AimOne of the few general laws in ecology is that species richness is a positive function of area. However, it has been proposed that area would merely be a proxy for energy. Additionally, habitat heterogeneity has been found to be an important factor determining species richness. Yet the relative importance of those relationships is little known, and it is still unclear how they are brought about. We aimed to dissect which factors drive the species richness of boreal forest birds, and to identify the most probable mechanisms. Location Forested protected areas in Finland.Methods Using bird line census data collected in 104 protected areas, we ran simultaneous autoregressive models to explain the species richness of forest birds. We explored the value of forest area, tree volume, tree growth, mean degree days and habitat heterogeneity as explanatory variables and used the species richness within different species groups, based on the predictions of hypothesized mechanisms, as a response variable.Results Energy, rather than area or habitat heterogeneity, seems to be the main driver of species richness in boreal forest birds. More specifically, productive energy was a better predictor of total species richness than solar energy. Among the tested hypothetical mechanisms, the sampling hypothesis received strong support. After accounting for sampling, solar energy had an effect on species richness.Main conclusions As productive energy, such as tree volume, is associated with species richness, high-energy areas should be prioritized in forest conservation planning. Reductions in productive energy may first lead to the disappearance of the rarest species due to the random sampling process. Climate change may result in increased species richness due to increasing amount of productive and solar energy in forests. However, the range shifts of bird species may not be fast enough to keep up with the temperature increases.

Climate change is driving species to shift their distributions toward high altitudes and latitudes, while habitat loss and fragmentation may hamper species ability to follow their climatic envelope. These two drivers of change may act in synergy, with particularly disastrous impacts on biodiversity. Protected areas, PAs, may thus represent crucial buffers against the compounded effects of climate change and habitat loss. However, large‐scale studies assessing the performance of PAs as such buffers remain scarce and are largely based on species occurrence data. Conversely, abundance data have proven to be more reliable for addressing changes in wildlife populations under climate change. We evaluated changes in bird abundance from the 1970s–80s to the 2000s inside and outside PAs at the trailing range edge of 30 northern bird species and at the leading range edge of 70 southern species. Abundances of retracting northern species were higher and declined less inside PAs at their trailing range edge. The positive effect of PAs on bird abundances was particularly marked in northern species that rely strongly on PAs, that is, their density distribution is largely confined within PAs. These species were nearly absent outside PAs in the 2000s. The abundances of southern species were in general lower inside PAs and increased less from the 70s–80s to 2000s. Nonetheless, species with high reliance on PAs had much higher abundances inside than outside PAs in the 2000s. These results show that PAs are essential in mitigating the retraction of northern species, but also facilitate northward expansions of southern species highly reliant on PAs. Our study provides empirical evidence documenting the role of PAs in facilitating species to adjust to rapidly changing climatic conditions, thereby contributing to the mitigation of impending biodiversity loss. PAs may thus allow time for initiating wider conservation programs on currently unprotected land.

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Human land-use effects on species populations are minimized in protected areas and population changes can thus be more directly linked with changes in climate. In this study, bird population changes in 96 protected areas in Finland were compared using quantitative bird census data, between two time slices, 1981–1999 and 2000–2009, with the mean time span being 14 years. Bird species were categorized by distribution pattern and migratory strategy. Our results showed that northern bird species had declined by 21 per cent and southern species increased by 29 per cent in boreal protected areas during the study period, alongside a clear rise (0.7–0.8°C) in mean temperatures. Distribution pattern was the main factor, with migratory strategy interacting in explaining population changes in boreal birds. Migration strategy interacted with distribution pattern so that, among northern birds, densities of both migratory and resident species declined, whereas among southern birds they both increased. The observed decline of northern species and increase in southern species are in line with the predictions of range shifts of these species groups under a warming climate, and suggest that the population dynamics of birds are already changing in natural boreal habitats in association with changing climate.

Anthropogenic climate change is rapidly becoming one of the main threats to biodiversity, along with other threats triggered by human-driven land-use change. Species are already responding to climate change by shifting their distributions polewards. This shift may create a spatial mismatch between dynamic species distributions and static protected areas (PAs). As protected areas represent one of the main pillars for preserving biodiversity today and in the future, it is important to assess their contribution in sheltering the biodiversity communities, they were designated to protect. A recent development to investigate climate-driven impacts on biological communities is represented by the community temperature index (CTI). CTI provides a measure of the relative temperature average of a community in a specific assemblage. CTI value will be higher for assemblages dominated by warm species compared with those dominated by cold-dwelling species. We here model changes in the CTI of Finnish bird assemblages, as well as changes in species densities, within and outside of PAs during the past four decades in a large boreal landscape under rapid change. We show that CTI has markedly increased over time across Finland, with this change being similar within and outside PAs and five to seven times slower than the temperature increase. Moreover, CTI has been constantly lower within than outside of PAs, and PAs still support communities, which show colder thermal index than those outside of PAs in the 1970s and 1980s. This result can be explained by the higher relative density of northern species within PAs than outside. Overall, our results provide some, albeit inconclusive, evidence that PAs may play a role in supporting the community of northern species. Results also suggest that communities are, however, shifting rapidly, both inside and outside of PAs, highlighting the need for adjusting conservation measures before it is too late.