to adversely affect the reproduction of birds, especially in northern latitudes. A recent study in Finland inferred that declining populations of black grouse, Tetrao tetrix, could be attributed to advancement of the time of mating and chicks hatching too early-supporting the mismatch hypothesis. Here, we examine the breeding success of sympatric capercaillie, T. urogallus, and black grouse over a 38-year period in southeast Norway. Breeding season temperatures increased, being most pronounced in April. Although the onset of spring advanced nearly three weeks, the peak of mating advanced only 4-5 days. In contrast to the result of the Finnish study, breeding success increased markedly in both species (capercaillie: 62%, black grouse: 38%). Both brood frequency and brood size increased during the study period, but significantly so only for brood frequency in capercaillie. Whereas the frequency of capercaillie broods was positively affected by rising temperatures, especially during the pre-hatching period, this was not the case in black grouse. Brood size, on the other hand, increased with increasing post-hatching temperatures in both species. Contrary to the prediction that global warming will adversely affect reproduction in boreal forest grouse, our study shows that breeding success was enhanced in warmer springs.

Habitat selection as a hierarchical spatial process: the green woodpecker at the northern edge of its distribution range.

Habitat selection can be envisaged as a hierarchical spatial process, from choice of home range to choice of dietary item. The green woodpecker (Picus viridis) is described as being closely bound to cultivated land and deciduous forests, mainly due to its summer diet composed of ants (Hymenoptera: Formicidae) found on meadows and pastures. To explore possible responses of this woodpecker to recent changes in land use practice, we studied home ranges, feeding habitats and food selection of a marginal population (four radio-marked males and five females) in a 30,000-ha conifer-dominated landscape at the northern edge of its distribution range in south-central Scandinavia. We asked: (1) Is the green woodpecker confined to areas with cultivated land and deciduous forest? (2) If so, are important food items (ants) particularly abundant or exclusively found there? (3) Can clearcuts and young plantations substitute for cultivated land as feeding habitat? Home ranges (mean=100 ha) were invariably confined to the parts of the landscape that contained cultivated land (<1% of the total area). In summer, birds preferred to feed in cultivated land, presumably due to a higher overall biomass of ants compared to forest habitats. They avoided clearcuts, but preyed extensively upon soil-dwelling ants in young conifer stands (16-30 years old). We failed to find preferences for particular ant groups (Lasius niger and L. flavus) associated with cultivated land. The principal summer food was Serviformica, an ant group that was equally abundant in cultivated land and forest habitat. A positive correlation between ant body mass and a preference index suggests that the birds selected the larger ant species independent of habitat type. In winter, birds fed exclusively on mound-building Formica rufa-ants in closed-canopy, older forest stands. Our results indicate that the green woodpecker successfully utilizes young conifer plantations as feeding habitat. At a larger scale, we hypothesize that green woodpecker populations fail to establish in managed forest tracts, not because of food shortage, but because the landscapes lack cultivated land serving as a key stimulus encouraging individuals to settle.