Mountain regions are globally important areas for biodiversity but are subject to multiple human-induced threats, including climate change, which has been more severe at higher elevations. We reviewed evidence for impacts of climate change on Holarctic mountain bird populations in terms of physiology, phenology, trophic interactions, demography and observed and projected distribution shifts, including effects of other factors that interact with climate change. We developed an objective classification of high-elevation, mountain specialist and generalist species, based on the proportion of their breeding range occurring in mountain regions. Our review found evidence of responses of mountain bird populations to climate (extreme weather events, temperature, rainfall and snow) and environmental (i.e. land use) change, but we know little about either the underlying mechanisms or the synergistic effects of climate and land use. Long-term studies assessing reproductive success or survival of mountain birds in relation to climate change were rare. Few studies have considered shifts in elevational distribution over time and a meta-analysis did not find a consistent direction in elevation change. A meta-analysis carried out on future projections of distribution shifts suggested that birds whose breeding distributions are largely restricted to mountains are likely to be more negatively impacted than other species. Adaptation responses to climate change rely mostly on managing and extending current protected areas for both
Timing reproduction to coincide with optimal environmental conditions is key for many organisms living in seasonal habitats. Advance in the onset of spring is a particular challenge to migratory birds that must time their arrival without knowing the conditions on the breeding grounds. This is amplified at high elevations where resource availability, which is linked to snowmelt and vegetation development, shows much annual variation. With the aim of exploring the effects of variability in the onset of local resource availability on reproduction, we compared key life history events in an Alpine population of the Northern Wheatear (Oenanthe oenanthe) between years of contrasting timing of snowmelt. Based on remote sensed images, we identified 2020 as an exceptionally early snowmelt and green-up year compared to the preceding year and the long-term average. Individuals tracked with light-level geolocators arrived well before the snowmelt in 2020 and clutch initiation dates across the population were earlier in 2020 compared to 2019. However, observations from a citizen science database and nest monitoring data showed that the arrival-breeding interval was shorter in 2020, thus the advance in timing lagged behind the environmental conditions. While hatching success was similar in both years, fledging success was significantly reduced in 2020. A trophic mismatch in early 2020 could be a possible explanation for the reduced reproductive success, but alternative explanations cannot be excluded. Our results show that, despite the timely arrival at the breeding grounds and a contraction of the arrival-breeding interval, Wheatears were not able to advance breeding activities in synchrony with environmental conditions in 2020. Earlier reproductive seasons are expected to become more frequent in the future. We show that the negative effects of changing seasons in Alpine migratory birds might be similar to birds breeding at high latitudes, despite their shorter migratory distance.
Predictions derived from species distribution models (SDMs) are strongly influenced by the spatial scale at which species and environmental data (e.g. climate) are gathered. SDMs of mountain birds usually build on large-scale temperature estimates. However, the topographic complexity of mountain areas could create microclimatic refuges which may alter species distributions at small spatial scales. To assess whether fine-scale data (temperature and/or topography) improve model performance when predicting species occurrence, we collected data on presence-absence of bird species, habitat and fine-scale temperature at survey points along an elevational gradient in the Alps (NW Italy). Large-scale temperature data, and both large-and fine-scale topography data, were extracted from online databases for each point. We compared species models (fine-scale vs large-scale) using an information-theoretic approach. Models including fine-scale temperature estimates performed better than corresponding large-scale models for all open habitat species, whereas most forest/ecotone species showed no difference between the two scales. Grassland birds such as Northern Wheatear Oenanthe oenanthe and Water Pipit Anthus spinoletta were positively associated with warmer microclimates. These results suggest that alpine grassland species are potentially more resistant to the impact of climate change than previously predicted, but that indirect effects of climate change such as habitat shifts (forest-and shrub encroachment at high elevations) pose a major threat. Therefore, active management of alpine grassland is needed to maintain open areas and to prevent potential habitat loss and fragmentation. SDMs based solely on large-scale temperatures for open habitat species in the Alps should be reassessed .
Climate change is leading to the advancement of spring conditions, resulting in an earlier snowmelt and green-up, with highest rates of change in highly seasonal environments, including alpine habitats. Migratory birds breeding at high elevations need to time their arrival and lay dates accurately with this advancement, but also with the annually variable spring conditions at their breeding sites, to maximize nest survival probabilities and reproductive output. Nest survival probability and mean nestling mass were analysed in relation to lay date and habitat conditions in an alpine population of the migratory Northern Wheatear Oenanthe oenanthe collected over six consecutive breeding seasons in the Western Italian Alps. This open grassland species showed the lowest nest survival probability in years with an early onset of spring conditions. Within-season, nest survival was highest when breeding late, at lower elevations, and when grass cover and grass height were higher. Both across-and within-season, severe weather conditions may indirectly lead to higher early season nest failure rates by increasing predation risk. By contrast, mean nestling mass, and thus the quality of the fledglings, was lower when breeding late. This might be driven by a mismatch with the peak in food abundance. Breeding early is thus generally advantageous in terms of chick quality in our highelevation population, but reproductive success is limited by the risk of nest failure that is higher in early springs and early in the season. This trade-off between breeding early and late may thus allow Northern Wheatears to maximize fitness under highly variable spring conditions. However, climate change may cause disruption to this trade-off, and shifts in phenology could become a threat for migratory alpine birds that might not be able to keep track of advancing spring conditions.
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